THE (LEPTDOPTERA) OF CANADA

CHRISTOPHER WILKINSON Department of Systematics and Zoogeography Vrije Universiteit Amsterdam, Netherlands

M. J. SCOBLE Transvaal Museum Pretoria 0001, R.S.A.

MEMOIRS OF THE ENTOMOLOGICAL SOCIETY OF CANADA - NO. 107

Pubtished bylHE ENTOMOLOGICAL SOCIETY OF CANADA, OTTAWA K1Z 7K9 1979 To My Friends and Colleagues of the Biosystematics Research Institute, Ottawa, Canada (C.W.) CONTENTS Pacs

Abstract I

Introduction I

Methods L

List of Abbreviations .

Acknowledgments . .

Taxonomic History

Family NEPTICULIDAE ... 8

Gerus t2 't3 GenusEctoedemia . ..

GenusODrzssa .... 98

Gents Glaucolepis r02

General Discussion and Summary 108

References 115

Index to Genera, Species, and Subspecies . .. . r29

Plates l.Therosaefoliellagtotp-mlnes... 'p'119 2. The crataeg{olielta gtovp-mines . ' p' 120 3. The saginella group-mrnes " " " ' p' l2l 4. The corylifoliella group-mines . . . ' p' 122 5. Thebifasciella group andquercipulchella group-'mines . . . " " p' 123 6. argyropeza, E. clemensella, E. platanella-mines . ' p' 124 125 7 . Ectoedemia rubifulietla, E. ulmella, E. quadrinotata, Glaucolepis saccharella-mines - . . . ' . p' 8. The rosaefoliella group a d the cratae7ifotiella gtoup-imagines . ' ' ' ' ' ' p ' 126 9. The saginella group, the corytifolielta group, and thebifuscielta group-imagines " " " " p' 127 lO. The procrastinella group, Ectoedemia canutus , E. Iindquisti, E. rubifulielln, E. quadrinotata Obrissaochrefasciilla,'O. sericopeza,Glaucolepis saccharella-imagines . " " p' 128 THE NEPTICULIDAE () OF CANADA Cur.tstopur:,n WtLxtNsoN aNp M. J. ScosI-E, Abstract A taxonomic revision, including descriptions, diagnoses, and bibliographies, is given of the genera and species of the family Nepticulidae (Lepidoptera) so far collected in Canada. In addition, extensive material from the U.S.A. and a number of specimens from Europe have been examined. Five new species and two new subspecies are described. Notes on the biology of the species have been collated from rearing records, label data, and the relevant literature. With one exception all existing types have been examined and, in addition, a number of types designated. A major f'eature of this work is a study of the genitalia of the never befbre undertaken extensively on the North American Nepticulidae. These structures give valuable diagnostic features and are of fundamental importance in the provision of modern descriptions and keys. As genitalia characters are thought to be the best taxonomic indicators of relationships between species in the Nepticulidae, the species-groups proposed have been mainly defined on these structures. The confusion over the nomenclature of the family name is discussed. Taxonomic histories of the genera are given and one generic synonymy is made. Taxonomic problems are discussed in relation to distribution and food-plant choice. The hoslplant associations of the family are reviewed, with reference to the mining habits of the Iarvae. A priori use of host-plant data in the of the Nepticulidae is found to be limited. However, the proposed species-groups illustrate a general conclusion that related species tend to feed on plants that are themselves taxonomically related.

INTRODUCTION The Nepticulidae is a cosmopolitan family of microlepidopteran whose larvae mine the parenchymatous tissue of plants or occasionally induce the formation of galls on the petioles of leaves. Most species are leaf-miners, but some mine bark and others fiuits. Nearly all the work on the systematics of the genera has been carried out in the Nearctic and the Palaearctic. However, because the entomologists of these respective regions worked rather independently of each other, there have been certain inconsisten- cies in the generic placement of a number of species. The Canadian species are regarded in this work as falling into four genera: Stigmella Schrank, Ectoedemia Busck, Obrussct Braun, and Glaucolepis Braun, principally on the basis of wing venation, genitalia, distribution, and also biology where known. Some changes in nomenclature are the result of a critical review of the taxonomic history. These are primarily the subject of a separate paper (1978) but the main conclusions are summarized here. The aims of this study are'.To list, describe, andprovide modern fteys of the genera and species of the Nepticulidae so far known from Canada, and to include any new species and subspecies. The work aims to provide descriptions of the external features using high intensity illumination to give both the ground colour and the reflections of all the species considered. The genitalia are used extensively to provide taxonomic characters for the descriptions. Leaf-mine pattern and, to a certain extent, choice of food-plant give additional diagnostic features. Although this study deals primarily with the Canadian members of the family, it is aimed to incorporate specimens collected in the U.S.A. of species which occur in both countries. This additional material comes largely from the Annette Braun collection (deposited in the Academy of Natural Sciences, Philadelphia) which includes most of the types of the North American Nepticulidae. To resolve certain problems of the taxonomic treatment of the group. Clemens and Chambers commenced the work on the svstematics of the Nearctic Nepticulidae last l.n,u.r. In severar and, on a number or occasions, described'"JJ. species from";#ffi';"';;"ffiiar the mines alone. The latter are often of little value in the differentiation of closely related species. Braun reared imagines of most of Clemens' species and also a large number of new ones. However she did not use genital characters, but relied largely on external colouration and form of the mines. In this study the genitalia (believed to be less affected by environmental vicissitudes) are used extensively in indicating relationships as well as providing good diagnostic characters. To relate the taxonomy of the family in the Nearctic to work carried out in the Palaearctic. The cosmopolitan nature of the family is clearly reflected by the distribution of the genera. Stigmella, Ectoedemia, and Obrussa are all represented in Europe as well as North America. There is considerably more literature on the European Nepticulidae, including studies of the genitalia, than on the North American representatives. However, although European workers have made a number of references to the Nearctic species, they have studied only a few of them at first hand. Consequently a further purpose of this study is to make changes, where necessary, in nomenclature and to suggest treatment for the much disputed supra-specific ranks. This includes examining a number of closely related European species and, in two cases, species occurring in both this region and North America. To collate from label data, rearing records and the literature, notes on the biology of each species and to discuss host-plant relationships.

METHODS The wing measurements give the alar expanse in millimeters. The moths were measured from the middle of the thorax to the wing-tip and this figure was doubled to give the full expanse. The very small size of these can cause problems in setting and any buckling in the wings makes accurate measurements impossible. As some specimens were not well mounted, we have only measured those which give a relatively accurate figure. The genitalia were figured by means of a drawing tube fitted to a Wild M20 microscope. The scale represents 1/10 mm unless otherwise indicated. The male genitalia are presented from a ventral aspect; the aedeagus is represented separately from the capsule. As the valves are curved, in many cases the back and front are represented by thick lines on opposite sides. Occasionally a lateral view of a valve is given. Black and white photographs of imagines and mines are provided. The extremely small size, and lighting difficulties, have proved problematic in producing satisfactory prints of the imagines. However, to a certain extent, these photographs illustrate the variation and convergence within the species examined. With certain exceptions, each species is described, and provided with a 'DIAGNOSIS': this is differential. The exceptions include taxa which are not sufficiently different from others to warrant repeating a full description. Instead 'DIAGNOSTIC FEATURES' are given. Single specimens showing variation not falling within the confines of a given species description are separated as specimens a, b, c, etc. The dates given in the 'SPECIMENS EXAMINED' are those of the emergence of the imagine, not the date on which the tenanted, mined leaves were collected. A section entitled 'BIOLOGY' follows the more formal taxonomy, in which additional information, collated from mined leaves, rearing records, and relevant literature is given. A number of species-groups are proposed in this study. We are agreed with Johansson (191 l) that to split the Nepticulidae into a large number of genera would be WILKINSON AND SCOBLE: THE NEPTICULIDAE OF CANADA J of little value. The more informal method adopted indicates relationships between species within a given group yet prevents the taxonomy from becoming unwieldy. Throughout this study the International Code of Zoological Nomenclature has been followed.

Tvpp MerEntaL. Besides selecting lectotypes and paralectotypes from syntypes, four neotypes have been recommended. In all cases the species were first described by Clemens in the last century from leaves mined by the larvae. The International Code accepts names founded on the 'work of an animal' as long as they were designated before 193 1 (Article 24b). Enquiries have shown that the mines from which these descriptions were made are not extant.

TnxoNolurc Cul.nectsns. In nearly all species the forewings of the imagines have reflections, the intensity of which varies with the angle of incident light. Sometimes they are bright and at other times weak and hardly visible. To standardize the descriptions, the ground colour of the forewings is given followed by the reflections that can be seen when the specimen is rotated under the light. High intensity illumination, together with a blue, daylight filter, was used for viewing. Names used in describing the genitalia are shown in the annotated figures 6 and 7 the male and female of rosaefoliella the first species described. Any character not present- in that species, which occurs -in a later one, is subsequently labelled in the respective figure (see Figs. 1 1, 12, 41, 42, and 51). The terms used in this study generally follow Beirne (1945) whose work on the males of the British Nepticulidae is frequently cited by other workers on the group. The lateral arms of the vinculum form either a "U", the ends of which articulate with the sides of the tegumen, or alternatively a complete ring. In species with a ring-shaped vinculum it seems that the tegumen has been extended into a projection, termed by Beirne a pseuduncus (see Fig. al). In such cases the true uncus is membranous and very difficult to discern. Beirne states that in Etainia (later in this work to be synonymized), a true uncus, rather than a pseuduncus is present. Having examined the genitalia from a lateral aspect, we feel that there is no separation between the 'uncus' and the tegumen and that the term pseuduncus again applies. The tegumen appears to curl over forming a lip; it is probably this feature that Beirne figured in his lateral, annotated view of the capsule. The valves are joined basally by a transtilla (Fig. 6). The lateral arm of this fuses with the transverse bar to form a rigid structure. Attempts to 'spread' the valves results in distortion. Other terms describing the male genitalia, and those used in the descriptions of the female, have come from Klots in Tuxen (1956), Eyer (1924), Busck and Heinrich (1921), and Snodgrass (1935, 1957). These authors together with McDunnough (1911) have provided descriptions of characters, and a history of the nomenclature. Regarding the female genitalia, there are two pairs of apophyses, one anterior and the other posterior. The posterior pair, arising from the 9th segment, is usually centrally situated and narrow. The anterior pair, arising from the Sth segment, is generally arcuate, broad basally, and lies outside the posterior pair. The ductus bursae is used in the wide (taxonomic) sense, see Klots, which includes the ductus seminalis of the morphologist. The membranous sac arising from the ductus is simply termed an accessory sac in this study. For labels see Fig. 7. All the Canadian genera may be separated on their wing venation (Figs. l-5). For comparison, a figure of Zeller is also included (after Klimesch, 1953) who reviewed the European members of the . MEMOIRS OF THE ENTOMOLOGICAL SOCIETY OF CANADA The larval and mine terminology adopted is that of Hering (195 l) which is explained by means of the tables below.

GTII Plant organ mined Classification of mg1: Stigmella Leaf Phyllonome Glaucolepis Leaf Phyllonome Ectoedemia Bark (cortical Caulonome parenchyma of twigs) Petiole Phyllonome Leaf blade Phyllonome Obrussa Fruit Carponome

Hering's Ming nomenclature Description Example Linear Ophionome Simple tract coryliJoLiella Blotch Stigmatonome eats around stigmaciella in all directions Linear/Blotch Ophistigmatonome clemensella

LIST OF ABBREVIATIONS BM(NH) British Museum (Natural History), London CNC Canadian- National Collection. Ottawa International- Code International Code of Zoological Nomenclature adopted by the XVth International Congress- of Zoology MCZ Museum of Comparative Zoology, Cambridge, Massachusetts, U.S.A. Phila- Academy of Natural Sciences, Philadelphia, U.S.A. Soo - Canada Department of Forestry. Sault Ste. Marie UBC - University of British Columbia, Canada USNM - United States National Museum, Smithsonian Institution, Washington D.C., U.S.A. ACKNOWLEDGMENTS Christopher Wilkinson. This study was started by C.W. during the tenure of a National Research Council of Canada, Post-doctoral Fellowship at the Biosystematics Research Institute, Ottawa rn 1910-i1. He would like to acknowledge the help at that time of Dr. David F. Hardwick, now Director of the Institute, the late Dr. Tom N. Freeman, Mr. Anthony Downes, Mr. George G. Lewis, and many other friends for helpful discussions. Since then C.W. has continued leaf-miner studies with the help of his research group. Malcolm J. Scoble. M.J.S. wishes to acknowledge the help and supervision of the senior author in his part of the work which was successfully submitted to the Council for National Academic Awards for the degree of Master of Philosophy (1974). He would also like to thank Dr. K. Sattler of the British Museum (Natural History) for his helpful discussions.

The authors are indebted to research assistant Mr. Philip J. Newton who is studying the Nepticulidae of the U.S.A. also to Mrs. Diana Wilkinson and Mrs. E. du Plooy for typing the manuscript. We wish to acknowledge the following persons and their respective institutes for loans of material: Dr. D.F. Hardwick, Agriculture Canada, Ottawal Mr. O. Lindquist, WILKINSON AND SCOBLE: THE NEPTICULIDAE OF CANADA 5 Environment Canada, Sault Ste. Marie; Dr. G.G.E. Scudder, University of British Columbia; Dr. D.R. Davis, United States National Museum, Washington D.C.; Dr. W.W. Moss, Academy of Natural Sciences, Philadelphia; Dr. A.F. Braun from Cincinnati, Ohio; and Dr. K. Sattler, British Museum (Natural History), London; also staff of the Museum of Comparative Zoology, Cambridge, Massachusetts in which a number of types are deposited. We are grateful to Portsmouth Polytechnic UK, the Biosystematics Research Institute, Ottawa, Canada and the Transvaal Museum, Pretoria, RSA for providing facilities to enable this research to be carried out.

TAXONOMIC HISTORY The nomenclature and systematics of the Nepticulidae have been the source of much controversy throughout the taxonomic history of the family. There has been an inconsistent use of the family name resulting in part from the debate regarding the valid name for the genus. Some have favoured Stigmella Schrank and others Nepticula Heyden (see Wilkinson, 1978). We use Stigmella here. Some species have also been attributed to wrong genera. For simplification, the views of several prominent workers on the Nepticulidae are presented in Tables I-III. The classification proposed for the Canadian Nepticulidae in this work is given in Table IV. The valid name of the family is Nepticulidae, Stainton (1854a, b) being the first to use it. However, other writers, e.g. Beirne (1945), have used the name Stigmellidae presumably because Stigmella Schrank (1802) was the oldest genus, from which they named the family correspondingly. Schrank (1802: 169) described the genus Stigmella. He immediately followed this description with the words "Ich meyne, dass die mir nicht hinllnglich bekannte Motte, welche die Rosenblhtter gangweise minirt, hieher gehcire". [I am of the opinion that the , not sufficiently known to me, which mines galleries in rose leaves belongs here.]

Table I. From Beirne (1945). STIGMELLIDAE

Stigmella Nepticula Dechtiria Scoliaula Levarchama Trifurcula Etainia Fomorru

(all of generic rank)

Table II. From Johansson (1971). NEPTICULIDAE

Genera: Nepticula (: Stigmella) Trifurcula

I Subdivided into species-groups

Subgenera Fedalmia Ectoedemia Fomoria Trifurcula Levarchama Scoliaula (: Dechtitia) (s. slr. ) 6 THEENI.M.LOGI.ALSocIETyoFCANADA 'EMOIRSots Walsingham (1907: 1008) assumed that Schrank, in this statement, was referring to a species that he (Schrank) described asTinea rosella 30 pages earlier in the same paper (p. 139). Schrank's vague referencetorosella is inadequate to fix it as the type-species of Stigmella. Article 67(c) of the International Code for Zoological Nomenclature states that "The term 'designation' in relation to the fixation of a type-species must be rigidly construed; a designation made in an ambiguous or qualified manner is invalid. (l) Mention of a species as an example of a genus does not constitute a

type-designation' ' . The type-species of Stigmella was actually fixed by Walsingham (1907: 1008) under the provision of Article 69 of the Code which deals with nominal genera established before 1931 without an originally designated or indicated type-species. In particular, section (a) (iii) of this Article states that "In the absence of a prior valid type-designation for a nominal genus, an author is considered to have designated one of the originally included nominal species as type-species, if he states that it is the type (or type-species), for whatever reason, right or wrong, and if it is clear that he himself accepts it as the type-species. That Walsingham did accept rosella as the type-species is clear from his comment: "It is however obvious that his [i.e. Schrank's] remark 'Ich meyne, etc. .', refers to rosella Schrank . .".

Table III. From Borkowski (1972n\. NEPTICULIDAE Genera -T --l Nepticula Johanssonia Obrussa Scoliaula+ Fomoria Etainia Fedalmia Trifurcula Levarchama (.: Stigmella)

Ectoedemia

Subgenera: Ectoedemia Zimmermania Dechtiria (Petiole-miner) (Bark-miner) (Leaf-miner) +j.syn. of Stainton (see Kloet and Hincks, 1972).

Table IV. Treatment of the Canadian Nepticulidae in this study

NEPTICULIDAE

Genera Stigmella Obrussa Glaucolepis (: Nepticula) (: Etainia)

Subdivided into specles-groups

E. populelkt the argyropeza group E. lindquisti the clemensella group THE NEPTICULIDAE OF CANADA

In the same paper, Walsingham pointed out that rosella was a junior synonym of anomalella Goze (1783) and so the type-species of Stigmella is anomalella Goze, subsequent designation by Walsingham (1907). The genus Nepticula Heyden, 1843 was synonymized by Walsingham (1907) because Stigmella Schrank was the oldest name. The issue is further complicated since Beirne (1945) regarded Nepticula and Stigmella as two separate genera. As both names had been previously used to refer to the same genus, it was rather an unfortunate decision. Also, the differences in wing venation and genitalia between Stigmella and the other Canadian genera compared with Stigmella serzsz Beirne and Nepticula sensr Beirne is disproportionately great. Borkowski (197h , b; 197 5) and Johansson ( I 97 I ) also unite Sti gmella and Nepticula in Beirne's sense, in one genus. Apart from Stigmella, Johansson (197 l) treated all the genera of the Nepticulidae with a tongue-shaped gnathos in the male and signa reticulata on the bursa copulatrix of the female, as subgenera ofTrifurcula (see Table II). Johansson suggested that to give generic status to the genera as given by Beirne, 1945 (see Table I) was unwarranted because the species of "Nepticula" could also be divided "into about thirty smaller units of about the same rank as the aforementioned subgenera" (i.e. as given in Table II). Actually, in Canada, Stigmella, Ectoedemia, Obrussa, andGlaucolepis can all be distinguished from each other by wing venation (see their respective diagnoses) although within the genera this is constant. Borkowski (1912n) working on the European Nepticulidae illustrated clear differences in venation between Beirne's genera. This paper puts the genera on a firmer basis than before. We rreatEctoedemia as a full genus: Trifurcula sensu stricto is not represented in the Canadian collections. Trifurcula was originally described by Zeller (1848) who included two species, pallidella and immundella. T. pallidella became the type-species by subsequent designation (Tutt 1899: 354). The European species were revised by Klimesch (1953). Ectoedemia was described by Busck (1901:91) who fixed E. populella as the type-species by original designation and monotypy. Svensson (1966) synonymized Dechtiria Beirne with Ectoedemiz. Borkowski (19'72{t) agreed with this synonymy and tentatively suggested dividing the genus into three subgenera, each reflecting a different feeding habit (see Table III). As the Canadian material does not include specimens of the bark-mining Ectoedemia species-which comprise the subgenus E. (Zimmermania) of Borkowski-in this study Ectoedemia is more simply subdivided into species-groups. Although the male genitalia are characteristic (see Fig. 4l and the description and diagnosis ofthe genus p.13-14) the homogeneity ofthe external characters has led to a number of incorrect generic identifications by those authors who did not examine the genitalia. In this study seven species previously regarded as'Nepticula' are now taken to be congeneric with Ectoedemia populella (the type-species). Actually both Busck ( 1913) and Braun (l917b) showed that they were aware that clemensella and its relatives had wing venation indistinguishable from E. populella. However, they did not include them in the genus Ectoedemia, mainly because the larvae mine purely in the leaf-blade and do not cause the formation of petiole galls as do populella larvae. Busck's formal description of Ectoedemirz does not specifically exclude leaf-miners; he discussed this in a more suggestive rather than emphatic tone later in the paper. It therefore seems justifiable to include these other species in Ectoedemia, thus somewhat extending Busck's original concept of the genus. Obrussa Braun (p. 98) is represented in Canada by two species. Braun (1915: 196) made the type-species Nepticula ochrefasciella Chambers by original designation and 8 uBMOTRS oF ENT.MOLoGICAL socrEty oF .ANADA 'HE monotypy. The characteristic genitalia (Figs. 56, 57) show that a species previously recorded only from Europe, which we have now found in North American collections, is congeneric. Glaucolepis Braun is monotypic. The genus was described by Braun (lgllb), type-species Nepticula saccharella Braun by original designation and monotypy.

NEPTICULOIDEA Nenticulidae Stainton. 1854a: 295. 1854b: 166 Stigmellidae Hampson, 1918: 387 F,tntlv DsscntprroN. Adult. ErternaL features. Head: antennae filiform with basal segment enlarged to fbrm'eye-cap'; palps extend beyond labrum; maxillaries twice length of labials; tongue very short: front of head and vertex tufted: 'collar' present between head and thorax, comprising two tufts of scales. Thorax and abdomen usually concolorous dorsally; ventral surface of abdomen pale. Abdomen with anal tuft in male, presumed to function in scent dispersal. Wings: aculeate. Forewing petiolate when denuded of scales, rounded apically when scales present; marked with fasciae or patches or unmarked; tegulum comprising flap of scales, concolorous with forewings overlapping base of forewings along their leading edge; retinaculum subcostal in male, subdorsal if present on female; fringe slightly shorter than width of wing. Hrndwing lanceolate; generally light brownish grey; fringe up to 4 times as long as width of wings. Frenulum: comprising single, strong spine in male, minute functionless bristles in female; costal spines present in female, function in wing-coupling. Retinaculum: in male usually in form of a series of heavily sclerotized, hooked scales on underside of subcostal vein; in female generally a series of long scales situated on lower surface of forewing, just above fold, but occasionally subcostal. Wing coupling in maie by frenulum-retinaculum association; in female by association of costal spines and retinaculum. Jugum sometimes present in both male and female. Venation (Figs. 1-5): reduced. Forewing: Rs with R2*3 always fused throughout lengths, R4 and R5 sometimes fused, otherwise represented as separate branches; Cu either separate tiom, or coalescent with M fbr a part of its length; M sometimes fused with Cu fbr part of its length, sometimes separate, but always fused with Rs for part of length, sometimes present as one branch, sometimes as Ml and M2; Cu separate, or fused with M for part of its length; 2A prominent, srtuated below fold. Hindwing: Sc fused with Rl;Rs and M coalesced basally; Rs unbranched; M either a single branch or present as Ml and M2. Legs: foreleg-epiphysis and tibial spurs absent: midleg with pair of apical spurs on tibia; hindleg with medial and apical pair of tibial spurs, the most proximal of which vary in position according to genus. (Tibial spur formula o-2-4.\ Genitalia d. Lateral arms of vinculum either U-shaped or forming complete rine; tegumen sometimes extended into pseuduncus, in which case true uncus is membranous; saccus large; gnathos, usually well sclerotized; valves often divided into a pointed style and broad cuiller, joined by transtilla comprising pair of lateral arms and horizontal bar; aedeagus large, sometimes twice length of genital capsule; vesica with cornuti generally spine-like, often adorned with striate thickening. 9. Genital aperture single, situated on sternite 8: ovipositor not attenuated; ductus bursae short with accessory sac sometimes adorned with denticles; signum, or paired signa, sometimes present in bursa copulatrix. Btor-ocv. Egg. Laid on the surface of the food plant, not inserted into the tissue. The chorion is translucent but the hatched egg may appear black because dark frass is ejected into it by the mining larva (Lindquist and Harnden 1970: l29l). Larva. Prognathous, eats its way out of the egg into the host-plant. By consuming parenchyma it makes a tunnel, called a mine, the pattern of r.r'hich may be characteristic of a given species. Most species are leaf-miners, but there are petiole-and bark-miners. One species feeds in the fruit of Acer. Cocoon. Oval and silken, spun by the mining larva. It is brownish or yellowish. The larva pupates within the cocoon. Voltinism. The first generation of adults emerges in the spring. In most species there can be two or three generations per year. WILKINSON AND SCOBLE: THE NEPTICULIDAE OF CANADA

COstal spines

2A 2 Rs

A

Ftcs. l-3. Wingventation: l,Stigmellaslingerlandella;2,TrifurculapaLlidella;3,Ectoedemiapopulella 10 MEMOIRS OF THE ENTOMOLOGICAL SOCIETY OF CANADA

sericoPeza 4 obrussa ']

$ ctaucoteois saccarella

FIcs. 4-5. Wing ventation: 4, Obrussa sericopeza;5, Glaucolepis saccharella' WILKINSON AND SCOBLE: THE NEPTICULIDAE OF CANADA il A Llsr op GeNBnA,, Specrps, lNo SusspeclEs IN Ca,Nnoe Stigmella Schrank, 1802 bfasciella Clemens, 1862 Nepticula von Heyden, 1843; j. syn. serotinaeella Chambers, 1873 rosaefoliella Clemens, 1861 intermedia Braun, 1917 r. rosaefoliella Clemens, 1861 querc ipulc hella Chambers, I 878 r. pectocatena ssp. n. terminella Braun, 1914 slingerlandella Kearfott, 1908 variella Braun, l9l0 fuscotibiella Clemens, I 86 1 pr o c r as tine I la Br aun, 1 927 ciliaefuscella I Chambers, 873 alba sp. n. discolorella Braun, 1912 Ectoedemia Busck, 1907 populetorum Frey and Boll, 1878 Dechtiria Beirne, 1945 aromella sp. n. T r ifur c ula Zeller ; s e ns u crataegfoliella Clemens, 1861 Johansson, 1971. [Partim] scintillans Braun, 1917 populella Busck, 1907 pomivorella (Packard 1870) chalybeia Braun, 1914 ar gyrop eza (Zeller, 1 839) sr:inanella sp. n. (a. argyropeza (Zeller, 1839). European) stigmaciella sp. n. a. downesi ssp. n. canutus sp. n. saginella Clemens, 1861 quercicastanel/a Chambers, 1873 lindquisti (Freeman, 1962) clemensella (Chambers, I 873) fuscocapitella Chambers, 1 873 castaneaefoliella Chambers, 1875 platanella (Clemens, 1861) latifasciella Chambers, I 878 rubifoliella (Clemens, I 860) macrocarpae Freeman, 1967 ulmella (Braun, 1912) quadrinotata (Braun, l9l7) corylfoliella Clemens, 1861 chlorantis Meyrick, 1928: 462* virginiella Clemens, 1861 1915 minimella Chambers, 1 873 Obrussa Braun, opulffoliella Braun, 1914 Etainia Beirne, 1945; syn. n. paludicola Braun, 1917 Tr ifur c ula Zeller ; s e ns u exasperata Braun, 1930 Johansson, 1971 [Partim] ostryaeJoliella Clemens, I 86 I ochrefasciella (Chambers I 873) caryaefoliella Clemens, I 861 seric opeTa (Zeller, 1839) obscurella Braun, 1912 Glaucolepis Braun, 1917 juglandfoliella Clemens, 1861 saccharella (Braun, 1912) myricafoliella Busck. 1900

&Is known from single I in BM(NH), not discussed here, but in paper in preparation.

Knv ro Tns GrNnnA, oF THE CnNlrr.q.N NnptrcuLroe.E, Gnathos with tongue-shaped medial projection in male genitalia; vinculum ring-shaped; tegumen extended into pseuduncus; true uncus membranous. Female genitalia generally with a pair of reticulated signa on bursa copulatrix, or if not then with paired signa comprising rows of characteristic pectinations as in Fig. 61. Forewing with M and Cu fused for part of their length, and R4 separate from R5 ..... 2 Gnathos usually lacking tongue-shaped medial projection in male genitalia; vinculum U-shaped, fusing with sides of tegumen; tegumen not extended as a pseuduncus; uncus sclerotized. Female genitalia with neither a pair of reticulated signa on bursa copulatrix nor signa as Fig. 61. Forewing with M separate from Cu and R4+5 fused . . Srigmella (p. 12) Male genitalia with pair of large spine-like processes arising from valves (e.g. Fig. 56). Female genitalia with antrum (e.g. Fig. 57) . . . . Obrussa (p. 98) Male genitalia without spine-like processes arising from valves. Female genitalia without antrum I Z MEMoIRS oF THE ENToMoLocIcAL soclETY oF cANADA

3. Aedeagus with characteristic, long, hooked spines extending nearly its entire length (Fig. 60). Female genitalia with paired signa on bursa as in Fig. 61. Forewing with M+Cu fusing with Rs to form R+M+Cu. Hindwine with M branching into Ml and M2 Glaucolepis (p. 102) - Aedeagus never with long, hooked spines. Female genitalia with pair of reticulate signa on bursa (e.g. Fig. 42). Forewing with M arising from M*Cu and meeting Rs to form R*M. Hindwingwith Munbranched ...... Ectoedemia(p.73)

GENUS STIGMELLA Schrank Svnonymy and selected references: Stigmella Schrank, 1802: 169. Type-species by subsequent designation (Walsingham, 1907: 1008), Phalaena Tinea anomalella Goze, 1783: 168 : Tinea rosella Schrank. 1802: 139 Stigmellet Schrank; Walsingham, 1907: 1008 Stigmella Schrank; Hampson, 1918: 387 Stigmella Schrank; Fletcher, 1929: 210 Stigmella ' ' Heyden' ' ; Beirne, 1945: 19'7 . (Incorrect author given by Beirne. Inconectly synonymized with Nepticula Heyden by Borkowski, l97h: 702) Stigmella Schrank; Klimesch, 1948: 49 StigmeLla Schrank; Borkowski, 1969: 96 Stigmella Schrank; Emmet in Heath, 1916 213 Stigmella Schrank; Wilkinson, 1978: l3 Nepticula Heyden, 1843: 208. Type-species by subsequent designation (Tutt, 1899: 184),Tinea aurella Fabricius, l'775:666 (syn. by Walsingham' 1907: 1008) Nepticula Heyden; Zeller. 1848: 301. (Incorrectly called new genus) Nepticula Heyden: Stainton. 1854a: 295 Nepticula Heyden: Stainton, 1854b: 166 Nepticula Heyden; Stainton, 1855: 2 Nepticula Heyden; Tutt, 1899: 184 Itlepticula Heyden; Dyar, 1903: 545 Nepticula Heyden; Busck, l9l3: 103 Neptic'ula Heyden; Braun, 19lla: 233 Nepticula Heyden; Braun, 19llb: 162 Nepticula Heyden: Braun ln Forbes. 1923: 83 Nepticula Hevdenl Petersen, 1930: I l,leptic:ula Heyden; McDunnough. 1939: 107 Nepticula Heyden; Beirne, 1945'. 201 l,{epticula Heyden; Klimesch, 195 l: 4 l{epticula Heyden; Johansson, l9'7 l: 211 Nepticula Heyden; Borkowski, 1912a: 690 Nepticula Heyden; Borkowski, 1912b: 161 Microsetia Stephens sen.su Kirby, 1897: 313. Incorrect designation Neptic'ula mic'rotheriella Stainton, 1854a: 302. Mic:rosetia, in any case, belongs to the Gelechiidae WTLKINSON AND SCOBLE: THE NEFTICULIDAE OF CANADA 13

DescrupuoN: External features: d, I . Head: antennae rather less than half length of wings. Wings. Venation as in Fig. l. Forewing with Cu separate from M; M not therefore crossing to Rs; R4 and R5 fused to form R4*5; hindwing with Rs and M each represented by single branch fusing half way along wing to form common trunk. Legs: hindleg with proximal pair of tibial spurs not below middle of tibia. Genitalia: d. Vinculum U-shaped; tegumen short, joining posterior ends of lateral arms of vinculum; gnathos generally M-shaped, U-shaped, or comprising pair of horns arising from basal sclerite;r aedeagus more variable in length than width; vesica usually with spine-like or denticulate cornuti, and often with striate thickening. I . Ductus bursae with accessory sac sometimes adorned with denticles; bursa copulatrix with variable density of minute pectinations; signa, if present, rather weakly demarcated.

DIecNosrs. Stigmella has R4*5 in the forewing united whilst in Trifurcula, Ectoedemia, Obrussa, andGlaucolepjs R4 and R5 are present as separate branches. Cu and M are not united in the forewing of Stigmella whereas they are always tused for a part of their length in these other genera. ln the hindwing, M which is unbranched inStigmella, is represented by Ml +2 and M3 in Trifurcula. In the male of Stigmella the lateral arms of the vinculum form a U-shape whilst in the other genera they meet so forming a ring. The female of Srigmella lacks signa reticulata on the bursa copulatrix which are present rnTrfurc'ula, Ectoedemia, and Obrussa.

Knv: SrrcuE,r-r-e. l. Male with snathos in form of inverted "V Female with signum of bursa comprising heavily sclerotized denticles . . . . the procrastinella group (p. 70) - Malewithgnathosnotuninu.rt.d"V".f.*uf.withsignumoibu.ruabsentor,ifpresent, not comprising heavily sclerotized denticles ...... 2 2. Female with accessory sac not as in Fig. 37. Male with aedeagus rarely more than twice length ofcapsule ...... 3 - Female with accessory sac in form of characteristic spiral (Fig. 37). Male with aedeagus almost twice length of capsule the quercipulchella group (p. 65) Female with signa of bursa as paired, weakly defined, patches of denticles. Male with aedeagus flask-shaped and with pair of characteristic plates at phallotreme'z (Figs. 27,29,31) . the c'orylifollella group (p. 50) Female with signa of bursa, if present, not as patches of denticles. Male with aedeagus not flask-shaped and without plates at phallotreme ...... 4 4. Male with homs of gnathos arising from large basal sclerite, the corners of which are produced into short lateral processes. Female genitalia: if signum on bursa present then accessory sac unadorned the rosaefoliellz group (p. 14) Male with gnathos lacking large basal plate. Female genitalia: if signum on bursa present then accessory sac of ductus adomed with denticles ...... 5 5. Male with well developed scale-like hairs arising from tip of style. Female with signum. or pair of signa, present on bursa the saginella group tp. 39t Male without well developed scale-like hairs arising from tip of style. Femaie without signum, or pairolsigna.onbursa ...... 6 Male: valves with clearly defined, rectangular, cuiller; cornuti of aedeagus densely covering vesica; uncus with pair of lobes .... the crataegfoliella group (p. 30) - Male: valves with poorly defined cuiller; cornuti of aedeagus never densely covering vesica; uncus in form of characteristic cross the bifasciella group (p. 59)

' S . braunella (Jones) comb. n. a species known from Califomia, appetrs to have a W-shaped gnathos simrlt to Ectoedemia species. Other features of the genitalia, and the wing venation, show that braunella falls into the genus Stigmelln. z A tem coined by Snodgrass: the opening of the endophallus, usually at the end of the aedeagus. I 4 MEMoIRS oF THE ENToMoLocrcAr, socrL r y ots crAN ADA

Kpv: the RosAEFoLTELLA GRoUP L Female with signum on bursa copulatrix ...... 2 - Female without signumonbursacopulatrix ...... 4 2. Forewings dark each with shining silver fascia. Male genitalia with valves not symmetrically bilobed .....fust'otibiella (p.23) - Forewings palelime-green. Malegenitaliawithvalvesbilobed ...... 3 3. Abdomen of male orange-ochreous, with invaginated scent pouches ventrally - atoo*"n;f -"i;darkish, unJiu.ttng r..nipou.t", ...... :: ,"r\iiTflr|frf.i 3Zl 4. Male: gnathos with homs closely juxtaposed; transtilla as in Fig. 8. Larvae form blotch mines .. slingerlunlellu tP l9\ - Male: gnathos with horns relatively widely separated; transtilla as in Fig. 6. Larvae form linear s illrt"l. *;in u"rr^ .opututri* ."".r"0 *iit .."1"p-r1-t*6 p".tlnuilon, rorr'.\"rt"{Ii^l':":t1 cornuti of aedeagus spiculate . . . . . r. rosaefolielLa (p. 11) - Female with bursa copulatrix covered with chain-like pectinations. Male with comuti of aedeagus in form of denticles ..... r. pectocatena (.p. 18)

The nosaERot-IEI-r-A GRoUP The group is characterized by the male and female genitalia. In the male the uncus is bilobed and papillate and the gnathos comprises a pair of medial, posteriorly directed horns and a pair of more widely separated, shorl lateral projections situated ventral to the horns. The cornuti on the vesica of the aedeagus are spine-like or spiculate. The ductus bursae of the female is rather short and leads to a wide bursa copulatrix, uniformly covered with minute pectinations, sometimes with a band-like signum. The larvae mine leaves of certain of the Rosaceae and Salicaceae.

Stigmella rosaefoliella (Clemens) comb. n. Nepticula rosaefoliella Clemens, l861: 85 Neptit:ula rosaefoliella Clemens; Chambers ln Hayden, 1878: 158 (listed) Nepticula rosaefoliella Clemens; Dyar, 1903: 547 (listed) Nepticula rosaefoliell.a Clemens; Braun, 1917b: 184 Nepticula rosaefolielLa Clemens; Braun in Forbes, 1923: 9l Dsscr.tpttoN. External .features: d, 9. Head: palps grey and lustrous; antennae grey; eye-caps white with some buff scales; tuft on front of head and vertex orange-ochreous; collar off-white. Thorax and abdomen brownish black. Wings: ground colour of forewings very dark greyish brown or brownish black with grey lustre and bronze and purple reflections; one white, postmedial fascia shining silver; fringe grey. Legs grey-brown, lustrous, with irregular off-white patches. Variation: the species has a variable fascia width, sometimes being very narrow and sometimes broad. This is not apparently a sexual or subspecific difference. Genitalia: d. Figure 6; tegumen bluntly rounded; saccus wide, bilobed; uncus with 2 papillate lobes; gnathos with a pair of long, horn-like spines each with a broad, short spine laterally; valves reaching uncus, divided into a broad rounded cuiller and small spine-like style; transtilla stout, if lateral arms are not joined medially they end in swellings; aedeagus robust, straight, with a patch of spine-like or spiculate cornuti and usually with rectangular striate plate. 9. Figure 7; anal papillae shorter than broad; bursa copulatrix covered with minute scallop-shaped or chain-like pectinations; signum absent; anterior apophyses arcuate, narrow, widening terminally into plate; posterior apophyses straight and narrow. Host-pla.nts: Rosa spp. in particular Rosa virginiana, R. setigera. Mine: upper surface ophionome. WILKINSON AND SCOBLE: THE NEFTICULIDAE OF CANADA l5

9nathos

Aedeagus

Fatera I -E I arm iI .:=l I venlrar > | Plats

Frc. 6. Stigmella rosaefoliella pectocatena, d genitalia.

DracNosrs. The species is close to both the North American slingerlandella which feeds on Prunus and the European anomalella Gijze which feeds on Rosa. However, rosaefoliella differs from anomalella in having a fascia, in the forewing ground colour, and in the colour of the tuft on the vertex. Differences in the genitalia are given under the diagnoses of the two subspecies into which rosaefoliella is here divided. S. rosaefoliella differs fuom slingerlandella in the wider separation of the horn-like spines of the gnathos and the shallower lobing of the saccus. The lobes of the uncus in siingerlantlello ute pointed and there are differences in the horizontal bar of the transtilla and in the^ot" spine-like cornuti. The ophionome of rosaefoliella contrasts with the ophistigmatonome of slingerlandella. DrscussroN. S. rosaefoliella was originally described by Clemens (1861: 85) from the mine, together with brief remarks on the larva. He failed to designate any types from his material. The original mined leaves do not exist and there is nothing to suggest that Clemens ever preserved them. The imagines remained unknown until Braun (1912: 89) described them from specimens she had reared from mines similar to Clemens' description. We designate as neotype, for the nominate subspecies, one of the specimens reared by Braun. This is a male, mounted together with a mined leaf which is viry probably the one from which it was bred. It bears the following data: "8.741 . Cincinnati, O. Annette F. Braun i. V.13.13. Nepticula rosaefoliella Clem." I6 MEMOIRS OF THE ENTOMOLOGICAL SOCIETY OF CANADA

Poster i o r a pophysis

anter i or aFoFhYsis

bursae

ssory sac

w b

a

Ftc. 7. Stigmella rosaefoliella pectocatena, I genitalia WTLKINSON AND TOBLB: IHb NEPTICULIDAE OF CANADA I1 On the basis of the aforementioned specimens together with material from the CNC, we regard rosaefoliella as being divisible into two subspecies. One of these has been found in the U.S.A. and Canada, and the other in Canada alone. DIsrnIsurroN. Canada: Ontario. U.S.A.: Ohio, Pennsylvania, Arkansas, New York, Michigan, Missouri, S. Dakota. Bror-ocy. Egg. Laid on the upper or lower surface of the leaf, often beside the mid-rib or a vein. Mine. For the first half of its length the mine may run alongside a vein and then double back on itself. The larva often follows the serrations of the leaf for part or all of its mining life. Occasionally, when the leaf is very small, the larva crosses the mid-rib. Frass usually fills the mine for the first third of its length and then the frass-line narrows relative to the width of the mine and the grains of frass tend to become disseminated in a wavy pattern (see Pl. 1.1). Yottinism. Canada: Rearing records suggest bivoltinism. U.S.A.: Trivoltine.

Stigmella rosaefoliella rosaefoliella (Clemens) comb. n. Wing measurementsi 6 . 4.0-4.4 mm (7); I. 3.8-5.0 mm (8); neotype. 4.4 mm' DlecNosrs. Differs from rosaefoliella pectocatena in having scallop-shaped rather than chain-like pectinations on the bursa copulatrix. (cf. Fig. 7a and b). The spiculate rather than spine-like cornuti are characteristic ofthe nominate subspecies and the striate plate is weakly defined or absent. The horizontal bar ofthe transtilla is shorter in the nominate subspecres. DrscussroN. Although the external features and the mine are indistinguishable from the next subspecies, on the basis of the specimens examined, the differences, particularly in the female but also in the male genitalia, suggest the division of rosaefoliella into the two subspecies. Because Clemens failed to cite type material we have based the description of the nominate subspecies on material which includes the specimens bred by Braun and from which she first made known the adults of the species rosaefbliella. DIsrnreuttoN. Canada: Ontario. U.S.A.: Ohio, Pennsylvania, Arkansas, New York. Michigan. Missouri. SpEcrnrsNs Exa.nrNnr. d neotype. In Phila: U.S.A: Ohio, Cincinnati; on Rosa setigera, B.'741; 13.v.1913 (Braun); USNM slide No. 17360. Other specimens: CANADA: Ontario, Simcoe; on Rosa; I d, 3 ? ? , 11-12.viii.1969 (Freeman). U.S.A.: Pennsylvania, West Chester; on Rosa; 26 6, 19, I ex, l-21.ii.1966 (Lewis); in CNC. Data as neotype, 5dd, 899, 6.v.-6.vi.1913, 28.iv.-6.v.1918,2? I, 8.473, 20,21.v1i.1909; in Phila. No locality data;on rose; 19, I ex, 14.iii.1889 (ex Murfeldt Coll.); Missouri, Kirkwood, I ex,20.iii.1906 (ex Murfeldt Coll.), Cornell University Lot 51. Sub 619); 2? I , I ex, 20.vi.1889, I ex, no date (?Murfeldt), no other data; I ex, from rose, no data; Michigan, I ?, 5.vi.1966 (Hillman); Arkansas, Washinqton Co., Devil's Den St. Park, 1d, 10.vii.1966 (Hodges); in USNM.

Specimen a There is a male with tuft on front of head pale ochreous to whitish; tuft on vertex dark brown. Although the genitalia show the spiculate cornuti and rather poorly defined 1 8 MEMOIRS OF THE ENTOMOLOGICAL SOCIETY OF CANADA patch on the vesica, characteristic of the nominate subspecies, the specimen is mentioned en passant to draw attention to the difference in colour of the tuft on the vertex from the usual bright orange-ochreous of rosaefoliella. Wing measurementi 4.0 mm.

SpsctueN Ex,q.uINr,o. U.S.A.: South Dakota, Ipswich; on Rosa; 1 6, 15.i.1973 (Lewis): in CNC.

Stigmella rosaefoliella pectocatena ssp. n. Wing measurements: 6 . 4.4-5.2 mm (9); I . 4.6-5.0 mm (6).

DrncNosrs. Distinguished from the nominate subspecies by the chain-like rather than scallop-shaped pectinations on the bursa copulatrix of the female (Fig' 7) and by the stout spine-like cornuti on the vesica of the aedeagus (Fig. 6b), together with the shorter horizontal bar of the transtilla in the male (see diagnosis of rosaefoliella rosaefoliella). It differs from the European species anomalella Gijze in having a bright orange-ochreous rather than reddish brown tuft on the vertex. The forewings are much darker and bear a fascia.

DrscussroN. We select a female as holotype because the pectinations on the bursa copulatrix, from which the subspecies derives its name, is the least variable and most clearly diagnostic feature from the nominate subspecies.

DrsrntsutIoN. Canada: Ontario.

SpecIr4nNs ExlltrNpo. t holotype; in CNC: CANADA: Ontario, Ottawa; on Rosa;24.ix.1962 (Freeman and Lewis); slide No. 3292; in CNC. Paratypes: 9d d , 8I I , data as holotype; l.vi.-4.ix. 1956, 11-28'ix.1962, 9-13.v.1963.

Ervuot-ocv. Catena (Gk.)-chain.

Specimen b There is a male specimen reared from Rosa, collected in Ottawa, which differs from the species description of rosaefoliella inthe following ways: tuft on front of head and vertex dark brown not orange-ochreous; genitalia with strongly curved apical hook on valves. The stout spine-like cornuti on the vesica of the aedeagus together with the locality indicates that it is \earer to pectocatena ssp. n. than to the nominate subspecies. Wing measurement'. 4.8 mm. Spnctur,N ExaMrNE,n. CANADA: Ontario, Ottawa; on rose; 1d, 4.vi'1956 (Lewis)'. in CNC.

Stigmella anomalella (Goze). [Type-species] fEuropean] Selected references: Phalaena Tinea anomalella Goze, 1783: 168 Nepticula anomalella (Goze); Tutt, 1899: 206 (Giize); Walsingham, 1907: 1008 Tinea rosella Schrank, 1802: 139 (syn. by Walsingham, 1907: 1008) Stigmella rosella (Schrank); Walsingham, 1901 1008 (designation of anomalella, senior syn. of rosella, as type-species of Stigmella by subsequent designation) WILKINSON AND SCOBLE:IHB NEPTICULIDAE OtsCANADA 19

The male genitalia of this species are figured by Beirne ( 1945) . We have examined Beirne's original slides of the male and female genitalia (for details see diagnosis): these are lodged in the BM(NH). The adult specimens from which these slides were made are missing, but we have checked the external features of specimens in the main collection of the museum. Borkowski ( 1969) gives a photograph of the mine of this species which does not appear to differ from the North American specimens of rosaefoliella. Both species mine Rosa. DtecNosrrc Fr.4run6s. d,9 . Head: tuft on front of head ochreous; tuft on vertex dark reddish brown. Wings: ground colour of forewings greyish brown with bronze reflections; fascia absent. Wing measurementsi d.5.6 mm (1); 9.6.0 mm (l).

Dr.q.cNosrs. S. anomalella has a reddish brown tuft whereas that of rosaefoliella is orange-ochreous. ln anomalelkt there is no fascia on the forewing which there is in rosaefoliella. The capsule of the male genitalia of anomalella is more elongate than in rosaefoliella.

DrscussroN. This species is not known to occur in North America but because of its similarity torosaefoliella and its considerable significance in the genus, an account has been included. The species was first well described, vlz: moth, larva and leaf damage, and figured by DeGeer in 1152 but this predates Linnaeus' 1Oth edition of Systema Naturae and so the original name is not available. DeGeer referred to it again in 111 | but Gcize was the first to name it in 1783 as "Phalaena Tinea anomalella". The species subsequently became the type-species of Stigmella Schrank. Schrank's species rosella is a junior synonym. S. rosaefolielLa and anomalella are treated as separate species rather than subspecies because we did not examine sufficient material of anomalella and because of the differences in the external features. The similarities in the genitalia, food-plant, and the mine suggest that in future anomalella and rosaeJoliella may be treated as subspecies of the same polytypic species. DrsrnIsurtoN. Europe: lncluding British Isles. Spr,crNrE,Ns Exe.utNr,o. ld, 1890 (Frey Coll.); ENGLAND: l9, 1910 (Walsingham Coll.). Genitalia slide, d, B P. Beirne 1944 (Fletcher Coll.) plus others in the collection of BM(NH).

Stigmella slingerlandella (Kearfott) comb. n.

Neptic:ula slingerlandella Kearfott, 1908: I 87 Nepticula slingerlandella Kearfott; Crosby, lgll: 219 (life history, pest status, and control) Nepticula slingerlandella Kearfott; Crosby, 1912: 25 (life history) Nepticula slingerlandella Kearfott; Braun, 1911b: 184 Nepticula slingerlandella Kearfott; Braun in Forbes, 1923: 9l

Dr,scrtpttoN. External .f'eatures'. d,?. Hcad: palps grey and lustrous; antennae grey to buff, lustrous; eye-caps white; tuft on front of head and vertex ochreous; collar off-white. Thorax and abdomen greyish brown. Wings: ground colour of forewings brownish black with single, narrow! white postmedial fascia shining silver; fringe grey. Legs grey, with oft'-white patches, lustrous. Pl. 8. I . Wing measurements: 6.4.2-6.0 mm (19); ?.4.5-5.4 mm (23); Lectotype 4.6 rnm. 20 MLM.IRS ots rHE BNI.M.LOGT.AL socIETy oF cANADA

Genitalia: d. Figure 8; tegumen rounded posteriorly, broader than long; saccus bilobed with lobes longer than wide; uncus comprising a pair of pointed papillae extending well beyond tegumen; gnathos with central spines juxtaposed and almost reaching as far posteriorly as uncus; valves divided into cuiller and arcuate, pointed style; transtilla broad, stout, and with short lateral arms; aedeagus cigar-shaped, wide with cornutus comprising short, denticulate spines and a striate, rectangular patch. ?. Figure 9; anal papillae shorter than broad; ductus bursae short; bursa copulatrix covered with minute scallop-shaped pectinations; signum absent; apophyses long and thin throughout length, broader basally. Host-plants: Prunus. Similar mines found by Crosby on Malus. M ine : an ophistigmatonome. DrRcNosts. Related to rosaefoliella but with a rather narrower fascia on the forewing. In the male of slingerlandella, the uncus has smaller papillae than rosae.foliella and the horns of the gnathos are closer together. The transtillae of the two species are different (cf. Figs. 6a and 8a). The mine of slin6erlandella is a blotch whilst that of rosaefoliella is a gallery. S. slingerlandella has an ochreous tuft and brownish black forewings whereas the closely related European species plagicolella has the tuft ochreous, tinged with red, and greyish brown forewings. ln slingerlandella the male genital capsule is less thin and elongate thanplagicoleLla, the apex of the valves are more gently, and less strongly, curved and the lobes of the uncus are not clearly subdivided.

Drscusstou. The species was first described by Kearfott (1908). There are three specimens labelled "TYPE" mounted on the same strip of polyporous. Besides this there is a separate specimen, also labelled "TYPE" to which Kearfott added a separate label "Nepticula plagicolella Stnt?" suggesting that he had second thoughts on its identity. However, we can find no difference between the genitalia of this specimen and those specimens on the first pin: we regard all of them as the type material of slingerlandella which is what Kearfott had originally implied. Unfortunately all that remains of the single specimen relabelled by Kearfott, apart from the genitalia, is a hindwing which is of little diagnostic value. We designate the male specimen from the pin bearing the three "TYPES" of slingerlandellct as lectotype and the others, including the specimen Kearfott separated, as paralectotypes. The lectotype has been remounted and labelled accordingly. Both Kearfott, by the implication of his labelling, and Crosby (191l) in his paper on the life history of slingerlandella have emphasized the close relationship of the species to plagicolella. We have examined a male genitalia slide prepared by Beirne for his work on the British Nepticulidae (1945) and also a male and female specimen from the main collection of the BM(NH). Both the externals and the male genitalia differ from slingerlandella (see plagicolella diagnoslic features) although they indeed show clear relationships. DtsrntsuttoN. Canada: Ontario. U.S.A.: Rochester, New York (type locality), Michigan, Ohio. SpEctnr,Ns ExnluNno. d lectotype; in USNM: U.S.A.: New York, Rochester; on plum; (Slingerland); genitalia, CNC slide No. 3472. Paralectotypes: 2d d, 1 ?, data as lectotype. Other specimens: CANADA: Ontario, Normandale; on Prunus nigra; 16, 19, 15,2l .iii.1951; on Prunus pensylvctnica; I 6 , I I , 3,5.iii.1958 (Freeman and Lewis); Simcoe; on Prunus 266,7 99, 30.iv.-11.v.196'1 (Freeman); in CNC, Ottawa; Prunus pensylvanica; 1 ex, 4.vi.1956 (Lewis); in CNC. Elmira; on Prunus nigra; 1766,16??, I ex,22-31.iii.1966, l'7.iti.-21.iv.1967; in Soo. U.S.A.: Michigan, W]LKINSON AND SCOBLE: THE NEPTTCULIDAE OF CANADA 2l

b

Frc. 8. Stigmella slingerlandella, d genitalia.

Van Buren Co., Tavt Cherry Orchard; 16, 2? 9, I ex, 10.vi.1966 (Hillman); Lawrence; same data; on Cherry; 5*13.v.1966; Barry Co'., ld, 19,28.v.1961 (Hodges); Ohio, Cincinnati, I ex, 21.v.1914 (Braun) in USNM. Btor-ocv. The information given here is supplemented by the work of Crosby ( 191 1 , l9l2). Egg. Crosby states that the egg is laid on the lower surface of the leaf. We have also found it on the upper surface. Mine. The linear part of this ophistigmatonome begins interparenchymally but later becomes upper surface. The blotch is upper surface and may lie directly above the linear tract. The frass-line in the linear part is broken, dark, and central. The blotch has the frass deposited in a central patch. Pl. 1.2. Larva. Greenish white with head light brown. Cocoon. Light brown. The larvae may remain in the cocoon for a considerable time before pupation. Pupa. The species overwinters in this stage. Voltinism. Crosby found the species to have a single generation per year in New York State, but Braun's specimens show that there are two generations in Cincinnati, Ohio. (Also see Braun 1911b.) In Canada the species is probably bivoltine. 22 MEMOIRS OF THE ENTOMOLOGICAL SOCIETY OF CANADA

Frc. 9. Stigmella slingerlandella, ? genitalia. other notes. crosb, ;:ffi;;"^',0',"*'i'";"T"j"? oru,o causing o^,rr:l defoliation and a lowering of quality and quantity of the fruit", crop. This was brought about by heavy infestations of mines in leaves as many as a dozen per leaf . The miner was found to have a preference for certain varieties- of plum. It is of interest that work by Porter on a leaf-mining species of Lithocollells on aspen showed that the host tended to comDensate defoliation bv the production of extra leaves (verbal communication)'

Sti gme lla p la gico le lla (Stainton) [European] Original reference: Nepticula plagicolella Stainton; 1854a:303 Wing measurements'. 6 . 4.4 mm (1); ?. 5.0 mm (1). Ducxosrs. The tuft on the head of plagicolella is ochreous, tinged with red and the forewings are greyish brown. In slingerLandella the tuft is ochreous and the forewings are brownish black. In pktgicolella the uncus is bilobed with each lobe subdivided into a pair of small papillae, the apex of the valves are sharply curved almost at right angles, und th" capsule is relatively thin and elongate. In slingerlandella the uncus is bilobed, but each lobe is not clearly subdivided, the apex of the valves are curved more gently and to a lesser extent, and the capsule is not particularly thin and elongate. DrscussroN. This species is European but is included here because of its similarity to slingerlandella. Data labels on American material indicate that some workers thought that slingerlandella was in reality an introduction of this old world species. DrstntsuttoN. Europe: Including British Isles. Sppcrl.rENs Exa.urNE,p. ENGLAND: Lancashire, Preston; 16, l?, x.1887 (Threlfall, Walsingham Coll.); in BM(NH). d genitalia slide No. 1575 B.P. Beirne (Fletcher Coll.) plus others in collection of BM(NH).

Stigmella fuscotibiella (Clemens) comb. n. Nepticula fuscotibiella Clemens, 1862: 133 Nepticula fuscotibiella Clemens; Chambers in Hayden, 1878:157 (listed) Nepticula fuscotibiella Clemens; Dyar, 1903: 546 (listed) Nepticula fuscotibiella Clemens; Braun, l9l7b: 185 Nepticula fuscotibiella Clemens; Braun ir Forbes, 1923: 92 Nepticula ciliaefuscella Chambers, 18'73: 128 (syn. by Chambers, 1875: 117) Nepticula discolorella Braun, l9l2: 86 (syn. by Braun, 1917b: 185) DnscnrprroN. Externalfedtures: 6 , 9. Head: palps grey and lustrous; antennae dark grey; eye-caps white, shining white; tuft on front of head and vertex orchreous; collar dull white. Thorax slate-grey with bronze reflections. Abdomen grey, with a silvery grey lustre dorsally and laterally and ventral surface shining white. Wings: ground colour of forewings slate-grey with- bronzy reflections, and with scales the tips of which reflect purple at a certain angle of illumination; one white, medial to postmedial fascia, of variable width, shining silver; fringe grey marginally, white at tip. Legs darkish grey, lustrous, with pale ochreous patches, tarsi grey' Wing measurements: 6.3.4-5.4 mm (7); 9.4.4-5.0 mm (7). [Pl. 8'2. Genitalia: d. Figure 10; tegumen broad and bluntly rounded apically; saccus large, bilobed; uncus reaching beyond tegumen, gradually tapering, bilobed with lobes papillate; gnathos heavily sclerotized, comprising a pair of medial, horn-like processes arising from stout base, the corners of which are produced into short, lateral processes; valves broad, bilobed apically; transtilla 24 MEMOIRS OF THE ENTOMOLOGICAL SOCIETY OF CANADA

5,t"4) t/ ///t \, [ ,/'r' ),i/ r,

Lat.r.l r3p.cl ol v.lv.

Ftc. 10. Stigmella.fuscotibiella, d genitalia. stout, with lateral arms long, and horizontal bar short; aedeagus straight; vesica with a number of stout spine-like cornuti and an arcuate, striate plate. I . Figure I l; anat papillae simplel ductus bursae short, approximately length of apophyses, with unadorned accessory sac; bursa copulatrix elongate, covered with short rows of minute pectinations and a signum comprising a band of scale-like pectinations almost encircling the bursa; anterior apophyses arcuate, broad for basal two-thirds of length, narrowing anteriorly, each with small process appearing to splinter from base; posterior apophyses thin, long, straight, swollen basally. Host-plants: Salix nigra, Salix discoktr. Mine: an upper surface ophistigmatonome. DtacNosrs. Differs from the related populetorum and aromell{z sp. n. in externals, valves, and gnathos of the male genitalia, see diagnosis of populetorum. Fuscotibiella is only known to mine Sal1.r, not Populus - the host of populetorum. DtscussroN. Braun synonymized discolorellct withfuscotibiella on the basis of the externals. Dissections of the genitalia of the cotypes of discolorella confirm this. DrsrnrsurIoN. Canada: Ontario, Nova Scotia. U.S.A.: Ohio, Pennsylvania. SplctnsNs ExeluNsn. d holotype; in Phila: "Type 7495 Nepticulafuscotibiella B. Clemens". "At light Aug. 11". Chambers No. "123". Second label added by Busck giving its type status and "AB 1902". Other specimens: CANADA: Ontario, Normandale; on Salix; | 6 , 29 ? , 4- l0.iii. 1958 (Freeman and Lewis). Nova Scotia, Ohio; on Sa/rx; 3d d, 3I 9, 29.vii-13.viii.1959 (Lewis and Freeman); in CNC. U.S.A.: Ohio, Cincinnati, on Salix discolor; I d, 1 I , syntypes of discolorella;25, 26.vii.1909 (Braun); I d, on Salix nigra;6.v.1913 I ex, 22.v|ii. 19 | 6 (Braun) ; Clermont Co., Stonelick Lake; on Salix nigra ; 2 9 9, 20.v|i. 1962 (Braun); in Phila. Pennsylvania, West Chester; on Salix; I d, I ex, 14.ii.-4.iii.1966 (Lewis): in CNC. WILKINSON AND SCOBLE: THE NEPTICULIDAE OF CANADA 25

I \ ._i

signum

Stigmella.fuscotibiella, ? genitalia 26 MEMoIRs oF THE ENToMoLocICAL socIETY oF cANADA Bror-ocy. Mine. Rather variable; sometimes an ophionome of gradually increasing width, but usually it can more accurately be described as an ophistig- matonome. The frass in the early linear part of the mine forms a dark, broken line but is much less compact in the blotch. (Pl. 1.3.) Voltinism. Probably bivoltine in both Canada and the U.S.A

Specimen c There is a male specimen with externals similar toJuscotibiella: however, in the genitalia the valves are not as deeply bilobed (Fig. 10c). The rearing records suggest that the specimen mined Shepherdia canadensls , a plant not related to Salix which is the normal host of/rscoribietta. Apparently the mine was overlooked in a series of collected leaves on which specimens of aColeophora species were feeding. The rearing record reads "l (serp) appeared (?) on foliage and not noticed": actually "serp" i.e. serpentine does not strictly fit the characteristic tract made by fuscotibiella. Unfortunately, we have been unable to locate the mine, and the accuracy of the host-plant data for this specimen is very dubious. SppcrnrnN ExnntNr,n. CANADA: Ontario, Normandale; on Shepherdia canadensis; 16 , 22.v1i.1956 (Freeman and Lewis); in CNC.

StigmelLa populetorum (Frey and Boll) comb. n. Nepticula populetorum Frey and Boll, 1818: 276 Nepticula populetorum Frey and Boll; Dyar, 1903: 546 (listed) Neptir:ula populetorum Frey and Boll; Braun, 1911b: 194 Nepticula populetorum Frey and Boll; Braun in Forbes 1923: 95 DsscnrprroN. External .t'batures:6 ,?. Head: palps whitish and lustrous; antennae dorsally dark greyish brown, ventrally light grey; eye-caps ofl'-white, weakly lustrous; tuft on front of head buff: tuft on vertex brown; collar white. Thorax off-white to buff. Abdomen dorsally grey. Wings: forewing ground colour with proximal two-thirds off--white to pale lime-yellow, weakly shining, variouily irrorate with brownish purple scales, distal third brownish purple; fiinge dorsally leaden-grey; ventrally rather yellowish. Legs pale ochreous. Pl. 8.3' Wing measurements: 6 .4.2-5.8 mm (6); ? .4.2-6.0 mm (l l). Gen-iatia.. d. Figure 12; as .fuscotibielLa except: gnathos bifurcate, median horn-like processes being closely juxtaposed and arising from biconcave sclerite; valves not symmetrically |ilobed but with a pointed style and a wider cuiller; juxta large, well sclerotized (see figure). I . As fuscotibiella. Host-plants: Populus deltoides, P. trichocarpa, P t'anescens, P. x canadensis' Mine: An upper surface ophistigmatonome. DrecNosrs. Differs fromfuscotibiella in the colour of the forewings, body, and legs. The male genitalia are distinguished by the unequal lobing of the valves in pipuletorum and the sharper and more closely juxtaposed medial horns of the gnathos. ln choice of host-plants populetorum has only been found on Populus whrlst fuscotibiella is aSallx feeder. Differs fromaromelLz sp. n. in the male: the abdomen of populetorum is dorsally grey not ochreous and the abdominal scent-scale pouches are absent. DrscussroN. Although Frey ( I 818: 271) stated that he and Boll reared a series of specimens, no types were designated. The Frey collection is mostly lodged in European institutes and we have located a specimen in the BM(NH) which is almost certainly one of Frey's original specimens of populetctrum. It is a male and is here designated lectotype: it has been labelled accordingly and bears the following data: "Lectotype. Frey Coll. Walsingham Collection 1910-421 , N. populetorum Fr. and Boll. Dallas". WILKINSON AND SCOBLE: THE NtsPTTCULIDAE OF CANADA 21

b

Ftc. i2 Stigmella populetorum, d genitalia

DtstrusurroN. Canada: Ontario, British Columbia. U.S.A.: Texas (type localiry), Ohio. SplclnrsNs Ex.qrurNn'r. d lectotype; in BM(NH); U.S.A: Texas, Dallas; 1910, onPopulus; (Walsingham Coll. ex Frey Coll.). Other specimens: CANADA: Ontario, Delhi; on Populus deltoides; 3dd, 89I, 3.viii.1966, 25.viii.-12.ix.1966 (.Freeman): Simcoe; on Populus canescens; I d, 7.iii.1958 (Freeman and Lewis): Blenheim; on Populus; l?, 21 .v1i.1961. British Columbia, Coldstream; onPopulus trichocarpa;1d, 3.vii.1951 (Freeman andLewis); in CNC. U.S.A.: Ohio, Cincinnati; on Populus x canadensis; 49 I , I ex, 28.vi.-8.vii.1911, I I , I ex, 28.vi.1914 (Braun); in Phila. Bror-ocv. Egg. Upper surface. Mine. Similar in both shape and variation to Juscotibiella. Pls. I .4,1.5.

Stigmella aromella sp. n. Wing measurements'. Holotype. 5.4 mm. Paratypes. 4.8-5.6 mm (3d d). DtncNosrs. The features which distinguish this species from populetorum are found in the males: dorsal surface of abdomen bright ochreous with no grey scales; ventrally with two medial, internal scent-scale pouches (see Fig. l3) packed with ovoid scales, formed by invagination of the intersegmental membranes one between segments I and 2, one between 5 and 6, and perhaps a small one between- 7 and 8. In MEMOIRS OF THE ENTOMOLOGICAL SOCIETY OF CANADA

a-. 2

1 i6 .' Ftc. 13. Stigmella aromella, d abdomen. WTLKINSON AND SCOBLE: THE NBPI'ICULIDAB OI CANADA 29 addition, there are patches of ovoid scales adhering to the abdomen which have, presumably, been exuded from the pouches. Host-plants: Populus deltoides, P. x c:anadensis. Mine; As far as is known indistinguishable from that of N. populetorum.

DtscussIoN. Although the invaginated pouches on the abdomens of the males have not been proved to be scent-pouches, it seems likely that they function as such: these pouches occur only in the males, as is general with lepidopteran scent-producing structures. A similar situation has been found in certain species of Striglina () by Whalley (1974). We regard these specimens as representing a separate species because specimens of both populetorum and rtromella have been collected from the same locality and reared from the same host-plant. Although the differences are relatively small the genitalia and wing colour being indistinguishable sympatric distribution of- the two forms suggests that they are probably not subspecies.- Further, scent is often species-specific, being used in courtship; such elaboration in the males of aromel/a suggests reproductive isolation. As diagnostic features have only been found in the males it is not possible to be certain about the identity of the females. We have tentatively identified those specimens from St. Williams asaromella, because all the males collected from this locality have scent pouches. Observations of courtship behaviour, collection of field data, and breeding experiments may, in the future, provide characters by which the females of the two species may be differentiated.

Dtsrntnurrou. Canada: Ontario.

SpnctupNs Exnutxr,o. d holotype; in CNC: CANADA: Ontario, Delhi; on Populus deltoides;26.v1li.1966 (Freeman); slide No. 23-M.J.S. Paratypes: CANADA: Ontario, Lake Erie, St. Williams on Populus canadensis;36 6 , 26.i't.-4.1ii.1962; tn Soo. Specimens of questionable identity: data as paratypes; 26 6 , I ex, l-4.iii.1962; in Soo. Erynror-ocy. (Gk.) Aroma - spice. Kny: the cRATAEGTFoLTELLA GRoup l. Forewings with extensive ochreous scaling ...... crataegifoliella (p.30) Forewingswithoutextensiveochreous scaling ...... 2 2. Forewings without markings, unicolorous, dull ...... 3 Forewings with fasciae or patches; brightly shining 4 3. Male with lateral bars of vinculum terminatrng in swollen knobs. Larva feeds on Pyrrzs - Mui; *irtr ru,"rur n"., ;i;;;;;t"- norierminating i" ,*"ri." o,roor. I-urul?K':"'n;"li and sometimes Crataegus . . pomivorella (p. 33) 4. Tuft on front of head and vertex dark chocolate brown. Reflections of bodv and forewinss heavily infused with purple . . scintillans (p.33) - Tuft on front of head and vertex orange-ochreous. Reflections of body and forewings heavily infusedwithred-gold ...... 5 5. Uncus of male with ear-like lobes. Accessory sac of ductus bursae without spiral thickening. Larva makes ophionome scinanella (p. 36) - Uncus of male with small, widely separated lobes. Accessory sac of ductus bursae with spiral thickening. Larva makes stigmatonome ..... stigmaciella (p.38) 30 MEM.TRS oF THE ENToMoLocrcAL socIETy oF .ANADA

The cnnrnecrFoLrELLA GRoUP Particularly characterized by the numerous denticulate cornuti which comprise a large patch on the vesica of the male. The lobes of the saccus are usually pointed, the uncus is bilobed and the gnathos more or less M-shaped. In the female the ductus bursae is long and the bursa copulatrix membranous, very delicate, usually only weakly pectinate and without a signum. The anterior apophyses are divided at the base giving rise to a small splinter-like process. The species feed on certain members of the Roseaceae and include the most brightly shining members of the family.

Stigmella crataegfoliel/a (Clemens) comb. n.

Nepticula crdtaegifoliel/a Clemens, 1861: 83 Nepticula crataegifuliella Clemens; Dyar, 1903: 546 (listed) Nepticula crataegifuliella Clemens; Braun, l9l4: l7 Nepticula crataegifoliella Clemens; Braun, l9llb: 193 Nepticula crataegfoliella Clemens; Bratn in Forbes, 1923: 94

DE,scrIprIoN. Externalfeatures: 6 ,?. Head: palps grey with grey lustre; antennae grey to ochreous; eye-caps white; tuft on front of head and vertex yellow-ochreous; collar as tuft. Thorax and abdomen: dorsal surface ochreous-orange, laterally and ventro-laterally grey. Wings: forewing with proximal third yellow-ochreous, medial third buff, distal third dark greyish brown; fringe grey. Legs: greyish to off-white, lustrous. Pl. 8.4. Wing measurements: 6.3.6-3.8 mm (2); ?.3.2-3.4 mm (2); neotype.3.6 mm. Genitalia: d. Figure 14; vinculum with ventral plate widely separating lateral arms giving capsule a rather short appearance; tegumen wider than long, rounded posteriorly; saccus comprising pair of well separated, pointed papillae; uncus with pair of small, blunt, well separated lobes; gnathos with a pair of lateral lobes and a pair of lateral, posteriorly pointing processes; valves with pointed style and rather rectangular cuiller; transtilla with horizontal bar wide; aedeagus with large patch of denticulate cornuti on vesica. ?. Figure l5; anal papillae as long as broad; ductus bursae long, with unadorned accessory sac; bursa copulatrix membranous with minute pectinations sparsely distributed; anterior apophyses relatively short, narrow at tip, wider at base; posterior apophyses also short, approximately same width throughout length. H o st- p lant : C r atae gus (Hawthorn). Mine: upper surface ophionome.

Dtecuosls. The external features are very easily distinguished by the extensive ochreous scales of the forewings. The form of the gnathos, tegumen and valves in the male genitalia separate crataegifoliella from pomivorella and other members of the group except scintillans. The broad ophionome separates crataegfoliella from the other Crataegus feeders, scintillans and stigmaciella.

DrscussroN. As with several of his other species Clemens failed to preserve the mines and larvae from which he described this species. The adult remained unknown until Braun (1914: 17) described it from specimens she reared from Crataegus. Consequently we designate one of these specimens as neotype. It is a male and bears the following label data: "Cincinnati O., Annette F. Braun, i.viii.14. lJ., on Crataegus mollis".

DrsrntsurroN. Canada: Ontario. U.S.A.: Ohio (type locality), Pennsylvania. SpEcIlrENs ExalrrNr,o. d neotype; in USNM: U.S.A.: Ohio, Cincinnati; on Crataegus mollis; l4.viii. l9l1 (Braun); genitalia, USNM slide No.16266. wlLKINSON AND SCOBLE: THE NEPTICULTDAE OF CANADA

FIc. 14. Stigmella crataegifoliella, d genitalia.

Other specimens: U.S.A.: 19, data as neotype; 3l.viii. 1917. Ohio, Clermont Co.; on Crataegus;20.v.1917 (Braun); in USNM. Pennsylvania, West Chester; onCrataegus; I d, 8.ii.1966 (Lewis); in CNC. CANADA: Ontario, Simcoe; on Crataegus; I d, I ?, l8.ii., 24.vi.1960 (Freeman and Lewis) in CNC.

Btor-ocv. Egg. Usually laid near the edge of the leaf. Mine. The mine follows theedgeof theleaf foraconsiderablebutvariablepartof itslength. Intheearlypartthe frass is dark and central, filling most of the channel. Later the mine widens rapidly and the frass line becomes rather irregular. Occasionally loops are formed indicating that the larva abandons its initially strictly linear feeding habit to consume tissue from a somewhat wider area. Pl. 2.1.

Specimen d There is a female specimen with genitalia indistinguishable from crataegifuliella, but rather different externals. DnscruprtoN. External features: ?. Head: palps white, lustrous; antennae whitish; eye-caps shining white; tuft on front of head and vertex off-white; collar white, Thorax white. Wings: ground colour of forewings white, lustrous, with one broad terminal to subterminal fascia with bronze reflections; fringe grey. Legs: grey with irregular off-white patches, lustrous. Wing measurement: 4.8 mm. SpEcIlrew ExnlrrNsp. CANADA: British Columbia. Summerland; 9, 2l.vi. 1935 (Gartrell): in CNC. J2 MEMOIRS OF THE ENTOMOLOGICAL SOCIETY OF CANADA

-r .lI ,./ i -\

Z\ _;

FIc. 15. StigmeLla trataegifoliella, ? genitalia W]LKINSON AND SCOBLE: THE NEPIICULIDAE OF CANADA 33 Stigmella scintillans (Braun) comb. n. Nepticula scintillans Braun, l9l7b: 161 Nepticula scintillans Braun; Braun ire Forbes, 1923: 86

DsscnIpttoN. External features'. 6 , ? . Head: palps dark grey and lustrous; antennae grey and lustrous, basally white; eye-caps white, shining white; tuft on front of head very dark chocolate brown and greyish brown; tuft on vertex very dark chocolate brown; collar dark chocolate brown. Thorax and abdomen brownish or purplish black with gold, purple and silver reflections. Wings: ground colour of forewings dark purplish brown, shining; one broad, strongly shining, silver, medial fascia; a basal patch and terminal band defined bf brilliant silver reflections; fringe grey. Legs grey, paling to off-white at tarsi, lustrous. Pl. 8.5. Wing measurements: 9 . 3.2-3.4 mm (4); holotype. 3.4 mm. Genitalia'. d, ?. As crataegfoliella. Host-plant: Crataegus mollis (Haw apple). Mine: upper surface ophionome, much contorted.

Dra,cNosrs. Differs from all the other species of the group in having a dark tuft. The brilliant reflections separate it from crataegfoliella, pomivorella, and chalybeia and the ground colour of the forewings differs from the two brightly shining new species of the group. Diagnostic features of the genitalia correspond to those of crataegifuliella from which they are indistinguishable. The nature of frass deposition in the mine is characteristic in the few specimens studied.

DrscussroN. Although the material available is very limited, the genitalia of this species do not seem to differ from crataegfoliella. The external colouration of both species is very different and since they feed onCrataegus in the same locality they are retained as full species.

DrsrnrsurroN. Canada: Ontario. U.S.A.: Ohio (type locality), Michigan.

SpncrltrNs ExalrrNEn. d holotype; in Phila: U.S.A.: Ohio, Cincinnati; on Crataegus mollis , B.448; 9.v .1914 (Braun); genitalia, USNM slide No. 17369. Paratypes: data as holotype; 16 ,29 I , 5,9.v.1914. Other specimens: CANADA: Ontario, Ottawa; on Hawthorn; I ?, 15.vi.1956 (Lewis), l?, 14.vi.1956 (Freeman); in CNC. U.S.A.: Michigan, Emmet Co., Sturgeon Bay, I d, 16.v. 1943'82006' (Braun\.

Bror-ocv. Egg. Upper surface of leaf. Mine. In early stages naffow with a compact central frass-line. Later widening with frass alternately deposited as arcs reverting to a more compact frass-line. In the later stages the mine sometimes crosses under the early part. Pl. 2.2.

Stigmella pomivorella (Packard) comb. n. Microptentx pomivorella Packard, 1870l. 237 Micropteryx pomivorella Packard; Chambers in Hayden, 1878: 157 (listed) Micropteryx pomivorella Packard; Walsingham, 1880: 83 Nepticula pomivorella (Packard) Busck, l90l: 52 Nepticula pomivorella (Packard); Dyar, 1903: 546 (listed) Nepticula pomivorella (Packard); Braun, l9l7b: l9O Nepticula pomivorella (Packard); Braun in Forbes, 1923: 93 34 MEM'TRS oF THE ENToMoLocICAL soclB'y ots cANADA Dnscruprton. External featt$es: d, ?. Head: palps whitish; antennae purplish grey; eye-caps white, shining; tuft on front ofhead and vertex orange-ochreous; collar buff. Thorax and abdomen purplish grey-brown. Wings: forewing ground colour as thorax and abdomen, whole surface reflecting bronze with some purple. Legs grey, paling towards tarsi, lustrous. Pl. 8.6. Wing measurements: 6 . 4.4-4.8 mm (3); ?. 4.6-5.0 mm (4). Genitalia'. d. Figure l6; as descriptionof crataegfoliella except that the capsule is rather longer and the aedeagus variable in width (see discussion). ? . As craraegifoliella.

H o s t- p lant s : M alu s ( Apple), C r at ae g us (Hawthorn) . Mine: an upper surface ophionome widening latterly.

DtacNosrs. Larger than chalybeia, lacking the swollen ends of the lateral arms of the vinculum in the latter and feedine on a different host. Differs from the other members of the species-group in having grey-brown forewings tinged with purple. It also differs from scinanella sp . n. in the form of the uncus. It may be distinguished from stigmaciella sp. n. in the form of the gnathos in the male and in lacking the spiral marking on the accessory sac of the ductus bursae in the female. It also shows some similarities to intermedia but differs in details of the uncus, the valves, the aedeagus, and the cornuti. In addition the fascia on the forewins of intermedia is absent in pomivorella.

DIscusstoN. Packard failed to designate any type-material. However, Busck (1901) states that he examined Packard's "type" in the Agassis Museum, Cambridge, Mass. Busck states that this was in "rather poor condition". We have not been able to locate this specimen, but Packard's description together with the work of subsequent authors seems to put the identity of the specimens we have examined beyond reasonable doubt.

d

b Frc. 16 Stigmella pomivorella, d genitalia WILKINSON AND SCOBLE: THE NEFTICULIDAE OF CANADA 35

The variable width of the aedeagus is shown in Fig. l6c and d. This difference in width may be correlated with distribution, i.e. in eastern localities the males of pomivorella seem to have a narrower aedeagus than those from the west. As insufficient material is available to be sure that this is a consistent difference, no separate taxonomic status is given to these variants. Three female specimens from British Columbia which feed on Crataegus rather than Malus have been examined. On the basis of external features and female genitalia they are indistinguishable from other specimens of pomivorella. They are the only available examples of this species reared from Crataegus. Unfortunately we have been unable to locate the mines and they have probably not been preserved.

DrstntnuttoN. Canada: Ontario, Nova Scotia, British Columbia. U.S.A.: New York, Washington, (Massachusetts, Packard).

Spsctl{rNs ExaurNnn. CANADA: British Columbia, Vancouver, Stanley Park; onCrataegus 3 ? I , l8.vi.l95l (McLeod); in CNC. Vancouver Is., Victoria; 1d, 5 ex, ll.vi.1923 (Blctckmore); in U.B.C. Nova Scotia, Round Hill, on apple; 1d, 119 ?, 18.iv.1911. Ontario, Niagara Falls; 2d d, 3,10.v.1900. U.S.A.: Washington, Olym- pia;on apple; No. 6112, 106 6, 129 9, 2 ex, 2l-25.iv.1894. New York, Castile; 19, 1 .vi.1925 (Bradley). Washington, Bellingham; on apple; I ex, v. 1918 (Newc:omer). No locality given; on apple; 19, I ex, iv.\922. 1d, (Fernald Coll.); I ex,9.vi.1963 (Fernald) [Cu Exp. No. 721] probably belongs here; in USNM. Brolocv. Mine. Braun (l9l1b) figures the mine of pomivorella and there appears to be no difference between this and the mine made by scinanella sp. n., the details of which are given below. We have been unable to examine any mines from which specimens of pomivorella have been reared.

Stigmella chalybeia (Braun) comb. n. Neptir:ula chalybeia Braun, l9l4: 20

DnscntpuoN. External .f'eatures: 6, 9. As po,nivorella except for lack of purple reflections in chalybeia and perhaps the smaller size of the latter. Wing measurements: 3. 3.8.-4.0 mm (3); lectotype. 3.2 mm. Genitalia'. d. Figure 11; aspomivorella except vinculum rather rectangular in shape with posterior ends of lateral arms terminating in swollen knobs; aedeagus long with denticulate comuti and a rectangular striate patch on vesica. ? . As t'rataegiJoliella. H o st- p lant : Pyrtrs (Pear). Mine: Upper surtace ophionome, as pomivorella.

DracNosrs. The presence of swellings on the lateral arms of the vinculum separates this species from the other members of the crataegfoliella group. In addition, the purplish grey-brown forewings of chalybeia distinguishes it from all of them except for pomivorella. S. chalybeia has a smaller wingspan than pomivorella. DtsrnrsurroN. Canada: Ontario. U.S.A.: Ohio (type locality). SpEcrrunNs ExaurNEo. We designate as lectoype the male of Braun's two syntypes. It bears the following data: "Cincinnati, O.B. 751 (on wild pear), i. [imagine emergedl 22.vri.l913 Annette F. Braun; CoTYPE, Collection of Annette F. Braun"; genitalia, USNM slide No. 16219; in Phila. Paralectotype: 1?, same data as lectotype; 23.vi|.1913. (Abdomen missing). 36 MEMOIRS OF THE ENTOMOLOCICAL SOCIETY OF CANADA

.s i^:4ts lrf_^ l\€ ,{(,/ ^'.--*d.

b Frc. 17. Srigmella chalybeiu. d genitalia.

Other specimens: CANADA: Ontario, Vineland Sta., on pear; I 6 6 , 1.934 (Walley); in CNC.

Bror-ocv. This species appears to feed only on pear, even where pear and apple grow in the same area (Braun 1911b). Unfortunately, Braun does not indicate whether the leaves of apple trees in such a situation are mined by pomivorella which would give evidence that sympatric populations of the two species do not interbreed.

Stigmella scinanella sp. n. DnscntprtoN. External features: 6 , ?. Head: palps grey; antennae dark greyish brown; eye-caps white, shining; tuft on front of head and vertex orange-ochreous; collar pale ochreous. Thorax and abdomen brownish black. Wings: ground colour of forewings blackish brown with striking iridescence but particularly red-gold and basally to sub-basally with brilliant silver or gold, depending on angle of illumination; postmedial fascia defined by brilliant silver reflections. Legs blackish, paling to off-white at tarsi, lustrous. Pl. 8.7. d.4.5mm(1); ?.accuratemeasurementsnotpossible; holotype.4.4 .rrntmeasurements: ^ Genitalia: d. Figure l8; uncus short, wide and with pair of large, ear-like lobes; gnathos bilobed with pair of lateral arms; valves reaching uncus, weakly divided into style and rectangular-shaped cuiller; transtilla stout; aedeagus long, narrow posteriorly, with large patch of denticulate cornuti on vesrca. ? . As crataegifoliella. Host-plant: Malus ( Apple). Mine: trpper surface ophionome, beginning narrow and then broadening quite rapidly about half way along length. WTLKINSON AND SCOBLE: THE NEFTICULIDAE OF CANADA )t

Frc. 18. Stigmella scinanella, d genitalia.

DrncNosts. This species has striking ref-lections. a postmedial fascia and darker legs which pomivoreLla lacks. In the male senitalia the uncus has large. ear-like lobes which are not widely separated. This feature, and the absence of the spiral marking on the ductus bursae, distinguishes it from both pomivorella and stigmaciella sp. n. The mine of the next species is a blotch on Hawthorn whereas scinanella makes a linear tract in the leaves of apple. The orange-ochreous tuft, forewing with postmedial fascia and large, shining silver, basal patch together with the golden-red reflections of scinanella distinguish it from scintillans.

DtsrnrsurtoN. Canada: Ontario.

Spnctlrr,Ns ExavtNpo. d holotype; in CNC: CANADA: Ontario, Normandale; on Malus, 56-154 20.1ri.1951 (Freeman and Lewis); genitalia CNC slide No. 2972. Paratypes: data as holotype; I ?, I ex, 8. iv.1958. Ottawa; I ?, 12.vi.1956 (Freeman) in CNC.

Ervlrolocy. Scinan (O.E.)-Shine. (Little Shiner.) 3 8 &MOTRS ots r HE LN l oMoloclcAI- socrETy oF cANADA Brolocy. Egg. Laid on upper surface of leaf. Mine. The frass-line in the early part of the tract is compact, central and occasionally broken. Later it becomes dispersed as separate grains. The mine broadens suddenly about half way through its length, being similar to crataegifoliella. However, in the specimens examined the larvae do not generally follow the edge of the leaf in the early stages as in the latter. Pl. 2.3.

Stigmella stigmat:iella sp. n. DlscntprtoN. External features: 6, 9. Indistinguishable from scinanelLa. Wing measurements'. 6. 5.0 mm (l); ?. 5.4 mm (1); holotype. 5.0 mm. Genitalia'. d. Figure 19; tegumen a wide band gently rounded posteriorly; saccus bilobed, each lobe rounded; uncus short and broad with a pair of small, widely separated papillae; gnathos wide, with a pair of large, laterally placed spines and a pair of short posteriorly directed spines; valves clearly divided into rectangular cuiller and pointed style; transtilla fairly well sclerotized, broad; aedeagus short and broad with very large arcuate, densely spinose cornutus and with some larger spines laterally. ?. Figure 20; ductus bursae approximately twice as long as apophyses, with an accessory sac, the wall of which is supported 6y a spiral striation; bursa copulatrix large, with a few short denticulate chain-like pectinations; signum absent; anterior apophyses short, broad at base; posterior apophyses also short, slightly flattened before entering body membrane, and continuing as a fine sinuous apodeme. H o s t- p lant : C r atae gus (Hawthorn). Mine'. tpper surface orthogenous stigmatonome.

DrecNosrs. The shining external features distinguish this species from the others in the crataegfotietla group except for scinanella and scintillans. S . stigmaciella has a bright ochreous tuft, a postmedial fascia and forewings with striking reflections, whereas scintillans has a chocolate-brown tuft, a medial fascia and weaker reflections. The lobes of the uncus of stigmaciella are very small and widely separated compared with those of scinanella which are large and less widely separated. This is the only species of the crataegifoliella group whose larva forms a blotch mlne.

DrscussroN. In external features this species shows remarkable convergence with scinanella, however, its genitalia are sufficiently different, to warrant full, specific status. The mine is very characteristic.

DrsrntnutroN. Canada: British Columbia.

SpEcrvrnNs ExenrNlo. d holotype; in CNC: CANADA: British Columbia, Vernon; on Crataegus; 24.vi.1957 (Freeman and Lewis); genitalia, CNC slide No. 3316. Paratypes: data as holotype; I d, I ?, I ex, 25,21.vi.1957; in CNC.

Etvnor-ocv. Stigma (Gk. )-Spot. Macian (O.E. )-Maker. (Little Spot-Maker. )

Brolocy. Egg. Laid on the lower surface of the leaf. Mine. The mine is made at the edge of the leaf and the frass comprises a central patch. Pl. 2.4.

Krv: the sAGTNEI-LA GRoUP l. Forewingswithcreamy-yellow scaling ...... ' 2 - Forewings grey-brown castaneaeJbliella (p 14) 2. Forewings with widc ochrcous fascia. Male with gnathos as in Fig. 25 . . . latifasciellu (p. 47) - *:*':*:*:i-r':l'"*l]'":i* *':n o::*" *":*l*::i:1"::: ':.0':,;;" th (p 3e) WILKINSON AND SCOBLE: THE NEPTICULIDAE OF CANADA 39

I :mm

Frc. 19. Stigmella stigmatiella, d genitalia.

The slcrNert-A GRoUP The male genitalia are particularly characterized by the bilobed uncus and the M-shaped gnathos. The valves are rather narrow with the cuiller and style weakly defined. The aedeagus has a striate plate on the vesica and, when present, the spine-like cornuti are few in number: there is a leuge juxta. The lateral arms of the vinculum are short and broad, giving the genitalic capsule a rather broad, squat appearance in ventral view. In the female the accessory sac of the ductus bursae is adorned with denticles and the signum on the bursa copulatrix is in the form of a U-shaped band or a pair of straight bands. The larva leaves the mine from the lower surface of the leaf and the adults all have at least some purplish brown scales on the forewings.

Stigmella saginella (Clemens) comb. n. Nepticula saginella Clemens, 1861: 85 Nepticula saginellct Clemens; Clemens, 1865: 146 Nepticula saginella Clemens; Chambers ln Hayden, 1878: 158 (listed) Nepticula saginella Clemens; Dyar, 1903: 547 (listed) Neptit:ula saginella Clemens; Busck, 1903: 220 (discovery of Clemens' "type") Nepticula saginella Clemens; Braun, 1911b: 195 Nepticula saginella Clemens; Braun in Forbes, '1923: 95 40 MEMOIRS OI- THE ENTOMOLOGICAL SOCIETY OF CANADA

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Ftc.20. Stipmella stismacieLla. ? eenitalia W]LKINSON AND SCOBLE: THE NEPTICULIDAE OF CANADA 41 Neptic:ula quercicastanella Chambers, 1873: 127 (syn. by Braun, 19llb: 195) Nepticula quercicastanella Chambers; Chambers ln Hayden, 1878: 158 (listed) Nepticula quercicastanella Chambers; Chambers, 1880b: 66 Nepticula querc:icastanella Chambers; Dyar, 1903: 517 (listed) Nepticula fuscocapitella Chambers, 1813: 128 (syn. by Braun, 1917b: 195) Nepticula fuscocapitella Chambers; Chambers ln Hayden, 1878: 157 (listed) Nepticula fuscocapitella Chambers; Dyar, 1903: 545 (listed)

DrscntprtoN. External .features: 6 , ?. Head: palps: labials dark grey-brown, maxillaries whitish; antennae dark grey; eye-caps dull white; tuft on front of head and vertex brownish black; collar dull white. Thorax and abdomen slate grey dorsally. Wings: ground colour of forewings creamy yellow, variously irrorate with brown scales which reflect purple. Legs grey, with ochreous patches, paling towards tarsi, lustrous. Pl. 9.1. Wing measurements: 6. 4.0-5.0 mm (7); ?. ,1.0-4.8 mm (7). Genitalia: d. Figure 2l; vinculum with medial posterior edge of ventral plate weakiy concave; tegumen in form of narrow band; saccus weakly bilobed; uncus bilobed with lobes widely separated; gnathos M-shaped with lobes widely separated; valves relatively narrow with cuiller poorly demarcated from pointed style-fiom latter arise a number of long straight scales, each terminating in flattened, digitate head; transtilla with short lateral arms each terminating in triangular plate, the apices of which just fail to join; aedeagus with striate arcuate plate on vesica. ? . Figure 22; as castaneaefoliella with the following exceptions: anal plate wider, lobes rounded; ductus bursae with accessory sac covered with small denticles; bursa copulatrix covered with scale-like pectinations; signum U-shaped, comprising band of spinose ridges. Host-pLants: Quertus prinus (Chestnut Oak), Q. alba (White Oak), Q. platanoides, Q. macrocarDa rBur Oakt. Mine: an upper surtace ophionome: larva emerges from lower surface of leaf to pupate.

Dr.c.cNosrs. Diffbrs from latifasc:ieLLa in that the latter has the creamy white scales of the forewing in the form of a wide fascia. Compared with both latifosciella and

a

Frc. 21. Stigmella saginella, d genitalia A '',) MEMOIRS OF THE ENTOMOLOCICAL SOCIETY OF CANADA

'f \.. / .....\ Y:t ; i --J'\-"1 - i-

Frc. 22. Stigmella saginella, ? genitalia. WILKINSON AND SCOBLE: THE NEPTICULIDAE OF CANADA 43 castaneaefoliella, in saginella the lobes of the uncus and arms of the gnathos are more widely separated and the valves have long, terminally digitate scales arising from the style. ln the female of saginella the markings on the accessory sac are slightly larger thanlatifusciella and the signum on the bursa is in the form of a U-shaped band rather than two separate elements.

DrscussIoN. This species was first described by Clemens from mines he collected on both Oak and Chestnut. As both saginella and castaneaefoliella make similar mines and also because adult specimens of the latter (as here diagnosed) have not been found on chestnut, saginella is regarded as an Oak but not a Chestnut feeder. Clemens eventually reared an adult specimen of saginella which he labelled "type", but not until 1865 which was after the original description had been made. We have examined this specimen but unfortunately the abdomen is missing so the genitalia cannot be checked. The specimen cannot be the holotype as it was reared after the species was first described from the mine and it is here designated a neotype, the original mine being no longer extant. It is lodged in the Academy of Natural Sciences, Philadelphia and bears the following label data: "Clemens No. 420, Type 1496, Nepticula saginella B. Clemens". A second label adds "A.B. 1902" to the name.

DtstnInurroN. Canada: Ontario, Quebec. U.S.A.: Ohio, New York; Virginia.

Spncrl.tENs ExAN,IrNEo. ? neotype labelled as 'Type' by Clemens; in Phila; (Clemens); Type 7496. Other specimens: CANADA: Ontario, La Passe; on Quercus alba;19, 5.ii.197 I (Lewis). Kinburn; on Quercus alba; | ? , 2l .viii. 1959 (Freeman and Lewis). Quebec, Val Tetreau; on Quercus alba; 19, 25.i.1961 (Freeman); in CNC. USA: Ohio, Cincinnati; onQuercus prinus; I6,26.v.1911 ; I ex, 9.viii.l9l3 (Braun);onQ. alba; ld,14.vii.l9O9(Braun);onQ.platanoides;466,39?,1,4.viii.1913,5-30.v.1914 (Braun); on Q. macrocarpa; 18.ix.1913 (Braun); on Quercus; I ex, 2.viii.1914, l6 , l?, 3.v.13 (Braun)t Adams Co., Peach Mountain; on Q. prinus; 1d, 19, l6,l8.vii. 1928 (Braun); in Phila. Virginia, Falls Church; on Oak; I d , 1 ex, 24.vi.1913 (Heinrich). New York, Ithaca; at light; I d , I I , 3. vi. , 24.vrii.1924; on Oak (no locality data); 2? ?, I ex, 1886, (ex Murfeldt Coll.); on Oak; 1d, 16.iv.1885; ld, I ?, 4.viii.1890; in USNM.

Brolocv. Egg. Laid on upper surface. Larva. Emerges from lower surface of leaf , at time of pupation. Mine. Appears rather wider than mines of castaneaefoliella but it is unlikely that this is a diagnostic feature. Pl. 3.1. Voltinism. In U.S.A' trivoltine; at least bivoltine in Canada.

Specimen e There is a female specimen probably incorrectly recorded from 'Scarlet Oak, Quercus coc:c'inea', which resembles saginella but differs in the following ways: forewings pale creamy yellow, barely irrorate with brown scales. Genitalia with apophyses long and sinuous. On the basis of available evidence the specimen is regarded as an aberation or variety of sagineLla.

SpncttttEN ExeltINnn. CANADA: Ontario. Simcoe; 9 , 21 .iii.l968 (Freeman); in CNC. 44 ,,,, ),,':;,::,;,': : . i,i "'i ; Jff ;;, ". Nepticula castaneaeJbliella Chambers, 1875: "I l7 Neptir:ula castaneaefolieLLa Chambers; Chambers in Hayden, 1878: 157 (listed) Nepticula castaneaefolieLla Chambers; Chambers, 1880a: 193 Nepticula c:astaneaefoliella Chambers; Dyar, 1903 545 (listed) Nepticula castaneaefoliella Chambers, Braun, l9l7b: 192 (partim) Nepticula castaneaefolielltt Chambers; Braun ln Forbes, 1923: 94

DsscntpttoN. External .features'. d, ?. Head: palps, maxillaries grey, lustrous, labials brownish; antennae grey and lustrous; eye-caps white, shining white; tuft on front of head and vertex dark brown; collar white with white reflections. Thorax and abdomen greyish brown with grey lustre, and some purple and bronze reflections. Wings: ground colour of forewings grey-brown with some golden and purple reflections, dusted with grey; fringe grey with purplish tinge. Legs grey with irregular buff patches, lustrous. Pl. 9.2. Wing meastrrements: 6. 4.0-4.8 mm (9); ?. 3.8-4.4 mm (15). Genitalia: d. Figure 23; vinculum with medial posterior edge of ventral plate with slight convex bulge; tegumen wide, bluntly rounded; saccus weakly bilobed; uncus comprising pair of long papillate lobes; gnathos M-shaped comprising pair of posteriorly pointing widely separate horns and pair of laterally directed spines; valves with weakly differentiated cuiller and pointed style from which arise a number of scales each comprising a long shaft, reaching approximately one-third of length of valve, not terminating in digitate lamina; transtilla stout, lateral arms short, straight each terminating in large triangular-shaped plate the apices of which just fail to meet, thus forming incomplete horizontal bar; aedeagus with striate, arcuate plate on vesica and patch of' shtrrt. stout spine-like cornuti.

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a

FIc. 23. Stigmella castuneae.foliella, d genitalia WLKINSON AND SCOBLE: THE NEPTICULIDAE OF CANADA 45

Ftc. 24. Stigmella castaneae.foliella, ? genitalia 46 MLM'IRS ots rHE tsNTOMoLocrcAL socrETy oF cANADA

? . Figure 24; anal plate with pair of triangular lobes; ductus bursae approximately same length as apophyses, widening gradually to bursa copulatrix, accessory sac adorned with denticles; bursa copulatrix large, with weak chain-like denticulate marks in area where it merges with ductus, otherwise covered by minute pectinations and with U-shaped signum formed by tncreased density and size ofthese marks; anterior apophyses broad basally; posterior apophyses long. straight and narrow. Host-plant: Castonea dentata (Chestnut). Mine: tpper surface, narrow ophionome. Frass-line narrow relative to width of mine, not reliably distinguishable frcm saginella.

DrlcNosrs. Differs from the closely related latfosciella and saginella in lacking the creamy yellow on the forewings, thorax, abdomen and patagia. The male genitalia of castaneaefoliella differ in the lobing of the valves, in the form of the gnathos, which is less heavily sclerotized than in latifasciella and the arms of which are more closely .;uxtaposed than in saginella, and in the ventral plate of the vinculum. The nature of the scales arising from the style of the valves, and the presence of spine-like cornuti on the vesica of the aedeagus are also diagnostic. The female may be distinguished from both saginella and latifasciella in the denticulate markings on the accessory sac of the ductus; the signum on the bursa is pectinate rather than ridged. Further, in latifasciella the ridged markings are represented by two bands which are not continuous and so do not form the U-shaped signum of castaneaefoliella.

DrscussroN. Braun reared several specimens from Chestnut and Oak but although she identified all these as castaneaefoliella, the oak-feeders are of doubtful identity as the female genitalia are indistinguishable from that of saginella. These specimens are treated separately, see below, and are not regarded as being castaneaefoliellct.

DrstntsurroN. Canada: Ontario. U.S.A.: Kentucky (type locality), Ohio, Virginia, Pennsylvania, New Jersey.

Spscrr{ENs Exe}rrwen. I holotype; in MCZ. U.S.A.: Kentucky (Chambers) Type No. 14956; genitalia, CNC slide No. 3492. Other specimens: CANADA: Ontario, Simcoe; on Castanea dentata;36 6, 29?, 4-'7.viii.1956, 3-4.iii.1958 (Freeman and Lewis). Normandale; on Castanea dentata; 19, 19.vii.1956 (Freeman andLewis). St. Williams, onCastanea dentata;49 ?, I ex, 4.viii.l956 (Freeman and Lewis); in CNC. Lake Erie, St. Williams; on Castanea; 26 6 ,l?,21.1i.1962;36 6 ,19I , I ex,2-20.iii.1962; in Soo. U.S.A.: Ohio, Adams Co., Beech Fork; on Castanea; I d , 1 I , 15,16.vii. 1928 (Braun). Cincinnati; on Castanea;19, l7.vii.19l3; l?,24.viii.1911, I ex, 9.v.1912 (Braun). Kentucky, Powell Co.; on Castctnea;1d, 1.viii. 1916 (Braun). Pennsylvania; 1 ex, 21 .vi.19l'7 (Haimbach);inPhila.Virginia,FallsChurch',onCastaneadentata;366,499,1ex, 2l-26.vi.1913;' 25.v|i.-9.viii. l9l3 (Heinrlclz); New Jersey, Montclair, 1 9, 17.vii (Kearfox); in USNM.

Brolocv. Egg. Laid on upper surface of leaf. Mine. The frass-line is usually dark and broken but in one specimen from the CNC and one of Braun's mines the frass is deposited in zig-zag arcs. Pls . 3 .2, 3 .4. Larva. Emerges from lower surface of leaf, at time of pupation. Cocoon. Buff. Voltinism. In Canada the material available (all collected in Ontario) suggests that the species is bivoltine. In the U.S.A. adults have been reared in May, June to July, and August, suggesting three generations per year. WILXINSON AND SCOBLE: THE NEPTICULIDAE OF CANADA 47 Stigmella latifasc:iella (Chambers) comb. n.

Nepticula latifasciella Chambers in Hayden, 1878: 106 Nepticula latifusciella Chambers; Dyar, 1903: 546 (listed) Nepticula latifasciella Chambers; Braun, l914: l8 Nepticula latfasciella Chambers; Braun, l9llb: 193 Nepticula latifusciella Chambers; Braun in Forbes, 1923: 94 Nepticula macrocarpae Freeman, 1967: 19 syn. n. Nepticula macrocarpae Freeman; Lindquist and Harnden, 1970: 1290 (biology)

DnscntprtoN. External features'. d, 9. Head: palps dull white; antennae dark greyish brown dorsally, lighter ventrally, and with grey lustre; eye-caps off-white and shining white; tuft on front of head pale ochreous; tuft on vertex dark brown; collar off-white. Thorax and abdomen greyish brown with grey lustre and some purple reflections. Wings: forewings sub-basally to postmedially, and at tip, creamy yellow with grey and pale golden reflections, basally to sub-basally and postmedially to terminally dark greyish brown with purple and golden reflections, fringe grey. Legs grey with buff patches, lustrous. Pl. 9.3. Wing measurements'. 6. 4.0-5.0 mm (l l); I . 4.0-5.2 mm (28). Genitalia: d. Figure 25; vinculum with medial posterior edge of ventral plate with slight convex bulge; tegumen wide, bluntly rounded posteriorly; saccus weakly bilobed; uncus robust, deeply bilobed; gnathos M-shaped with lobes close together and rather elongate, very heavily sclerotized; valves relatively narrow, with style and rounded cuiller; transtilla stout, with short lateral arms each terminating in a triangular plate, the apices of which are elongate and just fail to join, so forming long, weak horizontal bar; aedeagus straight, with arcuate, striate patch on veslca. ? . Figure 26; anal plate bilobed, ductus bursae approximately length of anterior apophyses; bursa copulatrix large, covered with small rows of minute pectinate marks; signum comprising

a

Frc. 25. Stigmella latifusc'iella, d genitalia 48 MEMOIRS OF THE ENTOMOI,OGICAL SOCIETY OF CANADA

Frc. 26. Stigmella latifasciella, ? genitalia THE NEPTICULIDAE OF CANADA 49 two bands of spinose ridges which fail to join, thus not forming U-shaped band; anterior apophyses shorter than posterior apophyses; posterior apophyses very long, and narrow. Host-plant'.Querr:usrubra(RedOak),0. palustris,Q.macrocarpa(BurOak),minesfound onQ. alba (White Oak) by Lindquist and Harnden (1970). Chambers (1878: 106) states that he collected the holotype from the trunk of Castanea americana (now : dentata Borkh.). Mine: upper surface ophionome.

Dracxosrs. ln latifasciella the creamy yellow scales of the forewing are concentrated to form a wide fascia whereas they comprise the whole forewing ground colour insaginella and are absent incastaneaefoliella. The shape ofthe uncus, gnathos and valves are diagnostic (compare fig.25a with 21a and23a). The markings on the accessory sac of the ductus in the female of latifasciella are smaller than in saginella and particularly so compared with castaneafoliella. The mine of latifasciella has a wider frass line relative to the mine width than the other members of the species group. S . latfosciella resembles alba sp . n . in the creamy yellow markings of the forewing, but the dark basal patch together with the brown tuft on the vertex of latifosciella distinguish it externally, and the genitalia are quite different.

DrscussroN. Freeman (196'7 : 19) described what he thought to be a new species. viz: "Nepticula macrocarpae" . However, as the genitalia and externals of the type material of this species have been found to be similar to Chambers' type of latifasciella, macrocarpae becomes a junior synonym.

DrsrnrsurroN. Canada: Ontario, British Columbia. U.S.A.: Kentucky (type locality), Ohio, Michigan, Virginia, Arkansas.

SpecrN{ENs ExlurNsr. I holotype; inMCZ. U.S.A.: Kentucky; (Chambers); type No. 1497; genitalia CNC slide No. 3497. 6 holotype of macrocarpae, in CNC; CANADA: Ontario, Lk. Simcoe, Orillia; on Quercus macrocarpa; No. 9539. 565-4100-01. l2.viii. 1965. Other Specimens: CANADA: Data as holotype of macrocarpae;206 6 , 349 ? , 4 ex, 12-15.11i.1965 30.vii., 12.viii.l965; 12.iv.1965; 10-19.iii. 1966 9-l'7 .viii.l966. Ontario, Bells Corners; on Red Oak; 1?, 16.iv.1959 (Freeman and Lewis). British Columbia, Victoria; on Oak;29 9, 30.v.1962 17.v.196'7 (Evans); ld, 6.vi.1965 at light (Evans); in CNC. 399, 2 ex, 4,14,21.vi.1923 (Blackmore); 29 9, 1 ex, 8.vi.1922 (Carter); in UBC. U.S.A.: Michigan, Cheboygan Co., Douglash, Maple Bay; ld, 12.viii.1936. Ohio, Cincinnati, 69 9, I ex, 12.vi.l903, 29'iv.1905, 15,22.v.19O6,25.vii.1914,6.ix.1954 (Braun). Spring Grove; onQuercus rubra; l?, 18.ix.1913 (Braun). Brachmann's1'onQuercus palustris;1?,25.iii. 1917 (Braun);in Phila. Salem Pike; on Quercus rubra;29 I , 12.vni.19l3. Virginia, Falls Church; 19, vi.l903 (Busck); Arkansas, Washington Co., Devils Den State Park, 16, 29 I, 24.v.1966, 30.v.1966 and 23.vi.1966; in USNM.

Btolocv. Supplemented by the work of Lindquist and Harnden (1970). Egg. Usually laid on the upper surface, but occasionally on the lower surface of the leaf. Many eggs may occur on one leaf. Mine. Early part of mine nearly filled by dark frass-line; later the frass particles become more disseminated. Pl. 3.3. Larva. Bright green, head pale with a black spot on each side. Vacates mine from under surface of leaf and overwinters in this stage in a cocoon. Cocoon. Light grey, spun in ground litter. Pupa. At first pale green, later darkening. Voltinism. Bivoltine in southern Ontario. 50 MEMOIRS OF THE ENTOMOLOGICAL SOCIETY OF CANADA

Kny: the coRYLrFoLrELL.q. cnoupg l. Male with ventral plate of vinculum very large; gnathos with arms sigmoid, unbranched M;i; *i,tr r.rrour pi.,. r", r"rg"' gnuito, *irrr ,r.. t:r:::l:::'f.i'ilt;;:t';". - "i "i"."r"- z. v"r".r.i.ur. nurro*. rr"ri)""i^r 0., oiir""r,ili" uli"u.i .. ,:"ryiit)iiit ,o ,ol - Valves of male wide, horizontal bar of transtilla not bilobed ...... ostryaefoliella (p. 54)

The conylrpot-rEt-r-A GRoUP In the species of this group the aedeagus is small and flask-shaped with a pair of plates at the phallotreme. The uncus, gnathos, and general shape of the genital capsule are characteristic. The bursa copulatrix of the female is pectinate with a pair of oval signa which are not heavily sclerotized. The wings have a single silver fascia and bronze or purple reflections. The range of host-plants is extremely wide, the significance of this fact is discussed more fully in the general discussion (p. I la). DtscussroN. Clemens (1861) described corylifoliella, ostryaefoliella, vir- giniella, juglandifoliella, and caryaefoliella from mines which he apparently failed to preserve. Chambers, Braun, and Busck subsequently described additional species based on very small differences in the external features and slight variation in the mine patterns. These authors tended, as a matter of course, to describe new species in this group if they reared specimens from previously unrecorded food-plants. A study of the genitalia has led us to synonymize a number of species. The external features of the adults, and the shapes of the mines are of poor diagnostic value. This fact, together with Clemens' habit of describing new species from the mines alone, has resulted in considerable confusion. We have used Clemens' species names, although we cannot be absolutely certain exactly which species caused the mines on which he based his descriptions. However, in the interest of stability of nomenclature it does not seem justifiable to propose new names. Owing to this uncertainty no neotypes are designated.

Stigmella t:orylfoliella (Clemens) comb. n. Nepticula corylifolieLLa Clemens, 1861: 83 Nepticula corylfoliella Clemens; Chambers in Hayden, 1878: 157 (listed) Nepticula corylfoliella Clemens; Packard, 1889: 356 Nepticula corylifuliella Clemens; Dyar, 1903: 545 (listed) Nepticula corylifoliella Clemens; Bratn l9l2'. 92 Nepticula corylfoliella Clemens; Braun l9llb: 179 Nepticula corylifuliella Clemens; Braun ln Forbes, 1923: 89 Nepticula virginiella Clemens, 1861: 83 (syn. by Braun 1917b: 179) Nepticula virginiella Clemens; Chambers in Hayden, 1878: 158 (listed) Nepticula virginiella Clemens; Dyar, 1903: 547 (listed) Nepticula minimeLla Chambers, 1873: 121 (syn. by Braui I9llb: 179) Nepticula minimella Chambers; Chambers ln Hayden, 1878: 158 (listed) Nepticula minimella Chambers; Dyar, 1903: 546 (listed) Nepticula opulfoliella Braun, l9l4:22 syn. n.

3 N. myricafoliella is not included in this key because it is lit(ely to be conspecific with ostryaefoliella-see under the discussion of this soecies. W]LKINSON AND SCOBLE: THE NEPIICULIDAE OI,'CANADA 5 I Nepticula opulfoliella Braun; Braun 1917b: 180 Nepticula opulifoliella Braun; Braun in Forbes, 1923: 90 Nepticula paludicola Braun, 19l7b: 177 syn. n. Nepticula paludicola Braun; Braun in Forbes, 1923: 89 Nepticula exasperata Braun, 1930: 17 syn. n.

DEscnrpttoN. Externalfeatures: 6 . Head: palps grey and lustrous; antennae purplish grey, lustrous; eye-caps white, shining, generally with brownish purple shading particularly around edges; collar pale ochreous or grey-purple; tuft on front ofhead and vertex pale ochreous to bright orange-ochreous. Thorax and abdomen dark greyish purple. Wings: ground colour of forewings dark greyish brown to purplish black sometimes with bronze and sometimes with purple reflections predominating; one broad, white, generally postmedial, sometimes subterminal, fascia, shining silver; fringe grey, irrorate with brown. Legs purplish grey, lustrous, and pale ochreous patches particularly on forelegs, midlegs, hind tarsi and tibial spurs. ? as d except for: eye-caps always shining creamy white. Wing measurements: 6.3.2-4.2 mm (14); 9.3.6-4.2 mm (21). Genitalia: d. Figure 27; tegumen in form of narrow, serrate band, slightly arcuate; saccus bilobed; uncus rectangular, weakly bilobed; gnathos with short branch arising from each horn; valves narrow, extending beyond uncus, with thin hairs arising particularly from inner apical aspect; transtilla with lateral arms forming bilobed horizontal bar; aedeagus flask-shaped with pair of palmate, arcuate plates at phallotreme and pair of heavily sclerotized spine-like processes on vesica. ?. Figure 28; ductus bursae short; bursa copulatrix with striations and scale-like markings which become dense and denticulate to form pair of rather weakly demarcated signa; anterior apophyses stout, arcuate; posterior apophyses slightly longer, straight and narrow.

W

b

Ftc. 27. Stigmella corylfoliella, d genitalia. 52 MEMOIRS OF THE ENTOMOLOGICAL SOCIETY OF CANADA

Frc. 28. Stigmella corylifoliella, ? genitalia.

Host-plants: Corylus (Hazel), Betula alba, B. alleghaniensl.s (now : lutea) (Btch), Alnus (Alder), Hamamelis virginiana (Witch-Hazel), Vacr:inium, Gaylussacia baccata, Orycoccus palustris (Cranberry), Physot'arpus opulifolius (Opulaster). Mine: a narrow, upper surface ophionome.

DrecNosrs. Differs from juglandfoliella in the overall form of the male genital capsule. S . juglandifuliella has a very large ventral plate and the arms of the gnathos are resssigmoid,^,'rt,tut;;;:H"-*ff;#;;,:;;:byitsnarrow.,,'."r'": valves, in the serrate nature of the tegumen and in the more shallow lobing of the saccus. The lateral bars of the transtilla are extended in corylfoliel/a giving the horizontal bar a bilobed appearance.

DrscussroN. The adults were first made known by Braun (19t4). This author described specimens she had reared from mines, similar in form and on the same host-plant (viz. Corylus) as given in Clemens' original paper. Subsequently she described opulifoliella, paludicola and exasperata, all of which we have reduced to synonymy. The last two species were originally described only from single specimens. All three feed on different food-plants. Actually Braun (1930: 17) believed that exosperata was most closely related to rosaefoliella but the genitalia show this to be incorrect. The genitalia of paludic:ola appear to be indistinguishable from corylfoliella. The species was described from a single male specimen, which had the fascia on the forewing subterminal in position rather than postmedial. This specimen is considered to fall within the confines of the variation exhibited bv corylifoliella and the soecies is reduced to svnonvmv.

DrstnlnuttoN: Canada: Ontario, New Brunswick, Quebec, British Columbia. U.S.A.: Ohio, New Jersey, Kentucky, Michigan.

SpecrN.IsNs Ex.c.N{rNEn. Types: Syntypes of opuliJoliella; in Phila: U.S.A.: Ohio, Cincinnati; on Physocarpus opulfolius; 7I ?, 3 ex, 29.vii.-4.vlii.19l3 (Braun).6 holotype of paludicola; in USNM: U.S.A.: New Jersey, Whitesbog; on Oryc:or:cus palustris; l6.viii.l9l5 (Scammel/); genitalia slide No. CNC 3466. 6 holotype of exasperata; in Phila: U.S.A.: Ohio, Adams Co., Beaver Pond; on Vaccinium; 5.vILi.1927 (Braun); genitalia slide No. USNM 11244. Other specimens of corylifoliella: CANADA: Ontario, Simcoe; onCorylus americarut; 299, 10,23.ri.1960 (Freeman and Lewis). On Vaccinium;39 ?, 9,12.iii.1958 (Freeman andLewis). La Passe; onBetula alba; | 6 ,22.i1i.19'71 (Lewis). Normandale; on Hamamelis virginiana; 26 6, 3 ? 9 , I ex, 22.v.-7 .vi.1962 (Freeman and Lewis). OnGaylussaciabaccata;766,9??,1ex,2'7.v.-10.vi.1962(FreemanandLewis). Quebec, Fairy Lake; on Yellow Birch; I ? , 5.vi. 1956 (Lewis). OnAlnus;26 6 ,30.vi., 5.vii.1956 (Freeman); 19, 21.vi.1956 (Lewis). Kingsmere; on Betula alba; l?, 2'7 .vi.1956 (Freeman); in CNC. New Brunswick, Northumberland Co., Quarryville, on White Birch; 1d, f.iv.1956; in Dept. Fisheries and Forestry, Fiton, New Brunswick. British Columbia, Field; on Betula; labelled "Nepticula ostryaefoliella Clem.", 19, 15.v.1916 (Braun); in Phila. U.S.A.: Ohio, Cincinnati; on Corylus;1d, l8.vii.l912, 16,21.v.1914 (Braun). On Opulaster',26 6,29 9, 1 ex, 11,12,23.vii.1916; l?, 5.viii.l9l4; l?,20.v.1919 (Braun). Champaign Co., Cedar Swamp; onPhysocarpus opulifulius 299, 18.v.1934 (Braun). Kentucky, Campbell Co., Claryville; on Physocarpus opulfolius; ld, 3??, 16-25.1943 (Braun); in Phila. New Jersey, Retreat; on Cranberry;19,29.v.7915 (Sc:ammel/); New Lisbon; on Cranberry; 19, l3.v.l9l5 (Scammell); Michigan, Bois Blanc, I ex, 18.v. 1943.

Btorocv. Mine. A long upper surface ophionome; the frass pattern and, to a certain extent the mine, show variation. This is exemplified in plates 4.1 and 4.2 which show mined leaves of corylfoliella onCorylus andVacciniur?? respectively. Voltinism. Probably bivoltine in Canada and the U.S.A. <^ SOCTETY OF CANADA Stigmella ostryaefoliella (Clemens) comb. n. Nepticula ostryaefoliella Clemens, 1861: 83 Nepticula ostryaefoliella Clemens; Chambers in Hayden, 1878: 158 (listed) Nepticula ostryaefoliellrz Clemens; Dyar, 1903: 546 (listed) Nepticula ostryaefoliella Clemens; Braun, 19 l7b: 176 (misidentification) Nepticula ostryaefoLiella Clemens; Braun in Forbes, 1923: 89 (misidentification) Nepticula caryaefoliella Clemens, 1861: 84 revised synonymy (incorrectly synonymized with Nepticula juglandfoliella Clemens by Braun, l9l7b: 181) Nepticula caryaefoLiella Clemens; Chambers in Hayden 1878: 157 (listed) Nepticula caryaefoliella Clemens; Dyar 1903: 545 (listed) Nepticula obscurella Braun, 1912l. 95 syn. n. Nepticula obscurella Braun; l9l7b: 116 Nepticula obscurella Braun; Braun ln Forbes 1923: 89

DsscnrpuoN. External features: As corylfoliella except for: d, eye-caps always lacking the brownish purple shading. Wing measurements: 6. 3.5-4.0 mm (8); ?.3.6-4.4 mm (9). Genitalia: d . Figure 29; tegumen in form of narrow band, posterior edge not serrated; uncus weakly bilobed but with lobes extended into short, inwardly pointed, arcuate papillae; gnathos with horns clearly branched; valves broad with stout, long hair-like scales arising from inner aspect of tip; transtilla with horizontal bar not significantly bilobed. ?. Figure 30; the figure represents the degree of variation from corylfoliella blut the differences shown are not reliably diagnostic. Httst-plants: Ostrya (Hornbeam), Carpirzzs (Hornbeam, Blue Beech), Carya (Hickory), Myrica (Blueberry). Mine: tpper surtace ophionome.

DrecNosrs. Differs fromcorylfoliella mainly in features of the male genitalia, in particular the nature of the valves and transtilla. In the externals, the eye-caps of the male always lack the brownish purple shading. The much smaller ventral olate of the vinculum in the male of ostryaefoliella d i stingu ishes it from j u g landifo t i'e t la.

DtscusstoN. Clemens' original description of ostryaefoliella stated that the frass-line of the mine was narrow. Actually, specimens reared from Ostrya (the host-plant from which the species takes its name) have the frass-line relatively wide in parts. However, considering the intraspecific variation of this character, these specimens are identified as ostryaefoliella. The only specimen labelled osrryaefoliella by Braun is a single male collected from British Columbia and reared from Birch. We have identified this specimen as coryl(bliella, mainly from a study of the male genitalia, by which it may be readily diagnosed. Corylifuliella is also known to feed on Birch whilst oslryaefoliella does not appear to occur on this host. The genitalia of the single male specimen reared on Myrica and described by Braun (1912) as a new species, viz'. obscurella, are indistinguishable from those of ostryaefoliella; S. obscurella is therefore synonymized above. It is very likely that the Myrica-teeding myricafoliella Busck is conspecific with ostryaefoliella. Unfortunately there are no males to verify this and so the species is not synonymized. Braun (l9l7b) synonymized caryaefoliella with juglandfoliel/a. However, the male genitalia of c'aryaefoliella are the same as ostryaefoliella but different from juglandfoliella. (Compare Figs. 21,29, and 31.)

DtsrnrsurtoN. Canada: Ontario. U.S.A.: Ohio. New Jersev. W]LKINSON AND SCOBLE: THE NEPTICULIDAE OF CANADA 55

Ftc. 29. Stigmella ostr!-aeJoliella, d genitalia.

SpEcrunNs ExrNttNE,t. d holotype of obscurella; in Phila: New Jersey, Montclair; on Myrica; (Braun Coll.). Other specimens: CANADA: Ottawa, Ontario; on Carya ovata; 26 d, 9,l0.viii.1955; 29 9, 1 ex, 74,1 8.vi.1956 (Freeman). Normandale; on Carya ovata; ld, 59 9, 24-21 .vii.1956 (Freeman and Lewis) in CNC. U.S.A.: Ohio, Cincinnati; on Ostrya; I ?, 10.vii.1912, l6 , 22.v.1917 (Braun). On Carpinus'. l6 , 22.v1i.1913, 19, 7.viii.1916 labelled "Nepticula corylifoliella Clem.", 1d, 23.vir.1913 (Braun). On Carya; 1d, 1 9 , 1.vri.I912 labelled "Nepticula juglandifoliella Clem.' ', 3d d, 1 ex, 7 - 12.vii. 19 12 (B raun\. Btorocv. Mine. The mine is generally very long and similar to corylfoliella. The only slight difference appears to be in the rather wider frass-line. Plate 4.3 shows the most extreme deviation from the normal Dattern in the material examined: the mine is short and the frass-line rather narrow. 56 MEMOIRS OF THE ENTOMOLOGICAL SOCIETY OF CANADA

Frc. 30. Stigmella ostryae.foliella, ? genitalia 57 Stigmella juglandifoliella (Clemens) comb. n.

Nepticula juglandifoliel/a Clemens, I 86 I : 84 NepticuLa juglandifuliella Clemens; Chambers in Hayden, 1878: 105 (incorrectly headed'n.sp' and 1878: 157 (listed) Nepticula juglandifoliel/a Clemens; Dyar, 1903: 546 (listed) Nepticula juglandifoliel/a Clemens; Braun, I9l2: 9l Nepticula juglandfoliella Clemens; Braun, l9l7b: l8l Nepticula juglandifoliel/a Clemens; Braun ln Forbes, 1923 9O Neptic:ula caryaefoliella Clemens, 1861: 84 (incorrect synonymy by Braun, l9l2:91)

Dr,scntptlott. External features'. 6. As corylifuliella except: Wings: ground colour of forewings blackish with reddish bronze and purple reflections; one postmedial fascia, shining silver and rather narrow. Legs dark grey, pale ochreous with a grey lustre. Pl. 9.4. 9. Unknown. Wing measurements'. 6. 3.0-3.4 mm (3). Genitalia: d. Figure 3l; as corylilbliella except: vinculum with ventral plate very large; saccus very weakly bilobed; uncus with a pair of widely separated horns; gnathos sigmoid, characteristic; valves broad with style pointed; transtilla without large triangular processes at ends of lateral bars. H o s tp lant : J u glans (Walnut). Mine: upper surf'ace ophionome.

DIecNosrs. Closely related to coryliJbliella and ostryaefoliella, but is distin- guished by the overall form of the genital capsule, in particular the large ventral plate of the vinculum, the form of the valves, the nature of the uncus- and the very sigmoid gnathos.

DrscussroN. Clemens failed to cite type-material and described the species from mines which are no longer in existence. Braun's material from which she described adults of jaglandfoliella, includes specimens of corylifoliella and ostryaefoliella. ln fact, those specimens from her collection which are here regarded as truly representative of juglandifoliella, are three males which fed on Juglans. They are diagnosed from corylfoliella on the basis of the genitalia. Unfortunately the female is unknown. Chambers ln Hayden, 1978:105 was apparently the first to rear specimens of juglandifoliella from mines fitting Clemens' description. Although we have not examined these specimens, Chambers' description appears to correlate with Braun's specimens.

DtsrmnurroN. U.S.A.: Ohio. SprctlreNs ExertNno. U.S.A.: Ohio, Cincinnati; on Jullans; 26 6, 12,17 .vri.1912; I d , 10.v. 1911 (Braun).

Btor-ocv. Mine. On the limited material available this appears to be indistin- guishable fromcorylfoliella. It is not known if many mines occurring on one leaf, as in plate 4.4, is a typical feature of this species.

Stigmella myricafoLiella (Busck) comb. n. Nepticula myricafoLiella Busck, 1900: 238 Nepticula myricafolielkz Busck; Dyar, 1903: 546 (listed) Nepticula myricafoliella Busck; Braun, 1917b: 118 58 MEMOIRS OF THE ENTOMOLOGICAL SOCIETY OF CANADA

b

Ftc. 31. Stigmella juglandfoliella, d genitalia.

Busck described the species from two specimens reared from M),rica cerifera. It is unlikely that the narne is anything other than a synonym of ostryaefoliella, now known to feed onMyric:a. It is not reduced to synonymy in this study for the following reasons: l Of the two syntypes, one, the male, has lost its abdomen and so the genitalia cannot be checked. 2. The female has rather more heavy ridges on the signa of the bursa copulatrix than is typical of ostrvaefoLiella. The problem might be resolved by the rearing of a male from the type-locality. WILKINSON AND SCOBLE: THE NEPTICULIDAE OF CANADA 59 Spncrr,rsNs Ex,qurNno. We designate as lectotype the female specimen of Busck's two syntypes. It is lodged in the USNM and bears the following data: "Type. No. 4948 U.S.N.M., 893 Fla on Myrica cerifera iss Mar 28.1900., Palm Beach, Fla [Florida]"; CNC genitalia slide No. 3473. Paralecrorype: d, on.same prn. Other specimens: CANADA: Nova Scotia, Tusket; on Myrica;2??, l,26.li.lg6l (Freeman andLewis). Sable Island; onMyrica, "Bayberry"; l9 ,29.11i.1968 (Martin); in CNC.

Specimens of Uncertain Identity in the corylifoliella group CANADA: Ontario, Sparow Lake; 12? 9, 22,23.vii.1926 (Braun), l g, same data,23.vii.l926 labelled "Nepticula corylfoliella clem.". I ?, l7.vii.1926 (Braun) labelled "Nepticula paludicola Braun." in cNC. U.S.A.: Maryland, plummers Isrand; I 9 , 5.viii.1914 (shannon). New Jersey, Pemberton; I I , 3l.vii.l9r5 (Scammeil); in USNM.

Kny: the BIFAscTELLA GRoup Head-tuft bright orange-ochreous. Male with saccus weakly bilobed, as in Fig. 32. Female with bursa copulatrix heavily pectinate bfascielta (p. 59) Head-tuft dark brown. Male with saccus deeply bilobed. as in Fie. 34. Female with bursa copufatrix weakly pectinare ...... -. ."..... intermediu (p.62)

The sIFasctELLA GRoup Two species, bifasciella andintermedia, are included in this group mainly because of similarities in the uncus and gnathos of the male genitalia. noth speiies are related to other species-groups, as given in their diagnoses, and bifasciella is even closer to ceanothi, a species not represented in Canadian collections.

Stigmella bifasciella (Clemens) comb. n. Nepticula bifasciella Clemens, 1862 133 Nepticula bifasciella chambers; chambers in Hayden, 1878: r5j (listed; incorrect original author given) Nepticula bifusciella Clemens; Dyar, 1903: 545 (lisred) Nepticula bifasciella Clemens; Braun ln Forbes, 1923: 8l Nepticula serotinaeella Chambers, 1873: 126 (syn. by Braun 1917b: 170) Nepticula serotinaeella Chambers; Chambers in Hayden, 1878: 158 Nepticula serotinaeella Chambers; Chambers, l8l9: 93 Nepticula serotinaeella Chambers; Dyar, 1903: 547

DEscnrpttoN. External Jbaturesi d, ?. Head: palps grey and lustrous; antennae dark brownish grey; eye-caps shining creamy white; tuft orange-ochreous, tending to be darker on front of head; collar very dark grey. Thorax and abdomen dark brownish grey with greyish lustre. Wings: ground colour of forewings dark greyish brown or brownish black with bronze and some purple reflections, particularly proximally; two shining silver fasciae, one antemedial, one postmedial; fringe leaden grey. Legs dark grey with some purple reflections, lustrous; tarsi pale ochreous. Pl. 9.5. Wing measurements: 6 . 3.2-4.6 mm (6); 9.3.2-4.4 mm (10); holotype.4.5 mm. Genitalia: d. Figure 32; vinculum with medial posterior edge of ventral plate with slight convex bulge; tegumen narrow; saccus relatively small, with pair of small, widely separaied papillae; uncus in form of a bridge with two small lobes; gnathos U-shaped, horns each with short 60 MEMOIRS OF THE ENTOMOLOGICAL SOCIETY OF CANADA

b

Frc.32. Stigmella b{hsciella, d genitalia.

branch arising approximately half way along length; valves broad, reaching beyond uncus with small, papillate style not widely separated from cuiller; transtilla with horizontal bar of each just meeting centrally; aedeagus flask-shaped with denticulate cornuti. ?. Figure 33; anal plate deeply emarginated; ductus bursae approximately twice length of apophyses, gradually widening into bursa copulatrix, with accessory sac adorned with denticles; bursa copulatrix without signum but with wavy rows of minute pectinations; anterior and posterior apophyses of equal length-anteriores arcuate and stout terminally, posteriores straight and nartow. Host-plants: Prunus serotina, P. nigra, P. pensylvanica, ? Populus. Mine'. tpper surface ophionome. DracNosrrc Fn.4tunEs. Externals with bright reflections; forewing with two fasciae. Genitalia: d; with U-shaped gnathos; aedeagus with rectangular plate, cornuti spiculate. 9 with denticulate accessory sac; bursa copulatrix pectinate, signum lacking. DrlcNosrs. The species shows similarities in genital morphology to several of the species-groups. The male is closest to the corylifuliella grottp but the aedeagus is longer and lacks the characteristic palmate plates of that group. It is also similar to intirmedia, which lacks the liberal covering of pectinations found on the bursa of the female of bifascielta. The denticulations on the accessory sac, and the presence of an anal plate, also resemble castaneaefoliella and its relatives, but these are the only similarities. Prunus, thehost-plantof bifasciella,isamember of theRoseaceaetyp\cal of the rosaefotietla and crataegifolielta groups. The species is also related to two WILKINSON AND SCOBLE: THE NEPTICULIDAE OF CANADA 6l

Frc. 33. StigmeLLa bifasciella, I genitalia 62 MEM.TRS oF THE ENToMoLoGTcAL socrETy oF .ANADA

Ceanothus-feeders that have so far only been collected in the United States, ylz. ceanothi and dffisciae. Its closest relative is ceanothi'. bifasciella may be distinguished by its less extensive saccus and in the smaller cornuti on the vesica. In the female, the accessory sacof ceanothimay haveweakerdenticulatemarkings thanbifascietta.The external features appear to be indistinguishable. S. diffasciae is a larger species with the fasciae of the forewings white, not shining silver. In addition, there are differences in the transtilla, gnathos and aedeagus in the male, and in the pectinations on the bursa copulatrix and in the accessory sac (which is not denticulate indiffasciae) in the female. DtscussroN. There is a male, collected by Downes, which is undoubtedly bfosciella yet has been reared fromPopulus. This plant is quite unrelated to Prunus , the host from which all the other specimens of bifasciella have been recorded. However, Downes has confirmed that the information is correct (verbal communication). This is discussed in more detail in the general discussion (p. 108). DrsrnteurroN. Canada: Ontario. U.S.A.: Pennsylvania, Ohio, New york. SpscrupNs ExnurNnn. d holotype; in Phila: U.S.A.: "Type No.1493Nepticula bfasciella. B. Clemens.". "specimen No. 124, AB 1902. At light Aug. 11.". CNC slide No. 3465. Other specimens: CANADA: Ontario, Ottawa; on Prunus nigra; 19, l5.viii.1964 (Lewis). On Populus; ld, i.vi. l9lo (Downes). Normandale; on Prunus;29, 1 ex, 17-l8.vii.1956 (Freeman andLewis). U.S.A.: Pennsylvania, West Chester; onPrunus pensylvanica; 26 6, 8? 9, 31.i.-l0.iii.l966 (Lewis); in CNC. I ex, 5.viii.l9l6 (Haimbach). 1d, 1 ex, labelled "Nepticula serotinaeella Cham., (Fernald Collec- tion)." Ohio, Cincinnati; on Prunus serotinai 6dd, 5?9.2 ex, ll-20.vti.1913. 30.vi.-11.vii.1916, 13.v.1922(Braun). AdamsCo.,BeaverPond;onPrunusserotina; 2?9,3-5.v.1928 (Braun); in Phila. New York, Ithaca, 6 ml. Creek ld, 13.v.1961 (Hodges).

Btorocv. Egg. Laid on lower surface of the leaf against the mid-rib. Mine. The larva crosses from the underside of the leaf to form an upper surface tract. In some specimens frass is deposited in zig-zag arcs for much of the length whilst in others the frass-line is darker due to a more compact deposition of the grains. Pl. 5. l. Voltinism. Trivoltine in U.S.A. and perhaps also in Canada.

Stigmella intermedia (Braun) comb. n.

Nepticula intermedia Braun, l9llb: 17 1 Nepticula intermedia Braun; Braun in Forbes, 1923: 81

Dpscnlpttorv. External features: d, ? . Head: palps off-white, lustrous; antennae slate grey, lustrous; eye-caps white with white reflections; tuft on front of head and vertex very dark brown; collar dark greyish brown. Thorax grey with silver and bronze reflections. Abdomen dark grey with grey lustre. Wings: ground colour of forewings brownish black with two white fasciae, shining silver, one postmedial the other broad and sub-basal, and only distinguishable at certain angles of illumination; fiinge grey. Legs grey, lustrous, pale at tarsi. Pl. 9.6. Wing measurements. 6. 3.4 mm (3); ?. 3.4-4.0 mm (2); holotype.3.4 mm. Genitalia: d. Figure 34: tegumen in form of narrow band; saccus large, bilobed, uncus heavily sclerotized, cross-shaped; gnathos robust, with pair of strong, posteriorly pointing horns well sclerotized at tips and pair of small, anteriorly pornting lateral papillae; valves squared-off apically, not divided into style and cuiller; transtilla with horizontal bar continuous, thin in middle, thickening at ends; aedeagus fairly short; vesica with cornuti in form ofstout spines near phallotreme and curved, striate rectangular plate. WILKINSON AND SCOBLE: THE NEPIICULIDAE OF CANADA 63

+'-ffi b Frc. 34. Stigmella intermedia, d genitalia. 64 MEM.TRS oF THE ENToMo'ocrcAL soclEl y oF .ANADA

? . Figure 35; ductus bursae merging imperceptibly with bursa; bursa copulatrix with few weak pectinations; anterior apophyses shorter and broader basally than posterior apophyses which are slightly longer, straight and narrow. Host-plant'. Rhus typhina, R. aromatictt (Sumac). Mine: tpper surface ophionome.

Frc. 35. Stigmella intermedia, ? genitalia. WILKINSON AND SCOBLE: THE NEPTICULTDAE Ots CANADA 65 Dr.q.cNosts. Distinguished from the very closely related rhofoliella (not rep- resented in the Canadian collections) by having a pair of fasciae on the forewing rather than a single one. The heavily sclerotized uncus and the shape of the gnathos resemble bifasciella, but intermedlc differs in the more deeply excavated saccus, the shape of the valves and the fewer and larger spine-like cornuti on the vesica in the male genitalia. In external features intermedia may be distinguished by its dark tuft. The bursa copulatrix of the female is only weakly pectinate. In this it resembles the crataegifuIiella group but the male genitalia are diagnostic. DrscussroN. The genitalia are indistinguishable from rhofoliella. However, there is a consistent difference in the wing markings. S. rhoifoliella apparently feeds only on Rhus toxidendron. DlstntnuuoN. Canada: Ontario. U.S.A.: Ohio (type-locality). Arkansas. SpscIr,InNs ExeutNno. d holotype; in Phila: U.S.A.: Ohio, Cincinnatil. oriRhus sp. B.410; 24.v.1914 (Braun); genitalia USNM slide No. 11311. Paratype: data as holotype; l6 ,2.vi.).914 (Braun); in Phila. Other specimens: CANADA: Ontario, Ottawa; onRhus typhina; 1 6 ,25.vi.1956,29 I , 27,30.vi.1956 (Freeman). U.S.A.: Ohio, Cincinnati; on Rhus aromatica; 19, 22.v.1917,1 ex, 9.v.1918 (Braun). OnRhus sp.; I ex, 3.viii.l917 (Braun); in Phila. Arkansas, Washington Co., Devil's Den State Park, 1?, 10.vii. 1966 (Hodges) in USNM. Brorocv. Egg. Laid on lower surface of the leaf. Mine. The frass-line nearly fills the mine throughout its length except for the emergence slit. Pl. 5.2. Voltinism. According to Braun (l9l1b) there are usually two generations, but occasionally a third, per year.

Kev: the eUERCIPULcHELLA cRouP - Reflections of the forewings weak; broad, off-white postmedial fascia. Hindwings of male with basal half covered by purple, petiolate scales ...... variella (p. 67) - Reflections of forewings brilliant; narrow, white postmedial fascia with silver reflections. Hindwings of male without petiolate scales ...... quercipulchella (p. 65)

The qunncrruLCHELLA cRouP The two species comprising this group, quercipulchella and variella, have a distinctive uncus and aedeagus in the male and a characteristic accessory sac arising from the ductus bursae of the female.

S ti g me lla quer c ipulche lla (Chambers) comb. n. Nepticula quercipulchella Chambers in Hayden, 1878: 105 Nepticula quercipulchrella (sic) Chambers; Dyar, 1903: 547 (listed) Nepticula quercipulchella Chambers; Braun, 1911b: 180 Nepticula quercipulchel/a Chambers; Braun in Forbes, 1923 90 Nepticula terminella Braun, l9l4:23 syn. n. Nepticula terminella Braun; Braun, l9l7b: 114 Nepticula terminella Braun; Braun in Forb.s, 1923: 88 DescrrpttoN. External .features: 6 , I . Head: palps whitish; antennae greyish brown with grey lustre; eye-caps white and shining white; tuft on front of head deep ochre; tuft on vertex 66 MEM.IRS oF THE ENToMoLoctcAL socIETY oF CANADA brownish black: collar dull white. Thorax and abdomen black with bronze reflections. Wings: ground colour of forewings greyish black with violet, bronze, golden, red and green reflections; one white, postmedial fascia, shining silver; fringe grey with purple reflections; hindwings greyish brown with purple dusting; fringe grey, with purple reflections. Legs darkish grey, becoming whitish with white lustre towards tarsi. Wing measurements'. 6 . 4.4-4.6 mm (6); ?. 4.6-5.0 mm (10)' Genitalia: d . Figure 36; tegumen bluntly rounded posteriorly; saccus comprising two widely separated papillae; uncus papillate; gnathos roughly M-shaped with lateral arms widely separated by long horizontal bar; aedeagus long, becoming weakly arcuate basally; cornutus coverrng vesica as sharp spines and denticles.

FIc. 36. Stigmella quercipulchella, d genitalia. WILKINSON AND SCOBLE: THE NEPI'ICULIDAB OF CANADA 61 ?. Figure 37; ductus bursae with accessory sac in fbrm of a large, characteristic spiral; anterior apophyses arcuate, stout; posterior apophyses narrow. Host-plant'. Quercus rzbra (Red Oak). Mine: upper surface ophionome.

Dta.cNosIs. S. quercipulcheLla has brilliant reflections on the forewing and a narrow, shining silver fascia, whllstvariella has weak reflections and a broad, off-white fascia. The wing span of quercipulchella is smaller than in variella . The hindwing of the male of quercipulchella lacks the deep purple, petiolate scales found invariella.

DrscussIoN. Braun (19l7b) noted the similarity between quercipulchella and terminella but kept the species separate because Chambers recorded the larva of quercipulchella as green, whereas the material of terminella studied by Braun was yellow. Chambers' account implies that only one larva was studied, from which the male, imaginal holotype was reared. As both the externals and genitalia of this type are indistinguishable from terminella, and as the colour of the larva of Chambers' specimen does not seem to have been a well evaluated character , terminella is reduced to a junior synonym of quercipulchella. The mis-spelling of quercipulchella as "quercipulc:hrella" by Dyar (1903) probably arose because the latin for beautiful is pulchra. However, although the original spelling suggests improper latinization, quercipulchel/a is retained (International Code, Article 32(a)1i). That Chambers' spelling is not a lapsus calami is indicated by his several references to the name in the same paper.

DrsrnrsurroN. Canada: Onrario. U.S.A.: Kentucky (type locality), Ohio. SpBcIur,Ns ExnltNr,o. d holotype; in MCZ: U.S.A.: Kentucky; "Chamb. Kentucky, 161". Type No. 14957; genitalia, CNC slide No. 3489. 16 syntypes of terminella, each labelled "Cotype". U.S.A.: Ohio Cincinnati;onQuercus;B.652. 16, 3.v.1913; 16 6,7 ? ?, I ex, l0-l9.viii.1913 (Braun); in Phila. Other specimens: 1 ex, data as Braun's syntypes; 28.iv.1917. CANADA: Ontario, Bells Corner; on Quercus rubra; I d, 1 I , 26.iv.1967 (Freeman and Lewis)i in CNC.

Btorocv. Mine. The tract is narrow at the start but widens considerably by the end. The frass-line is dark and remains central and compact throughout the length of the mine. Pl. 5.3. Larva. According to Chambers (1878: 105) the larvae are bright green. They are probably more usually yellow, see discussion. Chambers also suggested that the larva, from which he reared the holotype, made two separate mines on the same leaf. If this is true then, as Braun (l9l7b) points out, it is at variance with all other observations made on the familv.

Stigmella variella (Braun) comb. n. Neptir:ula variella Braun, 1910: 173 Nepticula variella Braun; Braun, 19l1b: 187

DescnIprtoN. ExternaL features as quercipukhella except for: tuft on front of head and vertex pale ochreous to ochreous; collar whitish. Thorax and abdomen brownish grey. Wings: ground colour of fbrewings greyish brown with weak bronzy and purple ret'lections: broad postmedial, off'-white fascia; irregular terminal band or patch of white scales overlapping fringe. Serual dimorphism'. 6. Hindwing with basal half covered by purple, petiolate scales. Variation: the variation suggested by the name of this species is evident in the fbrewings- MEMOIRS OF THE ENTOMOLOGICAL SOCIETY OF CANADA

1 -mm5

Frc. 37. Stigmella quercipulchella, ? genitalia WILKINSON AND SCOBLE: THE NEPTTCULIDAE OF CANADA 69 The ground colour is of variable intensity and the fasciae often weakly defined. Wing measurements'. 6 . 6.2-7.4 mm (6); I . 6.0-7.2 mm (2); holotype.7.4 mm. Genitaliat d, I . As quercipulchella. Host-plant'. Quercus agrifulia (California live oak). Mine: no specimens available. Braun (1917&) records it as an ophionome.

DIecNosIs. S. variella has weak reflections on the forewing and a broad, off-white fascia, whllst quercipulchella has brilliant reflections and a narrow, shining silver fascia. The wing span of variella is greater than in quercipulchella. The hindwing of the male of variella has deep purple, petiolate scales, but these are absent in quercipulche lla .

DrscussroN. There are no apparent differences between the genitalia of quercipulchella and variella, although the external features are easily distinguishable. S. variella has been collected only from California whilst quercipulchella (:terminella) is known from Kentucky, Ohio, and Ontario. On these criteria a case could be made for subspecific designation (see p. 67). Braun's type material of variella includes 12 specimens labelled as "Paratype" and five as "Type". Amongst the latter, one has an additional label "Cotype", one "Paratype", and one "Type". In her original description Braun stated "Type [i.e. singular] in my collection, paratypes in Californian Academy of Sciences and in Mr. W.D. Kearfott's Collection". In accordance with the International Code, Article 73(b), the specimen labelled "Type, Collection of Annette F. Braun" and bearing the additional label "Nepticula variella Type", is regarded as the holotype.

DlstnrsurroN. U.S.A.: California (type locality).

SpncIuBNs ExnutNno. d holotype. ln Phila; U.S.A.: California, Mills College, on Quercus agrfolia, iii. I 908 (P ilate, Braun Coll. ); genitalia, USNM slide No. 17 316. Paratypes: data as holotype; lOd d, 4? I , I ex, (10 ex labelled Alameda Co., no date given on 7 ex-presumed 1908), (Pilate, Coll. of A.F. Braun); in Phila.

Specimen f One of Braun's female paratypes of variella has the accessory sac considerably shorter and wider than the other specimens and, also, strikingly covered with numerous, well sclerotized spicules. Of great interest is Braun's breeding record which shows that the specimen was reared from "Tan Bark Oak". It is the only example of the species that we have examined from a food-plant other than Quercus agrifulia. Despite the variable nature of the external characters of variella, the genitalia are remarkably uniform except for this specimen which is here treated separately, see figure 38. Iiling measurements'. 9.7.4 mm. SppcIlrsN ExeurNno. U.S.A.: California, Mills College, on "Tan Bark Oak". l9 , 2l.vi.?19O8 (Pilate); paratype of variella labelled "TYPE, Collection of Annette F. Braun"; in Phila.

KEv: the PRocRASTINELLA GRoUP

Forewing off-white, except for purplish brown area medial to postmedial . . . . alba (p.73) Forewing grey-brown with irregularly-shaped, shining silver, postmedial fascra . prouastinella (p. 70) lv MEMOIRS OF THE ENTOMOLOGICAL SOCIETY OF CANADA

1 mm 2

Ftc. 38. Specimen f, ? genitalia

The pr.ocn,q,srINELLA GRoUP The male genitalia are characterized by a tongue-shaped, medial projection forming part of the gnathos. They resemble Ectoedemia in a few minor ways, but the group lacks the major diagnostic characters of that genus. On balance it is included in Stigmella in preference to erecting a new genus.

Stigmella procrastinella (Braun) comb. n. Nepticula procrastinella Braun, 1921: 59 DescnrprtoN. Erternal features: d. ?. Head: oalps off-white, lustrous; antennae extending just beyond half length of wings, greyish purple; eye-caps white, shining white; tuft on front of head and vertex orange-ochreous; collar as eye-caps. Thorax and abdomen grey-brown, lustrous. Wings: ground colour of forewings grev-brown with cupreous, and proximallv purple, reflections and a few white scales antemedially on anal edge of wing; two white, postmedial, WILKINSON AND SCOBLE: THE NEPTICULIDAE OF CANADA 1 I triangular patches shining silver, one costal, one anal which meet apically so forming biconcave fascia; fringe grey, proximally irrorate with forewing scales which form a clearly delineated band across terminal cilia; hindwings light brownish grey with cupreous reflections; fringe grey with cupreous reflections. Legs grey with cupreous reflections. Pl. 10.1. Wing measurements: 6 . 5.2-6.0 mm (7); ? . 5.2-5.1 mm (3); holotype.6'2 mm. Genitalia: d. Figure 39; tegumen rounded; saccus bilobed; uncus rounded, with median bifid papilla; gnathos in shape ofinverted Y, with lateral processes broad at end, then narrowing to fuse into single, pointed medial limb; valves reaching uncus with cuiller widely separated from and less pointed than style, with inwardly directed protuberance approximately halfway down each valve; transtilla with processes of lateral arms extending deeply; aedeagus with densely spinose and denticulate cornutus. 9. Figure 40: ductus bursae longer than apophyses: bursa copulatrix. or accessory sac. with signum comprising an irregular patch of heavily sclerotized denticles; apophyses broadening basally in both pairs and extending approximately two-thirds length of ductus, anteriores straight; posteriores gently arcuate. Host-plant: Ostrya virg,iniana (Hop-hornbeam). The adults were collected from this food-plant but were not reared-through from the larvae (Braun 1927: 59). Mine: unknown.

b

Frc. 39. Stigmella procrastinella. d genitalia 12 MEMOIRS OF THE ENTOMOLOGICAL SOCIETY OF CANADA

Ftc.FIG. 40. Stigmella procrastinella,procrastineLla, ? genitalia.

DtlcNosrs. The genitalia are characteristic, particularly the uncus, gnathos, and valves in the male, and the signa in the female. The forewings lack the expanses of off-white found in the new species immediately following, giving procrastinella a darker appearance. The tuft is a brighter orange-ochreolus in procrastinella. WILKINSON AND SCOBLE: THE NEPTICULIDAts Ots CANADA 13 DrstrusurroN. Canada: Ontario. U.S.A.: Virginia. DrscussroN. The form of the gnathos and the relatively long antennae resemble Ectoedemia; however, the wing venation, the vinculum, tegumen and uncus are typical of Stigmella. The signum, although very characteristic, is not like Ectoedemia. The genitalia of these specimens are indistinguishable from the new species immediately following, but the external characters of each differ clearly. SppctnnNs ExeurNno. d holotype; in Phila: CANADA: Ontario, Sparrow Lake; l2.vii.l926(Braun). Paratypes: data as holotype; 6d d,39 9, I ex, 12.v1i.1926l,19, 16.vii.1926 (Braun) in Phila. Other specimen: U.S.A.: Virginia, Mountain Lake; 1d, 23.v1i.194O (Milne and Milne); in USNM.

Stigmella alba sp. n. DsscnrpttoN. External features: 6 , ? . As procrastinella except: Head: tuft on front of head and vertex ochreous, slightly paler. Thorax off-white. Wings: forewing colour basal to medial, off-white, dull, irrorate with purplish brown scales particularly on costa; medial to postmedial purplish brown; postmedial to subterminal off-white, shining; subterminal to terminal purplish brown. Pl. 10.2. Wing measurements. 3.6.0-6.2 mm (4); ?. 5.4-5.6 mm (3); holotype. 6.0 mm. Genitalia: 6 , ? . As procrastinella . The off-white thorax and extensive pale markings on the forewings distinguish alba from procrastinella. DrscussIoN. Although alba has genitalia indistinguishable from those of procrastinella the external features are clearly different and alba is treated as a full species. Because their genitalia are so similar and their distribution allopatric, a subspecific distinction has been considered but rejected until more of the biology is known (see general discussion p. 108). DrstmsurIoN. Canada: British Columbia. U.S.A.: Arizona (type locality). SppcrlrnNs ExlutNso. d holotype; in USNM; U.S.A.: Arizona, Coconino Co., l6milesSWof Flagstaff, WestFork6,500ft; l3.vii.196l (Hodges). Paratypes:dataas holotype; 26 6, 1?, Jrlz miles NW of Flagstaff, Fort Valley, 7350 ft. ld, 19, 2l.vi.l96l (Hodges); in USNM. CANADA: British Columbia, Victoria; 2dd, '7.vi.1922, 16.vi.1923, 19,9.vi.1922 (Carter); in USNM. Victoria; 19, I ex, 8,12.vi.1922 (Carter);26 6, no data labels, probably same series. 29 ? , 21.vi.1923 (Blackmore); in UBC. Victoria; 3? ? , 14-21.vi.1923 (Blackmore) in Phila.

GENUS ECTOEDEMIA Btsck Ectoedemia Busck, 1901: 91 . Type-species by original designation and monotypy Ectoedemia populella Busck, 1907: 98 Ectoedemia Busck; Braun, 19l7b: 196 Ectoedemia Busck; Braun in Forbes, 1923 82 Ectoedemia Busck; Borkowski. 197%t: 693 Dechtiria Beirne; 1945 204. Type-species by original designation Tinea subbi- maculella Haworth, 1828: 583, Lepid. Br. Syn. by Svensson 1966l. 200 TriJurcula Zelle4 sensr Johansson, 197 l:245 (Ectoedemia given subgeneric rank) DsscnIpttoN. Efiernal .features: d, ?. Head: palps extend beyond labrum; antennae sometimes over, sometimes just under half length of wings. Wings. Venation as in figure 3. Forewing with M and Cu fused basally; M meeting Rs tinally zLrising towards apex as M; R4 and 14 MEMoTRS ots THE BNToMOLOGTcAL socrtsTy oF cANADA

R5 separate terminally; hindwing with Rs and M each arising as single branches apically tiom common trunk. Wing markings: if present, in form of fasciae or patches. Legs: Hinglegs with proximal pair of tibial spurs below, or in middle of tibiae. Genitalia: d. Vinculum ring-shaped; tegmen extended into prominent pseuduncus; true uncus membranous, difficult to discern; gnathos W-shaped in ventral view with middle member tongue-shaped; aedeagus with spine-like anellus. ? . Ductus bursae usually with funicular colliculum; bursa copulatrix with pair of large signa comprising reticulum of "cells". Dt,qcNosrs. Distinguished from all the other genera in details of the wing venation. In particular it differs fromstigmella inthe following ways: in the forewing M and Cu are fused basally, R4 and R5 are represented by separate branches, the tegumen of the male genitalia is produced into a pseuduncus and the gnathos is W-shaped in ventral view. The presence of signa reticulata in the female genitalia distinguishes Ectoedemia not only f rom Stigmella but also fuom Glaucolepis . M is not branched in the forewing of Ectoedemia whereas inObrussa M1 and M2 are present. The absence of spine-like processes associated with the valvae in the male, and the lack of an antrum on the ductus bursae of the female diagnose it further fromObrussa. In the forewing of Ectoedemia all branches of Rs arise from the same trunk whereas inTrifurcula only Rl and R2*3 arise from a common trunk. In the hindwine of Ectoedemia, M is not divided into Ml 12 and M3 as in Trifurcula.

KEv: EcroE,DEMrA l. Antennae extending just less than half the length of wings. Forewings with silver or white markings ...... 2 Antennae extending beyond half the length of wings. Forewings without silver or white markings ...... 3 2. Distal pair of spurs on hindlegs in middle of tibiae. Wing tip scales forming a continuous dark band overlapping the fringe ...... the clemensella group (p. 86) Distal pair of spurs on hindlegs below middle of tibiae. Wing tip scales do not form distinct band overlapping fringe ...... lindtlulstl (p. 83) 3. Forewings cupreous brown. Larva causes gall formation on petioles of leaves " " " PoPulella (P' 7a) Forewings predominantly grey, irrorate with cupreous brown. Larva mines petiole and leaf blade, forming only slight callus tissue in petiole . . the argyropez,a grotp (p. 77)

Ecto e demia p opu le lla Busck Ectoedemia populella Busck, 1907: 98 Ectoedemia populella Busck; Braun, l9l'7b: 191 Ec:toedemia populella Busck; Braun ln Forbes, 1923: 83 Ectoedemia populella Busck; Borkowski, 1912a:697 (discussion, d genitalia figured) DrscnrpttoN. External .fbatures: 6 , ?. Head: palps, maxillaries greyish brown with grey lustre, labials paler; antennae extend just over half length of wings with pale bronze reflections; eye-caps white; tuft on front of head and vertex ranging from pale ochreous to brown; collar dull white to pale yeltow. Thorax and abdomen brownish, lustrous. Wings: ground colour of forewings cupreous brown, dusted with grey, with copper reflections; fringe grey, variously irrorate with forewing scales; hindwings light brownish grey or greyish brown; fringe grey, cilia sometimes slightly paler atroots. Legs grey, lustrous; proximal pair of spurs on middle of hind tibia. Wing measurements: 6 . 6.2-8.0 mm (12); ? . 7.4-9.6 mm (6). Genitalia'. d. Figure 4l;vinculum with ventral plate large, weakly bilobed, extended to corners formed where ventral Dlate meets anterior ends of lateral arms: tegumen extended into WILKINSON AND SCOBLE: THE NEPTTCULIDAE OF CANADA 75

b

Ftc. 41. Ectoedemia populella, d genitalia.

tapering, blunt pseuduncus (see figure); gnathos with large medial projection; valves basally broad, heavily sclerotized, curving and narrowing towards apex; transtilla well sclerotized, lateral arms stout, arcuate, and ending in long straight process, horizontal bar narrower, arcuare; aedeagus straight with 2 pairs of stout anellar projections, one pair lateral, the other central as in figure; cornutus comprising arcuate, striate thickening. ? . Figure 42. anal papillae short; ductus bursae one and a half times length of apophyses; colliculum ring-shaped with rod-like sclerite and associated denticulate pocket, see figure; bursa copulatrix with pair of comparatively small, elongate signa; anterior apophyses each with broad basal plate; posterior apophyses narrow, slightly broader basally. H o s t- p lant s . P opulus tr e mulo ide s, P . gr andi de nt at a. Mine: the larva causes gall formation on the petiole (Busck 1907; Braun rgl'lb). - Dtacuosrs. Larger than all the other species here described. The cupreous brown forewings are also characteristic. Differs from canutus in features of the genitalia. The larvae induce the formation of galls on the petiole, but do not mine the leaf-blade. In this they differ from the larvae of argyropeza which mine both the leaf-blade and the petiole. They cause the production of some callus in the petiole, but, unlike populella, do not form rounded galls. The female genitalia of populella differ from lindquisti by having a denticulate pocket associated with the colliculum and less sinuate apophyses. , The gall forming habit of the larva distinguishes populella from the leaf-mining p-latanella group. In addition the antennae of populella are longer and the fringe on the forewing is variously irrorate with scales of forewing ground colour and not arianged in a band over the cilia as inplatanella and its relatives. In the female genitalia the signa 16 MEMOIRS OF THE ENTOMOLOGICAL SOCIETY OF CANADA

colliculum

t"

Ftc.12. Ectoedemia populella, ? genitalia WTLKINSON AND Sf,]OBLE: THE NEPTICULTDAE OF CANADA 17 and the "cells" that comprise them in populella are rather smaller. The species of Ectoedemia described by Busck (1913 102-104), so far not collected in Canada, is most closely related to populella, but, from Busck's paper, differs in external appearance, and also in the genitalia. DrscussIoN. Busck (1907: 98) did not designate a holotype. The'very large series of moths bred .' are thus all syntypes. M.J.S. has examined a pair of these syntypes deposited in the Transvaal Museum, Pretoria. A lectotype is not designated from these as this will be done later (in the series on the 'Moths of North America') from the large series in the USNM following the recommendation of the code. We have also compared drawings sent by Dr Davis, of the genitalia of a male and a female syntype from this series with the material we examined. DtstnrsutIoN. Canada: Ontario, S.E. Manitoba (Prentice et al . 1,965). U.S.A.: New York, Ohio (Braun), New Hampshire (Koebele), Massachusetts (Busck). SpEcruENs ExemuEo. I d, 1 I syntypes; in Transvaal Museum, Pretoria: U.S.A.: 6 '3238 15.v.1885 Poplar; Type No. 9904. U.S.N.M.' ? '3238 19.v.1885; Type No. 9904. U.S.N.M.' Transvaal Museum type nos. 4839 and 4840. Other specimens: CANADA: Ontario, Bells Corner 2? 9, 30.v.1965 (Sattler). Lake Erie, Cultus; on ; 19, l.iii.l962. La Passe; on Aspen; 1d, 10.xii.70; 1d, l.iii.197l (Lewis). Minden; 16,21.v.1931 (McDunnough); in CNC. Bradford; on Populus tremuloides; | 6 , 29 ? , I ex, 28.v.1970. On P. grandidentata; 26 6 , 59 ?, 3 ex, 28.v.1970. Nestor Falls; on Populus tremuloides; 26 6 , ll.ii.,7.iii.1963; in Soo. U.S.A.: New York, Monroe Co.; on poplar; 86 6 ,29 9,2 ex, 1-16.v.1948 (Kimbal/). Aweme, Man.; I ? , 24.v.1921 (Criddle); in USNM. Bror-ocv. The larvae form galls on the leaf petiole. These are swellings situated close to the leaf. The larvae mature in October and the imagines emerge in May of the following year (Busck 1907; Braun 1917b). The specimens in the CNC that emerged earlier than this were probably "forced" in the laboratory. Voltinism. Univoltine in both Canada and the U.S.A.

Ksy: the ARGyRopEZA GRoup l. Female with broad, flat ovipositor. Probably parthenogenetic . argyropeza 2 Female with ovipositor rounded, nor broad. Not parthenogenetic canutus (p. 81) 2. Forewings dark, heavily irrorate with purplish distally downesi (p. 80) Forewings rather pale, without heavy irrorations distally a. argyropeza (p. 80)

The ancynoPEZA GRoUP The two following species have been combined in a species-group: both feed on Populus and the female genitalia are similar. In external features they are difficult to separate, although there are slight differences.

E c t o e de mi a ar gy r o p e za (Zeller) Lyonetia ar gyropeza Zeller, 1839: 215 Nepticula argyropeza (Zeller) Zeller, 1848: 320 Nepticula argyropeza (Zeller); Petersen, 1930: 78 (figure of d genitalia; possible misidentification) Nepticula argyropeza (Zeller); Hering l95l: 232 (biology; figs. 148, 149) 78 MEMOIRS OF THE ENTOMOLOGICAL SOCIETY OF CANADA Nepticula turbidella Zeller sensu Herrich-Schdffer, 1855: 357; Downes, 1968: 1078 (misidentified; biology) Dechtiria argyropeza (.Zeller) Beirne, 1945: 205 D e chtiria ar gyropeza (Zeller) ; Emmet, 191 1 : 242 (biology) Dechtiria argyropeza (Zeller); Borkowski, 1912a: 698 Stigmella argyropeza (Zeller) Borkowski, 1969:. 107 Stigmalla (sic) turbidella (Zeller) sensuHenich-Schhffer 1855: 357; Cochaux, 1969: 12 (misidentified; biology) Trifurcula (Ectoedemia) argyropeza (Zeller) sensa Johansson, 1971: 245 (listed) (ZeIler) Emmet, 1,916: 189 DrscnrpIoN. Externalfeatures'. ? . Head: palps whitish and lustrous; antennae dark grey, extending more than half the length of wings; eye-caps white with white reflections; tuft on front of head and vertex pale ochreous to whitish; collar off-white. Thorax and abdomen brownish grey. Wings: ground colour of forewings grey, irrorate with cupreous brown, particularly distally, with copper and silvery grey reflections; sometimes white patch on costal margin; fringe grey with scales the colour of forewing ground colour forming well defined band across base of terminal cilia. Legs grey, with silvery grey lustre; proximal pair of spurs on hindleg just below middle of tibia. Genitalia: ?. Figure 43; anal papillae very broad; ductus bursae with colliculum having denticulate pocket; bursa copulatrix with signa sometimes extending nearly total length, sometimes less than half length of bursa; anterior apophyses with broad base, curved towards tip; posterior apophyses narrow, usually with slight swelling distally in at least one of the pair. Host-plants: , P. alba. Mine'. a petiole mine extending into leaf blade to form stigmatonome. Pls. 6.I,6.2. DrrcNosrs. Distinguished from "turbidella" (sensu marionella Ford) by fea- tures of the female genitalia: the posterior apophyses of turbidella, unlike argyropeza, are markedly swollen at their ends to form club-like structures; the anal papillae are pointed in turbidella, whereas in argyropeza they are flat and wide. Large patches of white on the forewings of "turbidella" are not present inargyropeza. It differs from canutus sp. n. in the considerably less wide ovipositor of the latter, in forewing ground colour (see Diagnosis of canutus sp. n.), and in being parthenogenetic. DrscussroN. The specimens in the CNC were identified asturbidella by Cochaux ( 1969). We have examined the external features of the type of argyropeza, lodged in the BM(NH), and have dissected a specimen from the type locality. The Canadian specimens have darker forewings than the type, but otherwise they seem identical. They are regarded as representing a subspecies of argyropeza.In a letter to Dr. K. Sattler, Mr. R. Johannson states thatE. argyropeza is "common and much variable" in Europe andZeller in his original description gives three "varieties". The species is apparently parthenogenetic in both Europe (Borkowski 1972a:698): and in Canada. Cochaux (1969: 12) states: "Reproduction was parthenogenetic; unmated females laid fertile eggs". The figuring of the male genitalia by Petersen (1930: 78) implies that males do exist although it is possible that his material was incorrectly identified. There is considerable confusion as to the identity of turbidella Zeller. In Mr. Johansson's letter he further states that ". . . Zeller' s description of turbidella is more in agreement with argyropeza than the species known as turbidella 1: populialbae . M. Her, and marionella Ford). The first descriptions of turbidella were published by Glitz (1812) and Wocke (1877), since then almost all authors have followed Wocke's interpretation". We have made a slide of a female specimen of turbidella sensu Glitz and Wocke. and. althoush related toars\)roDeza. this certainly differs in those features WILKINSON AND SCOBT,E: THE NEPTICULIDAE OF CANADA 19

Frc. 43. Ectoedemia argyropeza, ? genitalia. 80 MEM.TRS oF THts EN'oMoLOcrcAL socIETy ots .ANADA given in the diagnosis: the externals are clearly distinguishable. The situation is further complicated by Herrich-Shhffer's colour illustration of argyropeza which does not look like Zeller's type. The type of turbtdella is probably not extant.

Ectoedemia arByropeza argyropezG (Zeller). (European) DracNosrs. Differs from the new subspecies in having paler forewing colour. DrsrrrsurroN. EUROPE. Spr,crl.reNs ExlnrrNE,o. 9 lectotype in BM(NH) bearing the following data. "Gross Glogau, Silesia Zeller 1.8}.101291 Zeller Collection Walsingham Collection 1910-421 l0l29l, Argyropeza Zeller 1839:215. Lvcnetia A. Bucculatrix Isis 1839:

215" . Other specimen: Locality as lectotype; 19,22.v.1 853 (Zeller): Walsingham Coll. ex Zeller Coll., 1910-427 101289; in BM(NH). Other specimens in the BM(NH) series have been looked at.

Ectoedemia argyropeza downesi ssp. n. Wing measurements'. Holotype.6 mm; paratypes.5.2-6.4 mm (34? 9). DrecNostrc Fparunss. Wings: ground colour of forewings grey, heavily irrorate with purplish brown scales distally, more lightly irrorate proximally; some specimens with patch of white scales on costal edge forming incomplete post-medial fascia. Host-plant: Populus tremuloides (Trembling Aspen), ? P. grandidentata (Large Tooth Aspen). Mine: petiole mine, larva later forms stigmatonome in leaf blade. DrecNosrs. Differs from the nominate subspecies in having darker forewings. DtscusstoN. E. argyropeza is known to be variable in Europe and it is probably polytypic throughout its range. The subspecies is named after J.A. Downes who collected many of the specimens and, in addition, published valuable observations on the feeding habits of the adult, particularly the function of the tongue. DtsrmsurroN. CANADA: Ontario, Quebec. SpsctueNs Exetr,ttNno. I holotype in CNC; Ontario, Ottawa; on Trembling Aspen; 29.v.1969 (Downes). Paratypes: data as holotype: 1299,29.v.1969.3?9, 28.v.1968; 29 9, 31.v.1968; 39 9, 4.vi.1968:' 29 9, 7.vi.1968 (Downes) each labelled "Voucher specimen, J.A. Downes, Can. Ent. 100: 1078"; 29 9,28.v.19681 l?, 4.vi. 1968 (Downes) not labelled as voucher specimens. On poplar; 20? I, 6.v.1941 (Freeman). Ste. Anne de Bellevue, P.Q., Morgan Arboretum; 4? 9, I vi.l969 (Sheppard). Btor-ocv. The following is summarized from Coachaux (1969: 1,2). Egg. Laid on the petiole. The number of eggs per leaf depends on leaf size and the position of the leaf on the tree: eggs are more abundant on leaves at lower levels. Mine. Whilst a certain amount of callus tissue is formed in the petiole of Populus tremuloides, in P. grandidentata this may, apparently, block the mine, preventing the larvae from developing: the larvae seldom reach maturity on this host. A green zone extends along the mined portions of the leaf-blade, even after the leaf has turned yellow and fallen to the ground ("green-islands" of Hering). Larva. Remains in the mine until after leaf fall in late October and earlv November. Overwinters in sround litter or soil in cocoon. WILKINSON AND SCOBLE: THE NEPTICULIDAE OF CANADA 8 I Pupation. Occurs in early spring. Voltinism. One generation per year. Other notes. Cochaux (loc. cit.) suggests that the very high populations of this species may be due to the parthenogenetic reproduction.

Ectoedemia cnnutus sp. n. Drscnrptlor.r. Erternal features: d. Head: palps whitish, lustrous; antennae grey, extending just more than half length of wings; eye-caps white with white reflections; tuft on front of head and vertex yellowish white; collar as tuft. Thorax and abdomen dark brownish grey. Wings: ground colour of forewings pale greyish, irrorate with variable number of cupreous brown scales with weak copper and silvery grey reflections; fringe grey, irrorate with wing scales; hindwings with pale yellow tuft of cilia arising from base of costal margin. Legs grey, lustrous, paling at tarsi; proximal pair of tibial spurs on hind-leg just below middle. P1. 10.3. I . As male except for absence of yetlow tuft at base of hindwing. Wing measurements: 6. 5.8-6.6 mm (4); 9.6.0-6.2 mm (2); holotype. 5.8 mm. Genitalia'. d. Figure 44; vinculum with ventral plate narrow; saccus weakly bilobed; gnathos with lateral processes excavated terminally, median expansion thinner; valves tapering gradually towards apex; vesica of aedeagus with denticulate cornuti and striate thickening; anellar projections comprising a pair of stout, long, lateral, spine-like processes and a shorter, finger-like medial pair. ? . Figure 45 as argyropet.T except: ovipositor more rounded and less wide.

H ost- p lant : P opulus bal s amfer a.

b

Ftc.44. Ectoedemia canutus. d senitalia 82 MEMOIRS OF THE ENTOMOLOGICAL SOCIETY OF CANADA

F

Frc. 45. Ectoedemia (:anutus. ? genitalia. DrrcNosrs. r".,-.T;^.;";;Jffi:;;; in the more 0",",:: ovipositor: compare Figs. 43 and 45. E. canutus is darker than the nominate subspecies of irgyropeza ind differs from argyropeza downesi in the more uniformly scattered irrorations of the forewing which tend to give it a slightly darker overall appearance. E. canutus is not parthenogenetic. DrscussroN. It is possible that development on different host-plant species may cause observable differences in an . For example, parthenogenesis is often due to environmental effects and it is conceivable that argyropezd occlurs only as parth- enogenetic females when reared on P. tremuloides b.u.t as both males and females on P. balsamifera (thehost of canutus). However, forthepresent, argyropezaandcanutus are regarded as being two separate species due to the differences given in their respective diagnoses and because there is no evidence to suggest that males of canutus will mate with females of argyropeza and result in the production of fertilized, viable eggs. DtstxIsuttoN. Canada: Ontario. SpecrNIeNS ExnutNnl. d holotype; in Soo: CANADA: Ontario' Angus; on Populus balsamfera;4.tii.1969;68.5.3397.01, Soo No. 9. Paratypes: data as holotype: 6d d, 39 9, 3-11.iii.1969. Data as holotype: 705.0074.01; 5d d , l?, 2 ex, 28.v.19'lO:' in Soo. Ervuor-ocv. Canutus (L.)-Ash coloured. Brolocy. Apart from the host-plant, given above, we have no other data.

Ectoedemia lindquisti (Freeman) comb. n. Nepticula lindquisti Freeman, 1962: 522 Nepticula lindquisti Freeman; Lindquist, 1962: 524 (biology) This species is not included in the clemensella group, although it is extremely close. The very clear band of scales across the terminal cilia of the forewing is absent and the position of the proximal pair of tibial spurs on the hindleg is below the middle. The female genitalia show similarities to those of populella. just DEscnrpiloN. External features: 6, ?. Head: palps whitish; antennae less than half length of wings, pate ochreous, with grey reflections; eye-caps shining white; tuft on front of head and vertex ochreous to dark ochreous: collar grey. Thorax and abdomen dark brownish grey to black with grey reflections. Wings: ground colour of forewings dark brownish grey with grey, bronze and purple reflections, with single, emarginated, postmedial fascia with silver reflections; fringe grey, iriorate with scales of forewing ground-colour, which do not form a band across t"r1nlnut ciliu. L"gs dark grey, lustrous, with tarsi buff to ochreous; proximal tibial spurs of hindlegs below middle. Pl. 10.4. wing 3. 4.1-6'0 mm (26); ? ' 5'2-6'6 mm (12); holotype' 6 2 mm' Genitalia:^noturrments'. d. Figure 46; vinculum with ventral plate narrow; tegumen extended into long, tapering pseuduncus; gnathos with medial process very short, blunt; valves fairly broad, gently arcuate, iapering to pointed apex; transtilla with processes of lateral arms long; aedeagus with ^pair of bifid anellar projections; sometimes with arcuate, striate thickening on vesica. ?. Figure 47; as populella except: ductus bursae with colliculum lacking associated denticulate pocket; bursa copulatrix with minute scallop-shaped pectinations; apophyses less straight, anteriores with basal plates smaller. Host-plants'. Betula papyrifera, B. Lutea. Mine: an uooer surface stiqmatonome. 84 MEMOIRS OF THE ENTOMOLOGICAL SOCIETY OF CANADA

b

FIc. 46. Ectoedemia lindquisti, d genitalia.

Drl.cNosts. Differs from populella and canutus in having a fascia and in the colour of the forewings. In the male genitalia lindquisti has a more deeply bilobed ventral plate than populella, and the medial process of the gnathos is wider and shorter than both that species and canutus. The food plants differ and the larvae of lindquisti mine in the blade of the leaf at all times and not the petiole. DtscussroN. There are 38 specimens labelled "Paratype" deposited in the CNC, although Freeman (1962 523) only quotes 34 in the original description of the species. In addition, eight of these have a slightly different date from that quoted in the original citation. The exact specimens that comprise the paratypes of the type-series (International Code, Article 72b) are uncertain. E. Iindquisti is not included in the clemensella group because the very clear terminal band across the cilia of the forewing is absent, and because of the striking similarities in the female genitalia to populella. DIsrnrgutIoN. Canada: Ontario. U.S.A.: Maine. SpEcIvtnNs ExlnatNr,r. d holotype; in CNC: CANADA: Ontario, Lake Huron, Wiarton; on Betula papyrfera;13.vi.1959; Forest Insect Survey No. S59.0049.01; Type No. 7752. Labelled "Paratype": data as holotype; 266 6, 12? 9, S.vi- 1.viii.1959: in CNC. WILKINSON AND SCOBLE: THF NEPI ICULTDAE OT CA\ADA 85

Frc. 47 . Ectoedemia lindquisti, I genitalia 86 MEMOIRS OF THE ENTOMOLOCICAL SOCIETY OF CANADA Other specimens: 1?. data as types; S.vi.1959; in Phila. Ontario, Trout Creek; on Betulalutea,T?,24.11i.1958;inSoo.2dd,l9,29.vi.l946, Bethel,Maine(A.F.B.) Bror-ocy. Most of the following details have been summarized from the biological study of the species by Lindquist (1962). Egg. Laid on under surface of leaf. In heavy infestations over fifty eggs may be deposited on a single leaf. Mine. The blotch is more or less retangular and is macle between two lateral leaf veins. The frass is deposited in circular patches. Larva. Head pale yellowish; gfound colour of body whitish with integument translucent. Cocoon. Yellow-brown darkening with age. The species overwinters as a cocooned larva in the surface soil and litter. Voltinism' According to Lindquist there is only one generation per year in Ontario. A chart of the seasonal distribution of the stages of the species suggests that the adults are present from late June to late July. However, some of the paratypes were collected in early June and one on 1 August. The specimen from Betula Lutea that emerged in March was probably "forced".

Knv: the cLEMENSELLA GRoUP l. Forewings each marked with single fascia. Male without crown of thorn-like spines near phallotremeof aedeagus .."' 2 - Forewings marked with four small, silver patches. Male with crown of thorn-like spines near phallotreme of aedeagus . . . quadrinotata (p 95) 2. Fascia of forewings distinct, shining silver . . ' rubifulieLLa (p. 9O) - Fascia of forewings nor distinct but fbrmed by the union of two patches with dull or only weak reflections ...... '..3 3. Forewings mottled, fascia dutl. Female lacking colliculum . '.. ulmella (p 9l) - Forewings not mottled, fascia weakly reflecting. Female with colliculum present . ' . 4 4. Malewiihscale-likeprocesses,arisingfrominneraspectofvalves,dividedintoshaftandflat digitate head. Hindwings with chitinised plate on dorsal, costal margin . platanella tp. 8?) - l.lite with hair-like processes arising from inner aspect of valves. Hindwings with dorsal, costal margin lacking chitinised plate ...... clemensella (p 86)

The clslrpNSELLA GRoUP The following species have been separated as a species-group within the genus Ectoedemia. Collectively they differ from populella in the following ways. The maxillary palps, although well developed, are not as robust and the eye-caps, (in relation iolt're treaA-capJule), are slightly larger. The antennae are approximately half the length of the wings. On the fringe of the forewing there is a distinct band formed by scales which are the ground colour of the wing. Further, all the species have wing markings in the form of fasciae or patches. The spurs on the tibiae of the hindlegs are not situated below the middle. In the male genitalia the vinculum is not as extensive. A11 the members of the clemensella group mine entirely within the leaf blade. Although argyropeza, canutus and lindquisli share a number of the above characters with clemensella and its close relatives, they are not included in the species-group for the reasons given under their respective discussions.

Ectoedemia clemensella (Chambers) comb. n. Nepticula clemenseIla Chambers, 1873:125 Nepticula clemensella Chambers; Chambers ln Hayden, 1878: 157 (listed) Nepticula clemensella Chambers; Dyar, 1903: 545 (listed) Nepticula clemensella Chambers; Braun, 1917b 188 N e p tic ula r: leme ns e lla Chambers; Braun lrz Forbes, 1923: 93 WILKINSON AND SCOBLE: THE NbPIICULIDAB OT CANADA 87

DsscnIpttoN. External .features: d, ?. Head: palps whitish, lustrous; antennae buff to pale ochreous; tuft on front of head and vertex ochreous; eye-caps white, shining; collar pale ochreous. Thorax and abdomen purplish black with silvery reflections. Wings: ground colour of forewings purplish black with bronzy and purple reflections; two medial white patches weakly shining silver, one on costa, one on anal edge, usually meeting to form fascia. Legs grey, with pale ochreous patches, lustrous.

Wing measurements: 6 . 4.4-4.8 mm (3); ? . 4.4-4.6 mm (4).

Genitalia'. d. Figure 48; tegumen produced into tapering, tuberculate pseuduncus; saccus bilobed, each lobe pointed; gnathos with truncated medial member; valves broad, not reaching tip ofpseuduncus, with hair-like scales arising from inner aspects and each divided into three prongs; transtilla with horizontal bar wide; aedeagus long, narrow, widening slightly at base, with rasp-like, arcuate structure on vesica; anellus, a deeply divided plate, each division with tubercle posteriorly. 9. Figure 49; ductus bursae appoximately twice as long as apophyses; colliculum ring-shaped with associated denticulate pocket; bursa copulatrix with pair of elongate signa; anterror apophyses arcuate, broad basally; posterior apophyses more naffow. Host-plant: Platanus occidentalis (Sycamore). Mine: an upper surface ophistigmatonome comprising long linear tract terminating in small blotch.

Ftc. 48. Ectoedemia clemensella, d genitalia 88 MEMOIRS OF THE ENTOMOLOGICAL SOCIETY OF CANADA

Frc. 49. Ectoedemia clemensella. ? eenitalia. WILKINSON AND SCOBLE: lHE NEPTICULIDAE OF CANADA 89

DrecNosrs. E. clemensel/a differs from platanella in having a more strongly bilobed saccus, shorter broader valves, and much more weakly developed hair-like scales arising from them. The species has a smaller wing span thanplatanella. The mine of clemensella is a very long, straight linear tract which broadens suddenly into a terminal blotch, whereas in plcttanella the linear mine is very short and the blotch massrve. DrsrnrsurroN. Canada: Ontario. U.S.A.: Ohio. Marvland. Pennsvlvania (Cle- mens), Kentucky (Chambers). SpsctnsNs ExlrttNEp. I lectotype; inMCZ: U.S.A.: Kentucky; (Chambers); genitalia, CNC slideNo. 3490. Paralectotype: I ex, "Peak. Acad.," "Type 14955".In MCZ, Other specimens: CANADA: Ontario, St. Williams'. onPlatanus occidentalis; 3dd, 299; l6-26.ii.1960 (Freeman and Lewis); in CNC. Lake Erie, St. Williams: on Platanus; ld, 20.iii.1962; in Soo. U.S.A.: Ohio, Cincinnati: on Sycamore; ld, 1 3.vii. 19 16, 2 9 9, 25,26.v1i.1912, 2 6, 1.vni.1902, |d, 25.vii. 1908, l ?, 8.vii. 1912, 1d, l9 ,26,27.vi.l916, in Phila. Maryland, Plummer's Island; I ex, v.1906 (Busck); in USNM. Brorocv. Mine. Comprises a very long linear tract which suddenly terminates in a blotch. The frass-line, central in the ophionome, is absent in a large part of the blotch. Pl. 6.3. Voltinism. Trivoltine. Ectoedemia platanella (Clemens) comb. n. Selected references: Nepticula platanella Clemens, l86l: 83 Nepticula platanella Clemens; Clemens, 1862: 133 Nepticula platanella Clemens; Clemens, 1865: 146 Nepticula platanella Clemens; Chambers, 1873: 125 Nepticula platanella Clemens; Chambers in Hayden, 1878: 158 Nepticula platanella Clemens; Dyar, 1903: 546 Nepticula platanella Clemens; Busck, l9O3: 209 Nepticula platanella Clemens; Braun, l9l7b: 181 Nepticula platanella Clemensl Braun in Forbes, 1923: 92 Nepticula platanella Clemens; McDunnough, 1939: 107 [No. 9760] Nepticula maximella Chambers, 1813 126 (syn. by Braun, l9l7b: 181) Nepticula maximella Chambers; Chambers in Hayden, 1878: 158 Nepticula maximella Chambers; Dyar, 1903: 546 There is a single male specimen of platanella in the CNC, the genitalia of which had already been mounted. The slide is much distorted but can be distinguished from clemensella in the following ways: the scale-like processes that arise from the inner aspect of the valves are divided into a shaft and flat digitate head. In addition, the valves are longer and more narrow and the saccus is only weakly bilobed. Although the external features are very similar, platanella has a chitinized plate on the upper surface of the costal margin of the hindwing extending from the base nearly to the middle. The mine is an ophistigmatonome, but the linear part is very small and the blotch large. (See Pl. 6.4.) A full description and diagnosis of the species based on comprehensive material from the U.S.A. is in preparation, but the following information is also pertinent here. Chambers (1873: 126) described adults which he named maximella and associated them with a mine earlier described by Clemens (1862b: 149) as sycamore miner No. 3 90 MEMorRs oF THB tsN l oMOLmrcAL socIETy oF CANADA and thought to be the product of a different species from platanella. In spite of this Braun (1917b: 187) synonymized maximella withplatanella blt gave no reasons. The mine of maximella is characteristic in the quadrate shape of the terminal blotch as opposed to the round blotch of platanella. The male syntype of maximella does not have the bifurcate anellar process and the so markedly bilobed saccus of platanella. On the basis of limited material (1d, l9) we think that maximella is just as referable to clemensella and is more probably a distinct species. However, we will not increase nomenclatorial complexity until we can be sure of its status. Thus here maximella isleft in synonymy with platanella. Host-plant: Quercus rubra. The usual food-plant for platany'lla is Platanus occidentalis: if the label data are correct this is an interesting addition. SpsctN,IsNs ExeuINr,o: CANADA: Ontario, Normandale: on Quercus rubra l6 , 16.vi.1962. (Freeman and Lewis.) Syntypes of maximella: l6 , "Nepticula maximella Cham."; Type No. 525 USNM; slide No. CNC 3477; 19, "Cham. Typ""; slide No. CNC 3502: in USNM.

Ectoedemia rubifuliella (Clemens) comb. n. Nepticula rubifoliella Clemens, 1860 214 Nepticula rubfoliella Clemens; Chambers ln Hayden, 1878: 158 (listed) Nepticula rubfoliella Clemens; Dyar, 1903: 547 (listed) Nepticula rubifoliella Clemens; Busck, 1903: 208 Nepticula rubfoliella Clemens; Braun, l9Ilb: 183 Nepticula rubifoliellct Clemens; Braun ln Forbes, 1923: 91 DnscruprroN. External features: 6 , I . Head: palps whitish, lustrous; antennae purplish grey; eye-caps shining white; tuft on front of head and vertex ochreous; collar off-white to pale ochreous. Thorax and abdomen brownish black. Wings: ground colour of forewings brownish black with single, shining silver, medial fascia; fringe terminally broad and rounded, heavily irrorate with dark brown scales. Legs grey, lustrous paling at tarsi. Pl. 10.5. Wing measurements: 6 . 4.4 mm (1); ? . 4.0-4.2 mm (4); neotype. 4.2 mm. Genitalia'. d. Figure 50; tegumen rounded, produced into short pseuduncus; saccus weakly bilobed; gnathos with pair of long curved horns tapering only slightly posteriorly, joined medially; valves reach base of pseuduncus, broader basally, curving to tip, rather squat; transtilla with lateral arms widely separated by long horizontal bar; aedeagus with pair of anellar projections; cornutus comprising a curved striate thickening. ?. Figure 5l; anal papillae with pair of setose lobes; ductus bursae less than twice length of apophyses; bursa copulatrix covered with small pectinations, with two large signa; anterior apophyses slightly curved and joined by long, thin sclerite; posterior apophyses long and thinning towards the setose lobes. H o s t- p lant : Rabu.s (Blackberry). M ine : tpper surface ophistigmatonome. DracNosrs. This is the smallest of the clemensella group. The shining silver fascia on the forewing and the overall shape of the male genitalia are its most distinguishing features. The female genitalia lack a colliculum on the ductus. DtscusstoN. The species was described from the mine by Clemens ( I 860) . As the mines were not preserved we designate one of Braun's specimens, from which the adults were first made known, as neotype. It bears the following data: "8558, Cincinnati, O. Annette F. Braun i.v.23.16". DrsrnrsurroN. Canada: Ontario, Quebec. U.S.A.: Ohio, Kentucky, Pennsyl- vania (Clemens). WILKINSON AND SCOBLE: THE NEFTICULIDAE OF CANADA 9l

Ftc. 50. Ectoedemia rubfoliella, d genitalia.

SppcIl{eNs ExeuINE,n. d neotype; in Phila: Ohio, Cincinnati; on Rubus; 23.v.1916 (Braun); genitalia, USNM slide No. 17339. Other specimens: data as neotype; 2I ? , 18.viri.1912, l.vi.1918 (Braun). Kentucky, Red Bird River; I d, l9.viii.1933 (Braun); in USNM. CANADA: Ontario, Simcoe; on Rubus; ld, 14.ii.1966 (Freeman). Quebec, Hull; on Rubus; 2? ?, 21.iii.1957 (Freeman and Lewis); I ex, 31.iii.1957 (Freeman); in CNC. Bror-ocy. Mine. In the linear tract the frass-line is broken and central. This part of the mine then widens into a rather elongate blotch with the frass irregularly dispersed. Pl. 7.1. Ectoedemict ulmella (Braun) comb. n. Nepticula ulmella Braun, 1912 87 Nepticula ulmella Braun; Braun, 1911b: 186 Nepticula ulmella Braun; Braun ln Forbes, 1923 92 DsscntprtoN. External .features: d. Head: palps off-white; antennae greyish; eye-caps whitish; tuft on front of head ochreous; tuft on vcrtex darker; collar as vertex. Thorax and abdomen greyish brown. Wings: covered by grey scales with purplish brown tips. giving a variously mottled ground colour of purplish brown and grey, with bronze reflections; fascia single, medial, with weak reflections, formed by union of costal and anal patches; fringe grey; hindwings with large basal patch ofbuffto pale ochreous androconia. Legs offwhite, lustrous. ? . As male but lacking androconia on hindwings. Wing measurements: 6 4.-5--5.6 mm (6): ?. -5.0-5.4 mm (5); lectotvDe.4.8 mm. Genitalia: d. Figure 52; tegumen extended into blunt papillate pseuduncus; saccus large, briobed; gnathos with medial pro.lection narrow, papitlate; valves narrow, separated by long, thin, 92 MEMOIRS OF THE ENTOMOLOGICAL SOCIETY OF CANADA

Frc. 51. Ectoedemia rubifoliella, ? genitalia. WILKINSON AND SCOBLE: THE NEFTICULIDAE OF CANADA 93

b

Frc. 52. Ectoedemia ulmella, d genitalia. transverse bar of transtilla rather narrow apically but developing a lobe basally; aedeagus gradually broadening anteriorly; vesica with denticulate cornuti and arcuate striate thickening; anellar projections-a pair of medial spines, arising from large basal processes, and pair of thickened elongate processes laterally. ?. Figure 53; ductus bursae rather short; bursa copulatrix large with oval signa; apophyses approximately length of ductus bursae, anteriores broader than outwardly curving posteriores. Host-plants. Ulmus Julva, U. americana (White Elm), U. racemosa (now:thomasi Sargent). Mine'. tpper surface'ophistigmatonome'. Dte.cNosrs. In the male the shape of the valves and their lobed projections distinguish ulmella from the other members of the group. The female genitalia lack a colliculum and in this resemble rubfoliella. However, the latter has a smaller alar expanse and a more brightly shining fascia. 94 MEMOIRS OF THE ENTOMOLOCICAL SOCIETY OF CANADA

Et'toedemia ulmella, I genitalia WILKINSON AND SCOBLE: THE NEPTICULIDAE OF CANADA 95 DrsrnrsurIoN. Canada: Ontario, Quebec. U.S.A.: Ohio (type locality), Pennsvlvania. Kentuckv (Braun). SpBcturNs ExArarNen. We designate as lectotype one of Braun's syntypes labelled "COTYPE": d, in Phila: U.S.A.: Ohio, Cincinnati; on Ulmus 8'578; l8.viii.l9l2 (Braun, coll. of Annefte F. Braun); USNM genitalia slide No. 16251. Paralectotype: Ohio, Clermont Co.; on Ulmus;19, l0.viii.l9l2 (Braun, Coll. of Annette F. Braun): in Phila. Other specimens: Ohio, Cincinnati; on(Jlmus; l9 no date, (Braun). 16,24'vi.l905; 19, 8.vi.l9l2; ld, 19, 3.vii.1907; I ex, 6.ix.1954 (Braun); in Phila. Pennsylvania, Pittsburg; | 6 ,23.v1i.1906 (En7el); New York, Cornell U. , on Ulmus , I d , Lot 451 Sub 618 (Murfetdt); in USNM. CANADA: Quebec, Kingsmere; onUlmus rubra (:fulva)' 26 6 , 39 9 , 2 ex, 12-30.vii.1956 (Lewis). Ontario, La Passe; on Ulmus americana; 19, 25.ii.1911, 16, 22.ri.1971 16, 26.i1i.1971 (Lewis). Overbrook; on Ulmus americana; I ex, 27.vii.1955 (Lewis); in CNC. Brorocy. Mine. Begins as a much contorted, frass-filled tract then expands into a false blotch in which the frass is deposited in both dark patches and lines. PI. 7.2'

Ectoedemia quadrinotata (Braun) comb. n. Nepticula quadrinotata Braun, l9l7b: 168 Nepticula quadrinotata Braun; Braun in Forbes; 1923: 86 DnscnrprroN. Externalfeatures'. 3 , I . Head: palps grey-brown with grey lustre; antennae dark greyish brown paling at tip; eye-caps white with silver reflections; tuft on front of head and vertex dark-ochreous; collar greyish brown. Thorax and abdomen greyish brown with grey lustre. Wings: ground colour of forewings dark greyish brown with greenish bronze reflections; four patches reflecting silver, two on costa at antemedial and subterminal positions, two on hind margin one postmedially, one basally; fringe grey. Legs grey, with buff patches, lustrous. Pl. 10.6. I(ing measurements: 6 . 4.0-5.2 mm (8); 9 . 4.0-5.0 mm (5); holotype' 4.8 mm. Genitaliai d. Figure 54; tegumen bluntly extended into short tuberculate pseuduncus; saccus weakly bilobed; gnathos comprising a pair of stout lateral horns fusing at bases to form a very thick, short projection; valves reach beyond pseuduncus, curved apically and tapering to blunt points; transtilla with lateral bars extended into long projections anteriorly; aedeagus long, slightly narrowing anteriorly, with a crown of thorn-like spines, surrounded by longer spines and processes, comprising anellar projections. ?. Figure 55; anal papillae elongate; ductus bursae extending beyond apophyses, with colliculum lacking associated denticulate pocket; bursa copulatrix covered with pectinations, signa elongate; anterior apophyses broad, arcuate, fusing to form basal bar; posterior apophyses sinuous, broadening basally, appearing to arise from basal plate. Host-plantst Carpinus caroliniana (Hornbeam), Corylus americana (Hazel), Ostrya virginiana (Ironwood), Betula alleghaniensis (now:8. lutea) (Yellow Birch). Mine: upper surface; beginning as ophionome later broadening. Dr.LcNosrs. The wing markings, the characteristic gnathos and anellus of the male and the long anal papillae of the female differentiate this species from all the other Ectoedemia species considered in this revision. DlstnrsutroN. Canada: Ontario. U.S.A.: Ohio (type locality Cincinnati). Arkansas, Kentucky (Braun). SprcrurNs ExeuINBn: I holotype; in Phila: U.S.A.: Ohio, Cincinnati; on Carpinus, B.538; 28.v.1914 (Braun, Braun Coll.); genitalia slide No. 17326. 96 MEMOIRS OF THE ENTOMOLOGICAL SOCIETY OF CANADA

FHry

Frc. 54. Ectoedemia quadrinotata, d genitalia

Paratypes: data as holotype; 26 6, 19, 27-30.v.1914 (Braun). Sugar grove; on Corylus; 1d, 1 9, Lvi.l9l5 (Braun). Other specimens: U.S.A.: Ohio, Cincinnati; on Carpinus; ld, 30.v.1914, 19, 31.v.1917, 26 6 , 19,2,9.vi.1917 (Braun); Ark., Washington Co., 19, Devils Den St. Pk., 11.v.66 (Hodges)1in USNM. CANADA: Ontario, Sparrow Lake; 8d d, 19, l2.vii.l926 (Braun), in USNM. Severn; on Ironwood; 26 6, 19, l6-20.vi.1925 (McDunnougft); in CNC. Trout Creek on Betula lutea;29 9, 1g.iii., 3.iv.1958; in Soo. Bror-ocy. Mine. The mine begins as a narrow, linear tract extending along the mid-rib or between two leaf veins and broadens so as to fill the area between them. In the late stages the mine is virtually a blotch as it widens still further. The frass is dark, initially comprising a broken line then becoming less compact and later diffuse. Pl. 7 .3. WLKINSON AND SCOBLE: THE NEPTTCI-LTDAE OF CANADA 97

Frc. 55. Ectoedemia quadrinotata, ? genitalia. 98 MEMoTRS oF THE ENToMoLocrcAL socIETy oF cANADA GENUS OBRUSSA Braun Obrussa Braun, 1915: 196. Type-species by original designation and monotypy Nepticula ochrefasciella Chambers, 1873 128. (ochrefasciella misspelt 'ochrifus- ciella' by Braun, 1915) Obrussa Braun; Braun, 1911b:200 Obrussa Braun; Braun ir Forbes, 1923l. 82 Etainia Beirne, 1945: 208 syn. n. Type-species by original designation. Lyonetia sericopeza Zeller, 1839 215 Etainia Beirne; Borkowski, 1912a: 692 Trifurcula Zeller;. sensa Johansson, l97l:241

DnscruprroN. External featuresi d. Head: palps extend beyond labrum; antennae approximately four-fifths of wing length. Wings. Forewing: venation; d as in Fig. 4; M and Cu fused at base before M diverges to meet Rs and later arises as Ml and M2; R4 and R5 separate terminally. Underside with brightly coloured androconia forming large patch, with streak, scaleless except for single row of scales, running longitudinally. Hindwings: venation with Rs and M each represented by single branch fusing half way along wing to form common trunk; ground colour light brownish grey. Legs: hindlegs with proximal pair of tibial spurs below middle of tibiae. 9. As male except: androconia absent. Genitalia: d. Vinculum ring-shaped; tegumen extended into pseuduncus; saccus rounded; valves each with characteristic, long, spine-like process arising from outer aspect and extending across capsule; anellus with well developed projections; vesica of aedeagus with several thickenings. ?. Ductus bursae with antrum and other sclerotizations; bursa copulatrix with pair of signa comprising reticulum of small "cells".

DracNosIs. Differs from Ectoedemia, Stigmella, Glaucolepis andTrifurcula as iI has a large scent-scale patch on the ventral surface of the forewings of the male. In the forewing of Obrussa the branches arising from Rs are R4, R5, Ml , andM2;inStigmella R4*5 and M, and inEctoedemia R4, R5, and M. lnObrussa Cu continues as a separate vein after M branches to Rs, whereasinGlaucolepis it does not. The separation of Rl and R2+3 from R4 and R5 on different trunks in the forewing, and the presence of l|dl+2 and M3 in the hindwing of Trifurcala distinguishes this genus from Obrussa. The long spine-like processes associated with the valves and the shape of the valves are characteristic of Obrussa.In the female the presence of an antrum is diagnostic as is the habit of the larvae in mining the keys of sycamore. In addition Obrussa differs from Stigmella and Glaucolepls in having signa reticulata on the bursa of the female and in the smaller size of the aedeagus in the male. It is further distinguished from Stigmella by its ring-shaped vinculum and the presence of a pseuduncus.

KBy: OsR.ussn Dorsal surface of forewing with single, broad, antemedial fascia otherwise unmarked. Male with ochreous patch of scales on ventral surface of forewing- and dorsal surface of hindwing. Female genitalia without V-shaped anal plate ... ochrefasciella (p.99) Dorsal surface of forewing with broad antemedial, irregularly defined fascia, and three patches. Male with blue-black patch of scales on ventral surface of forewing and dorsal surface of hindwing. Female genitalia with V-shaped anal plate . . . . . sericopeza (p. lol) WILKINSON AND SCOBLE: THE NEPTICULIDAE OF CANADA 99

O br us s a o c hrefas c i e lla (Chambers) Nepticula ochrefasciella Chambers, l8'l 3: 128 Nepticula ochrefasciella Chambers; Chambers in Hayden, 1878: 158 (listed) Nepticula ochrefasciella Chambers; Dyar, 1903: 546 (listed) Obrussa ochrefasciella (Chambers) Braun, 1915: 196 Obrussa ochrefasciella (Chambers); Braun, 19llb: 201 Obrussa ochrefasciella (Chambers); Braun in Forbes, 1923 82 DnscnrprroN. External features'. d. Head: palps off-white; antennae grey to pale ochreous; eye-caps white, shining whitel tuft on front ofhead and vertex ochreous. Thorax and abdomen purplish grey. Wings: ground colour of forewings purplish black, irrorate with pale grey, with weak bronzy reflections and single, broad, weakty shining, antemedial fascia; ventral surface with ochreous androconial patch; hindwings with basat patch ofpale ochreous androconia dorsally. Legs grey, with pale ochreous patches lustrous. Pl. 10.7. 9. As male, but without androconia. Wing measurements'. 6 . 6.2-7.4 mm (5); ? . 5.8-'7.4 mm (8). Genitalia: d. Figure 56; vinculum with large ventral plate and medial U-shaped emargination; saccus rounded; pseuduncus papillate, truncated; gnathos W-shaped, broad at base;

a '1 t' 3-

Frc. 56. Obrussa ochrefetsciella, d genitalia. 100 MEMOIRS OF THE ENTOMOLOGICAL SOCIETY OF CANADA

l- r,,, 5

Ftc. 57. Obrussa ochrefasciella, ? genitalia. WTLKINSON AND TOBLE: THE NEPTICULIDAE OF CANADA IOI valves blunt, reaching gnathos, with hair-like threads arising posteriorly; associated spine-like process attached near middle of outer edge serrate; transtilla wide, lateral arms very short with short processes; anellus with two pairs of stout spine-like projections. ?. Figure 57; ovipositor lobes blunt; ductus bursae long with unadorned narrow accessory sac; sclerotizations of ductus: antrum comprising pair of mandible-like, well sclerotized plates (see figure) point at which ductus enlarges into bursa copulatrix with numerous pectinate thickenings; bursa copulatrix relatively small with signa reticulata oval. Apophyses short, both pairs widening at base, anterior apophyses shorter. Host-plant: Acer sar:charum (Sugar Maple).

DI.q.cxosts. O. ochrefasciella may be distinguished from sericopeza in the colour and markings of the forewings and the ochreous rather than purplish black scaling of the ventral surface. The male genitalia are diagnosed by the truncated pseuduncus, the W-shaped gnathos, the shape of the valves and the serrate nature of the associated spine-like processes, and the form of the transtilla. The female genitalia have a better developed antrum and lack the invaginated anal plate of sericopeza. The genitalia have been checked against Dr. Davis' (USNM) type drawings.

DIsrntsutIoN. Canada: Quebec, Ontario. U.S.A.: Ohio, Pennsylvania, New York, Illinois.

SpecurlENs ExnurNeo. CANADA: Quebec, Knowlton; l9, l.vii.l929 (McDunnough); in CNC. Ontario, Desboro; on Acer saccharum; 1d, 29 ? , 26.vi.1967; in Soo. U.S.A.: Ohio, Cincinnati; 19, 23.vi.1947:- 16, 11.vi.1912 (Braun);inPhila.Pennsylvania,AllegroCo.,OakStation, 19,28.vi.1908; 16,299, l1.vi.l9l1(Marloj!). Pittsburgh; 46 6,29?,12.vi.-24.vii.1906 (Engel). New York, Ithaca; 1d, 30.vi.1931; 19, 9.vii.1924. lllinois, Putnam Co., 19, 13.vti.1942 (Glenn). ? New Hampshire; ld, xi.l929 (Busck). Maryland, Plummers Is., 19, 28. vii. 1962 (Hodges) (Photo).

Brolocv. We have no rearing records, mines, nor references about the life of the larva. However, it is strongly suspected thatochrefasciella makes a carponome inAcer keys, like sericopeza. Braun (1917b:. 201) stated that the early stages of the moth were entirely unknown: she may never have considered the possibility of the genus being a fruit miner.

Obrussa sericopeza (Zeller) comb. n. Lyonetia sericopeza Zeller, 1839: 215 Nepticula sericopeTa (Zeller) Zeller 1848: 325 N e pticula s ericopeza (Zeller) ; Petersen 1930:' 7 5 Nepticula sericopeza (Zeller); Hering 1951:22,44 (mining habit) Nepticula sericopeza (Zeller); Drrighia l9l 4: 213 Etainia sericopeza (Zeller) Beirne 1945: 208 Etainia sericopeza (Zeller); Borkowski 1972: 698 (Figure of wing venation) Trifurcula (Etainia) sericopeza (Zeller) Johansson 1971 246 Stigmella sericopeza (Zeller) Borkowski 1910: 543 (with photograph of imagine)

DlscnrptIoN. External features: d. Head: palps off-white; antennae pale ochreous; eye-caps white with white reflections; tuft on front of head creamy yellow, tuft on vertex yellow; collar cream. Thorax and abdomen dark grey, lustrous. Wings: ground colour of forewings very dark grey, irrorate with pale grey and with bronzy reflections; fascia single, rather irregular, broad, creamy white, antemedial, three creamv white patches with silver reflections, one postmedial on costa, one subterminal on anal edge, one basal; ventral surface with blue-black 102 ffiMorRS oF EN.roMOLoclcAL socrETy oF cANADA 'HE androconial patch; hindwings with basal patch of purplish black androconia dorsally. Legs grey, with pale ochreous patches, lustrous. Pl. 10.8. L As male, but androconial patches absent, Iling measurements: 6.5.8-7.4 mm (26); 9.6.2-7.4 mm (15). Genitalia'. d. Figure 58; vinculum with large, rounded ventral plate and medial U-shaped emargination, tegumen extended into long tapering pseuduncus; saccus rounded, not bilobed, gnathos wide, U-shaped; valves triangular, gradually tapering to apex extending to point just short of tip of pseuduncus; associated spine-like processes arising from outer edges at base of valves, not serrate; transtilla with long extension of lateral arms; anellus with two pairs of stout projections. ? . Figure 59; ovipositor wide; anal plate comprising a bifid V-shaped plate, the base of which is upturned; ductus bursae long with unadorned accessory sac; sclerotizations of ductus: antrum comprising pair of sclerotized plates; wall of lower third of ductus with longitudinal rows of pectinations; bursa copulatrix with signa reticulata pear-shaped; anterior apophyses broad at base, narrow, arcuate at tip; posterior apophyses narrow broadening into a pair of gently rounded lobes basally. Host-pLant: Acer platanoides (Norway Maple). Mine: in the fruit, forming a carponome in the parenchyma. DrncNosIs. Differs from ochrefasciella in wing markings, in the longer, tapering pseuduncus and U-shaped rather than W-shaped gnathos. In the female the invaginated anal plate is characteristic. See also ochrefasciella diagnosis. DrscussIoN. This species is widely distributed, occurring in many parts of the Holarctic. As there must be several barriers between populations the species is likely to be polytopic with genetic variation not being revealed in the phenotypes. DIstnrsurroN. Found in the Nearctic and the Palaearctic. SpEcrlmNs ExaurNpn. I holotype in BM(NH), bearing the following data: "Zeller Coll., Walsingham Collection, l9lo-427 .101340. Nepticula sericopeza

Zell.type.Teste Wlsm. " . Other specimens: CANADA: Ontario, Toronto; onAcer keys; 17dd, ll9I I ex, 7-l3.vii.l96} (Freeman and Lewis). Lake Erie; Port Dover; 566,299,24- 29.v1i.1966; in CNC. Collingwood; onAcer platanoides; ld, l6.viii.1967; in Soo. U.S.A.: New York, Ithaca, Six Mile Creek; 16,21.v.1959 (Hodges). Connecticut; on bark of Norway Maple; 299,22.v.1930; ld, I ex, 22.v.1929;6 ex,2,28.vi.1930 (B.T.R. Lab. Col.) 19, (Hofmann Coll.); in USNM. EUROPE: Estonia; on Acer platanoides;29 9, 16,26.vii. 1930 (Petersen); in Phila. , Frankfurt; onAcer; 16, l9 ,24,30.vi.1865 (Schmid, Stainton Coll.); in BM(NH). Bror-ocv. According to Hering the mine has a wall of frass along the edges.

GENUS GLAUCOLEPIS BTaw Glaucolepis Braun, 1917b:201. Type-species by original designation and monotypy Nepticula saccharella Braun, l9l2 91 Glaucolepis Braun; Braun in Forbes, 1923: 82

DrecNosrs. In the forewingGlaucolepis has R4, R5, Ml, and M2 arising on the same limb, and M+Cu is fused basally. By contrast Stigmella has R4 and R5 fused and M represented by a single branch and is not fused with Cu at all. Ectoedemia also has M as a single branch, and is further differentiated from Glaucolepis by having Cu continuing as a long branch after M has diverged from M*Cu. In lacking this feature WILKINSON AND SCOBLE: THE NEPTICULIDAE OF CANADA 103

b

a !.',. 5

Frc. 58. Obrussa sericopez.a, d genitalia. 104 MEMOIRS OF THE ENTOMOLOGICAL SOCIETY OF CANADA

4.'n 5

Frc. 59. Obrussa sericopeTa, ? genitalia. WILKINSON AND SCOBLE 105

Glaucolepis may be distinguished fromObrussa. The separation of R1 and R2*3, from R4 and R5 on different trunks in the forewing inTrifurcala distinguish this genus from Glaucolepis. The presence of Ml and M2 in the hindwing further distinguishes Glaucolepis from Stigmella, Ectoedemia, and Obrussa. In the male the aedeagus is very much longer relative to the genital capsule compared with the other genera. The female genitalia lack the characteristic sclerotized adornments on the ductus bursae found in Ectoedemia and Obrussa and the signa on the bursa copulatrix are not reticulate. The very long linear mine of Glaucolepis is also diagnostic.

DtscussroN. Braun (l9l7b: 2Ol) erected this monotypic genus mainly on the basis of wing venation. The genitalia are also characteristic. The venation is more complex than any of the other genera treated here. The presence of a cross-vein together with the ring-shaped vinculum indicates a closer relationship to Ectoedemia and Obrussa than to Stigmella. The leaf-mining habit in Stigmella and Glaucolepis has probably arisen independently.

G lauco le p is s ac chare lla (Braun)

Nepticula saccharella Braun, l9l2l. 9'7 Glaucolepis saccharella (Braun) Braun, 19llb: 20I Glaucolepis saccharella (Braun); Braun ln Forbes, 1923:82

DescruprroN. External features: 6 . Head: palps extend beyond labrum, relatively long, whitish, lustrous; antennae extend just under half length of wings, purplish black; front of head and vertex brownish purple; eye-caps brownish purple with purple reflections. Thorax and abdomen purplish brown with silver reflections. Wings. Venation: as in figure 5; forewing with Cu*M fused, meeting Rs; R4 and R5 separate terminally, M divided apically into Ml and M2; hindwing ovate; Ml and M2 united half way along wing to form common trunk at basal half of wing. Ground colour of forewings purplish black with reddish bronze reflections, with poorly defined basal to sub-basal, shining silver patch; single postmedial fascia shining silver; fringe grey, irrorate with scales of forewing ground colour which form band across terminal cilia. Ground colour of hindwings greyish brown with oval patch of ochreous androconia proximally on dorsal surface; fringe grey. Legs grey with ochreous patches, lustrous. Pl. 10.9. 9. As male except: eye-caps with only a few brownish purple scales confined to distal edge. Wings: hindwing not ovate; ground colour grey; lacking androconial scale-patch. Wing measurements'. 6 . 4.2-4.4 mm (3); ?. 4.0-4.8 mm (3); lectotype.4.0 mm. Genitalia'. d. Figure 60; vinculum ring-shaped, ventral plate not large; tegumen extended into pseuduncus; saccus large, weakly bilobed; gnathos comprising pair of lateral arms which articulate with lateral arms of vinculum, meet and fuse to form inverted V-shaped medial process; valves more or less rectangular with hook-like style extending to pseuduncus; transtilla lacking horizontal bar but lateral arms present; aedeagus broad and long with long, hooked, spine-like processes extending nearly entire length, and with striate thickening. 9. Figure 61; anal papillae simple; ductus bursae very short, less than length of apophyses; bursa copulatrix long with pair of long signa comprising rows of characteristic pectinations; anterior apophyses broad basally; posterior apophyses more narrow. Host-plants: Acer saccharun (Sugar Maple), Acer rubrum (Red Maple), Quercus rubra. Mine: long, upper surface ophionome.

DrsrntnuttoN. CANADA: Ontario, Quebec. U.S.A.: Ohio (type locality). SpEctunNs ExaltrNno. 9 lectotype, designated from one of Braun's syntypes; in Phila: U.S.A.: Ohio, Cincinnati; onAcer saccharum B.406. "CoTYPE, Collection of Annette F. Braun"t senitalia. USNM slide No. 17351 . 106 MEMOIRS OF THE ENTOMOLOGICAL SOCIETY OF CANADA

Frc. 60. Glaucolepis saccharella, d genitalia.

Other specimens: U.S.A.: Ohio, Cincinnati; on Acer saccharum; I ex, l5.viii'1914 (Braun); in Phila. CANADA: Ontario, Ottawa; oiAcer saccharuml I d , 9.vii.1956, on Acer rubrum, I d, 9.vii.1956, I ? ,21 .vi.1956 (Lewis). Normandale onAcer rubrum; 1?, 10.vi.1962, Aylmer, Id, 26.1i.1960 (Freeman andLewis). Simcoe; onQuercus rubra; l6 ,23.Ii.1971 (Lewis). Quebec, Hull; 2? ?, 20.vi.1955 (Lewis). Bror-ocy. Mine. The larva often crosses the mid-rib of the leaf, sometimes about half way through its mining life, but also at quite an early stage. The frass is deposited at first as a dense but broken line and later it is more dispersed' Pr' 7 '4' WILKINSON AND SCOBLE: THE NEPTICULIDAE OF CANADA t01

FIc. 61. Glaucolepis saccharella,9 genitalia r08 MEMOIRS OF THE ENTOMOLOCICAL SOCIETY OF CANADA

GENERAL DISCUSSION AND SUMMARY In this work, species with similar genitalia are regarded as being closely related. Yet although the genitalia are invaluable they are not without their limitations. For example some species can only be distinguished from their closest relatives by external characters. Such situations have led to a number of problems in the taxonomy. These are discussed below. S. crataegfoliella andS. scintillans have always been separated at species level but their genitalia seem to be identical. Their diagnosis depends only on external markings which suggests subspecific ranking to be more appropriate. However, they occur in the same region so further biological work is necessary to establish the level of relationship. Only very limited material was available but it is also possible that a quantitative study of many specimens might reveal discontinuities in genitalia. For example, in, an examination of extensive material found the uncus width of two closely related species Choristoneura () to differ. The genitalia of quercipulchella andvariella also seem to be indistinguishable, but uinllke crataegfoliella and scintilLanr they are allopatric. We appreciate that this could make them subspecies, but feel that the differences in the external features are great enough for them to be called species. The same applies to procrastinella andalba. However, Johansson (1971) treated the European basigutella and cerricolella as subspecies of basigutella because their genitalia are the same and their distribution different. Small, but apparently constant differences in the male and female genitalia distinguish ro saefoliella pectocatena from the nominate subspecies. As the dissimilarity in these characters is very slight, and because pectocatena was collected north of rosaefoliella, implying two possible centres of distribution, the two are called subspecies. The possibility that they are seasonal variants was considered. Label data and rearing records show that the nominate subspecies emerged in spring and summer whrlst pectocatena emerged in summer and fall. Further records, particularly of pectocatena, would be useful although maintaining a colony would be the best way of checking for seasonal forms. In the absence of such information rosaefoliella and pectocatena are called subspecies. E. argyropeza downsi has darker forewings than its nominate subspecies, but whereas downsi occurs in Canada, the nominate subspecies is European. Again' as the difference between them is very small, and because the distribution is so different, they are regarded as being subspecies. The maintenance of consistency within the general framework of the text influenced our decisions on the problems discussed above. In the future it is intended that follow-up work on some of the problems will be undertaken. This would be biological and morphological. Further rearing records are needed for more detailed information on distribution. Of great value would be a detailed study of features used in mate recognition, one of the best criteria for deciding on the reproductive isolation of a species. A comparative study of the ultra structure of scent-scales from the wings and anal tufts would be particularly worthwhile. Although most of the taxonomic work on the Nepticulidae has been done on the adults it is the larval stages to which the name leaf-miners is given. The leaf-mining mode of life is polyphyletic being found in many families. Amongst the larvae there are several instances of convergence between leaf-mining families (Trhgfirdh l9l3; Jayewickreme 1940). For example, in nearly all cases the larvae are prognathous and the teeth on the mandibles have, in consequence, become modified in order to meet. Both the above authors divided the larvae into those which feed on the sap released by WILKINSON AND SCOBLE: THE NEPTICULIDAE OF CANADA I09 piercing the cell walls of the leaf (sap-feeders) and those which also eat the cell walls (tissue-feeders). The Nepticulidae are tissue-feeders but they are somewhat atypical of them in having a flattened head and body, features more characteristic of the sap-feeding forms. The extremely delicate nature of the larvae of the Nepticulidae imposes certain limitations on the mining habits. The mid-rib and other major leaf veins restrict the movement of mining larva which often avoid crossing these obstacles. Sometimes the larvae follow the serrations of the leaf, e.g. as in plates 1.1 and 2.1 , and frequently the mid-rib or a major vein is followed but not crossed (plates 3.2;3.3;4.1 and 4.4). Sometimes larvae consume all the tissue between two veins, thus forming a blotch, e.g. plates L2 andT .1. Blotch mines are often found in the non-gall-forming members of the genus Ectoedemia, so although making a blotch sometimes avoids the necessity of crossing major veins, this seems to be superimposed upon a phylogenetic habit. Hering (1951) believed that the gall-forming habit, although polyphyletic, was derived from leaf-mining. He cited a number of examples in which callus tissue is found in present day mines and suggested that this reflects a pre-adaptive route along which the larvae of early miners evolved. By eating the callus in order to clear the mine, they finally became dependent on the production of callus for food. The genus Ectoedemia (in the sense of this study) includes both leaf-mining and gall-forming species, and also species which are ecologically intermediate between the two. E. populella forms petiole galls and all members of the clemensella group are leaf-miners. E. argyropeza which represents the intermediate situation, mines firstly in the petiole and then moves into the leaf-blade. The relatives of E. argyropeza produce a slight thickening in the petiole. They do not appear to rely on this callus tissue for food (Hering) and so are not strictly gall-formers. The reduced wing venation suggests that Stigmella developed from Ectoedemia (Borkowski 1972), and so the evolution of the feeding habits can be explained in two ways. Either a leaf-mining Ectoedemia stock, like the clemensella group, could have given rise to the leaf-mining genus Stigmella on the one hand, and to the gall-formingEctoedemia populella on the other by way of an intermediate species l1ke argyropezd; or the gall-forming habit of -Ectoedemia is primitive and gave rise to leaf-mining through the intermediate species. The antiquity of the leaf-mining way of life in general is evident from Opler (1973), who published information on lower Eocene fossil leaf-mines (including Nepticulidae). Freeman (1965) records a mine of Gracillaria from a late Eocene leaf, and Hickey and Hodges (1975) report aPhyllocnistis species from the late early Eocene. Mining larvae leave evidence of their feeding habits on the host-plant. Hering likens this to leaving behind a "visiting-card". This implies that there are data of potential diagnostic value which can be used as a means of identification. The special characters available are: the species or genus of host-plant mined, the vertical extension of the mine, the horizontal extension of the mine and the way in which the frass is deposited. The value of these characters depends on either (a) members of the same taxon consistently making mines similar to each other but different from those made by other taxa or (b) different taxa, which make similar mines to each other, feeding on different hosts; l.e. having a sufficient degree of host-specificity. In general, related species of Nepticulidae feed on host-plants that are themselves related. However, a number of taxonomic errors have been made by using host data as a priori characters. Hering (after Voigt, 1932, Ztschr. Pflanzenkrankh 42: 313) subdivides the generally wider terms monophagy, oligophagy, and polyphagy in order to reflect, more sensitively, the degree of host-specificity found in miners. Not all these categories are I l0 MEMoIRS oF THE ENToMoLocrcAL smrEly ots .ANADA represented by the Canadian Nepticulidae, but we have essentially adopted the scheme. The terms used in the following discussion are defined as follows: Monophagy. Feeding on plants restricted to a single plant genus. The subdivisions of this term are not used as they depend on more detailed data of species of host-plant than is available to us. Oligophagy. First degree: Feeding on several but related host-plant genera of the same family. Second degree: Feeding on different but related famities of plants of the same order. Third degree: Feeding on plant genera of different but related orders. Disjunctive oligophagy. Consistent feeding on several plants belonging to different orders. Polyphagy. Indiscriminate feeding on plants belonging to different orders. Table V gives the host-plant data collated from the breeding records and label data of the North American species treated in this study. The supra-generic categories of the plants, given in Table Vl and in the following remarks, follow Cronquist (1968). Although the exact degree of host-specificity in the Canadian Nepticulidae is not known, the food-plant records available suggest monophagy. There are, however a number of exceptions. The usual host of pomivorella is Malus but the species has also been reared from Crataegus'. this suggests first degree oligophagy. S. pomivorella is the only species of the crataegfoliella group that feeds on plants from two different, albeit closely related, families. It has not been recorded from Pyrus, which is the food-plant of the very closely related chalybeia, even where both pear and apple are found growing together, and even though both plants are themselves closely related. S. pomivorella and chalybeia probably diverged from a common, possibly oligophagous, ancestor and were then isolated on different plants, each subsequently becoming specialized to its own host. The only other Canadian species of Nepticulidae exhibiting first degree oligophagy is Ectoedemia quadrinotata. This species feeds on three different genera of the Betulaceae. All the specimens of bifasciella we examined have been reared from Prunus, except for a single male collected by J.A. Downes fromPopulus. Downes reared a large number of leaf-mining specimens from Populus and we are assured that the host data is correct. As the intensive collecting and rearing of miners from this tree failed to produce other specimens of bifasciella, Populus is not regarded as a regular host, and consequently bfosciella is regarded as monophagous. The fact thatPrunus andPopulus are in different plant subclasses (the Rosidae and Dillenidae respectively) indicates that Stigmelld species in general may be potentially capable of living on plants outside their usual host-range, even if the hosts are unrelated phylogenetically. Work by Buhr (quoted by Hering, 195 1: 313) in transferring mining larvae to hosts other than their own has shown that there is a variable, but sometimes considerable degree of tolerance in food-plant. A number of larvae introduced into the 'wrong' host survived and were reared through to the adult insect. That larvae are capable of feeding on a number of species of plants in the laboratory, even when confined to one species of host in the field, has been shown by Downey (1962) in the case of Plebejus icarioides (Lycaenidae). This species fed on many species of lupin in the laboratory, but in the field 'preferred' lupin species with pilose leaves, because the female tends to oviposit on hirsute leaves. Although the food-plant difference is not as marked as between the aforementioned example of Populus and, Prunus, it illustrates the principle that the factors affecting food-plant choice in insects are notprimarily nutritional (Dethier 1952; Thorsteinson 1960; Ehrlich and Raven 1964, and others). Ecological criteria such as host-plant distribution and WlLKINSON AND SCOBLE: THE NEPIICULIDAE OF CANADA lll

Table V. Guide to host plants

Acer A. platanoides O. sericopeza A. rubrum G. saccharella A. saccharum O. ochrefasciella, G. saccharella Alnus sp. S. corylifoliella Betula B. alba S. corylifoliella B. alle ghaniensis (.now : lutea) E. Iindquisti, E- quadrinotata B. papyrifera E. lindquisti Carpinus C . caroliniana E. quadrinotata Carpinus sp. S. ostryaefoliella Carya sp. S. ostryaefoliella Castanea C. dentata S. castaneaefoliella Corylus C. americana E. quadrinotata Corylus sp. S. corylifoliella Crataegus C. mollis S. cratae gifoliella, S. scintillans Crataegus spp. S . pomivorella , S. stigmaciella Gaylussacia G. baccata S. corylifoliella Hamamelis H. virginiana S. corylifolielLa Juglans J. nigra S. juglandifoliella Malus spp. S. pomivorella, S. scinanella Myrica M. cerifera S. myricafoliella Myrica sp. S. ostryaefoliella Ostrya O. virginiana S. proc rastinella, E. quadrinotata Ostrya sp. S. ostryaefolieLla Oxycoccus O. palustris S. corylifoliella Physocarpus P. opulifolius S. corylifuliella Platanus P. occidentalis E. clemensella, E. platanella Populus P. balsamifera E. canutus P. x canadensis S. aromella, S. populetorum P, canescens S. aromelia, S. populetorum P. deltoides S. populetorum P. grandidentata E. argyropeza downesi, E. populella P . tremula E. ar gyropez,a ar gyropeza (European) P. tremuloides E. argyropeza downesi, E. populella P. trichocarpa S. populetorum Populus sp. ?5. bifosciella Prunus P. cerasifera S. plagicolella (European) P. domestica S. plagicolella (European) P. nigra S. bifasciella, S. slingerlandella P. pensylvanica S . bifasciella , S. slingerlandella P. serotina S. bfasciella P. spinosa S. plagicolella (European) Pyrus P . communis S. chalybeia 1t2 MEMOIRS OF THE ENTOMOLOCICAL SOCIETY OF CANADA

Table V. \Concluded)

Quercus Q. agrifolia S. variella Q. albcr S. lat{asciella, S. saginella Q. macrocarpa S . latifasciella, S. saginella Q. palustris S. latifasciella Q. platanoides S. saginella Q. prinus S. saginella rubra Q. S . latifasciella, S. quercipulchella , E. platanella, G. saccharella 'Tan Bark Oak' Specimen/ Rhus R. aromatica S. intermedia R. typina S. intermedia Rosa spp. S. anomalella (European), S. rosaefoliella R. setigera S. rosaefolielLa R. virginiana S . rosaefoliella Rosa sp. Specimen a, specimen & Rabzs spp. S. rubifoliella Salir S. discolor S. fuscotibiella S. nigra S. fuscotibiella Ulmus U. americana E. ulmella U. thomasi E. ulmella Vaccinium sp. S. coryllbliella

Table VI

HOST-PLANT \ErTlcuqD4E Genus Family Order Subclass Stigmella The nosarrol-tELLA GRoup S. rosaefoliella Rosa Roseaceae Rosales Rosidae S. slingerlandella Prunus Roseaceae Rosales Rosidae (?Malus) S. fuscotibiella Salix Salicaceae Salicales Dilleniidae S. populetorum Populus Salicaceae Salicales Dilleniidae S. aromella Populus Salicaceae Salicales Dilleniidae The cteratclpoLIELLA GRoup S. crataegifoliella Crataegus Roseaceae Rosales Rosidae S. scintillans Crataegus Roseaceae Rosales Rosidae S. pomivorella Malus, Roseaceae Rosales Rosidae Crataegus S. chalybeia Pyrus Roseaceae Rosales Rosidae S. scinanella Malus Roseaceae Rosales Rosidae S. stigmaciella Crataegus Roseaceae Rosales Rosidae The s,qctNEr-re cnoup S. saginella Quercus Fagaceae Fagales Hamamelidae S. castaneaefoliella Castanea Fagaceae Fagales Hamamelidae S. latifasciella Quercus Fagaceae Fagales Hamamelidae WILKINSON AND SCOBLE: THE NEPTICULIDAE OF CANADA 113 Table VI. qConcludedl

The coryr-rrolrELLA cRoup S. corylifoliella Corylus Betulaceae Fagales Hamamelidae Betula Betulaceae Fagales Hamamelidae Alnus Betulaceae Fagales Hamamelidae HamameLis Hamamelidaceae Hamamelidales Hamamelidae Vaccinium Ericaceae Ericales Dilleniidae Gaylussacia Ericaceae Ericales Dilleniidae Orycoccus Ericaceae Ericales Dilleniidae Physocarpus Roseaceae Rosales Rosidae S. ostryaefoliella Ostrya Betulaceae Fagales Hamamelidae Carpinus Betulaceae Fagales Hamamelidae Carya Juglandaceae Juglandales Hamamelidae Myrica Myricaceae Myricales Hamamelidae S. juglandifoliella Juglans Juglandaceae Juglandales Hamamelidae The nrrescrEr-r-A GRoup S. bifasciella Prunus Roseaceae Rosales Rosidae ?Populus Salicaceae Salicales Dilleniidae S. intermedia Rhus Anacardiaceae Sapindales Rosidae S. quercipulchella Quercus Fagaceae Fagales Hamamelidae S. procrastinella Ostrya Betulaceae Fagales Hamamelidae

Ectoedemia E. populella Populus Salicaceae Salicales Dilleniidae The encynoprzA GRoup E. argyropeza Populus Salicaceae Salicales Dilleniidae E. canutus Populus Salicaceae Salicales Dilleniidae E. lindquisti Betula Betulaceae Fagales Hamamelidae The clBrraENsBI-LA GRoup E. clemensella Platanus Platanaceae Hamamelidales Hamamelidae E. platanella Platanus Platanaceae Hamamelidales Hamamelidae Quercus Fagaceae Fagales Hamamelidae E. rubifuliella Rubus Roseaceae Rosales Rosidae E" ulmella Ulmus Ulmaceae Urticales Hamamelidae E. quadrinotata Carpinus Betulaceae Fagales Hamamelidae Corylus Betulaceae Fagales Hamamelidae Ostra Betulaceae Fagales Hamamelidae Betula Betulaceae Fagales Hamamelidae

Obrussu O. sericopeza Acer Aceraceae Sapindales Dilleniidae

Glaucolepis G. saccharella Acer Aceraceae Sapindales Dilleniidae

growing season, the egg-laying habits of the female, and secondary plant substances (Fraenkel 1959), are regarded as being of first importance in food-plant preference. _ The species with the greatest host-plant range are corytifotiella andostryaefoliella. lh-ev are closely related to each other and to the apparently monophagous jugtan- dfoliella. Three species are newly synonymized with corylifulielta ti thii study. The 114 MEMoTRS oF THE ENTOMOLOGT.AL socIETy oF cANADA food-plants of those synonymized are different from each other and fromCorylus, the previously accepted host of corylfoliella. However, the genitalia seem to be the same and the external features fall within a tolerable level of variation. We have avoided using host-plant data as a priori characters because the exact degree of host-specificity in the family is not known and so feeding on different plants is not a reliable character for separating species. As there are clear differences between corylifoliella and ostryaefoliella it would be inconsistent to treat the synonyms as different species. It would also be inconsistent to call them species within the framework of what has been regarded as specific rank in this work. S. corylifoliella feeds on genera belonging to orders of the Hamamelidae, on Vaccinium and Gaylussacia which belong to the subclass Dillenidae and on Physocarpus, a member of the Rosidae. Apart from the possibility of this being disjunctive oligophagy, it is the only example of polyphagy in the Nepticulidae of which we are aware. Possibly an ecological factor explains this situation . Vaccinium is often found as a ground cover in deciduous woods where at least some of the other food-plants of corylfoliella occtr. If hosts co-exist in the same community then barriers of a distributional nature do not exist. This increases the chance of females ovipositing on the 'wrong' host. This, together with the possibility that the larvae may have a potentially wider food-plant tolerance than is usually exhibited (discussed above), may partly explain the host-range of corylfoliella. S. ostryaefoliella, at least, shows a degree of selectivity in its food choice. It feeds on genera which, according to Cronquist, represent three plant orders, viz: the Juglandales, the Fagales, and the Myricales. This is an example of third degree oligophagy as the plant orders mined by ostryaefoliella are related, all belonging to the subclass Hamamelidae. This close relationship is emphasized by the fact that the families Myricaceae and Juglandaceae have, at times, been associated in a single order. Both of them are allied to the Fagales. Host-plant data may provide evidence of the phylogenetic relationships within the corylfolietkt group. Of the three familes of plant mined by ostryaefoliella, it shares the Juglandaceae in common with juglandfoliella and the Betulaceae with corylfoliella. It is possible that the original hosts of ostryaefoliella belonged to the Myricaceae. From these the insect may have spread to plants ofthe Juglandaceae and the Betulaceae giving rise to juglandifuliella on the one hand and corylfoliella on the other. Nevertheless at this stage we agree with Downey (1962) that the "taxonomic value of food-plant data is thought to be only potential or confirmatory". If more sophisticated techniques e.g. comparative work on ultra-structure of scent-scales and other parts of the insect, and electrophoresis of blood proteins, distinguish moths feeding on different species of plant, then the degree of host-specificity in the Nepticulidae could be established. Apart from the above examples, monophagy is general in both the Nearctic and the Palaearctic Nepticulidae. Also, species that are closely related tend to feed on host-plants that are themselves taxonomically related. This is not at variance with the well documented principle of the prime importance of non-nutritional factors in the choice of host-plant. As Southwo od (1912) has pointed out, transfer to a new host will not only be affected by ecological factors of the plant, but also by the 'predilection' of the insect for the change. As this is a reflection of the biochemistries of both the new and the original host and the metabolism of the insect, the degree of predilection may vary in direct proportion of the botanical affinity of the two hosts. For example, Eastop (1973) states that most species of aphid aregg%o host-specific. He suggests that although these insects alight upon a large number of 'wrong' food-plants, only a few of them are 'captured'. Plants successfully colonized are usually taxonomically related. WILKINSON AND SCOBLE: THE NEPTICULIDAE OF CANADA 115 SUMMARY l. A revision of 38 taxa of the Nepticulidae of Canada, as represented in Canadian collections, has been carried out. This has been enhanced by the loan of material from the U.S.A. 2. Three European species have been included because of their importance in nomencla- ture. 3 . A case has been made for stabilizing the generic name as ,ttigr?? elLa and notNepticula . It is proposed that the family name be Nepticulidae, and not Stigmetlidae. 4. Genera are separable by venation and genitalia. 5. Species are usually separable on male genitalia and sometimes on female genitalia. The majority are figured. 6. Life histories, mine characteristics, and larval habits are given where known. 7. Tables illustrate former classifications in addition to the one used here. 8. A guide is provided to the host-plants and it has been shown that related species tend to feed on different but related plant species. 9. Clemens described and named some species only from the damage caused by the larvae. Some of the resulting problems were resolved by neotype designations. 10. Palaearctic and Nearctic species have been examined together for the first time. Changes in classification reconcile the different approaches of past workers 11. Stigmella comprises only leaf-blade miners. Ectoedemia, in this account is taken to include leaf-miners of the lamina as well as the petiole. Obrussa mines fruits of trees sycamore keys. The only species of Glaucolepis is a leaf-blade miner. - 12. The larvae of the Nepticulidae are tissue-feeders rather than sap-feeders, although the veins and mid-rib often restrict their movements. 13. Ectoedemia petiole feeders are not dependent on callus tissue and are not therefore true gall formers. 14. Most Nepticulidae are monophagous. The exceptions are discussed in the General Discussion. 15. Although mostStigmella species have different genitalia, some pairs do not. In these cases, however, wing markings and often size differ markedly. They are also allopatric. The taxonomic implications are discussed. 16. The female genitalia of Ectoedemia and Obrussa are shown to be characterizedby a pair of large signa in the form of a reticulum of'cells'. 1'7. There is evidence that stereoscan studies would be rewarding to reveal the patterns of fine sculptures on the inside surface of the bursa copulatrix in females. 18. Ectoedemia argyropeza, previously thought to be European, is shown to have a North American sub-species. It has been contused in the past with turbidella. I 9. A numerical analysis of the Nepticulidae has been carried out (not detailed here) which confirms our classification. 20. This work will be supplemented shortly by an account of those Nepticulidae so far only recorded from the U.S.A.

REFERENCES

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Braun, A.F. 1910. New species of Tineina from Califomia. Ent. News 2l(4): l7l-l'19. 1912. Notes on North American species of Nepticula with descriptions of new species. J. Cincinn. Soc. nat. Hist. 2l(3): 84- 101, I 3 figs. 1914. Notes on North American species of NepticuLa with descriptions of new species (Lspidoptera). Can. Ent. 46(24): 17-24,8 tigs. 1915. New genera and species of Tineina. lbid. 47(6): 188-197, 6 figs. 191'la. Observations on the pupal wings of Nepticula with comparative notes on other genera. Ann. ent. Soc. Am. l0(3):233-239, I pl. 1917b. Nepticulidae of North America. Trans. Am. ent. Soc. 43(2): 155-209, 4 pls. 1919. Wing structure of Lepidoptera and the phylogenetic and taxonomic value of certain persistent trichopterous characters. Ann. ent. Soc. Am. l2(4):349-366, I pl. 1923. Superfamily Family 5 Nepticulidae. pp. 79-98 in Forbes, W.T.M. (Ed.), Lepidoptera of New York and Neighbouring States. 10 figs. New York. 1924. ThefrenulumanditsretinaculumintheLepidoptera. Ann.ent.Soc.Am. lT(3):234-256. 1921 . New from Ontario. Can. Ent. 59(3): 56-59. 1930. Notes and new species of Microlepidoptera from the Mineral Springs region of Adams County, Ohio. Trans. Am. ent. Soc.56(1): 1-17. Busck, A. 1900. New species of moths of the Superfamily Tineina from Florida. Proc . U .5. natn. Mus. 23:225-254, I pl. 1901 . Nepticulapomivorella Packard; aliasMicropteryrpomivorella Packard. Can.Ent.33:52. 1903. Notes on Brackenridge Clemens' types ofTineina. Proc. ent. Soc. Wash. 5(3): l8l-22l. lgoi. New American Tineina. Ibid. 8: 86-99, 11 figs. 1913. Two Microlepidoptera injurious to Chestnut. Ibid. l5(3): 102-104, I fig. 1914. On the classification of the Microlepidoptera; and Descriptions of New Microlepidoptera of Forest Trees. Ibid. 16: 46-54, 143-150. 1931. On the female genitalia of the Microlepidoptera and their importance in the classification and determination of these moths. Bul/. Brooklyn ent. Soc. 26(5): 199-216, 5 pls. Busck, A. and C. Heinrich. 1921 . On the male genitalia of the Microlepidoptera and their systematic importance. Ibid. 23(6): 145-149,2 pls. Cain, A.J. 1954. Animal Species and their Evolution, ix + 190 pp. Hutchinson. Chambers, V.T. 1873. Micro-lepidoptera. Can. Ent. 5(7): 124-128. 1875. Tineina of the Centfal United States. Cincinn. Q.JI Sci. 2(2): 91-121. 1878. New Tineina from Texas: and Index to the described Tineina of the United States and Canada. In Hayden, F.V. (Ed.), Bull. U.S. geol. geogr. Surv. Territ. 4:19-1O6, 125-16'7. i879. Micro-lepidoptera. Can. Ent. 11(5): 89-93. 1880d. Descriptions of sorne new Tineina, with notes on a few old species. J. Cincinn. Soc. nat. Hist. 2(4: 179-194. 1880b. Psyche, Camb.3: 63-68. Clark, J.F. Gates. 1950. The date of "A list of North American Lepidoptera" by Harrison G. Dyar. Proc. ent. Soc. Wash. 52(6): 308. Clemens. B. 1860. Proc. Acad. natn. Sci. Philad. 12:203-221. 1861. Micro-lepidopterous larvae. Notes on a few species, the imagos of which are probably undescribed. Proc. ent. Soc. PhiLad. l(4):15-81. 1862. New American Microlepidoptera. Ibid. 1(5): 131-137. 1865. North American Micro-Lepidoptera. Ibid. 5: 133-14'7. Cochaux, P. 1969. A European leaf-miner (Stigmella (Nepticula) turbideLla) in Quebec. Bi-Mon. Can. Dep. Fish For. Res. Notes 25(2):12. Comstock, J.H. 1918. The Wings of Insects.xviii + 430 pp., 427 figs., 10 pls. Cronquist, A. 1968. The Evolution and Classification of Flowering Plants. x + 395 pp. Nelson. Crosby, C.R. l9ll. The Plum (Nepticula slingerlandella Kearfott). Bull. Cornell Univ. agric. Erp. Srn 308: 219-227, 14 figs. 1912. Notes on the life history of Nepticula slingerlandella Kearfott (Tineidae). Can. Ent. M(.1):25-27. DeGeer, C. 1152. Memoires pour servir a l'histoire des insectes l. Stockholm. pp.440-458, pls. 29-31. 1171. rbid. rr(i. Dethier, V.G. 1952. Evolution of feeding preferences in phytophagous insects. Evolution (Lancaster, Pa.) 8: 33-54. Downes, J.A. 1968. A nepticulid moth feeding at the leaf-nectaries of Poplar. Can. Ent. 100(10): 1078- 1079. Downey, J.C. 1962. Host-plant relations as data for classification. Syst. Zool. ll: 150-159, 2 fiss. WILKINSON AND SCOBLE: THE NEPTICULIDAE OF CANADA tt7 Dr:ighia, l. 1974. Neue Angaben iiber die Morphologie und Biologie der minierenden Raupen-Schddlinge der Knospen und Friichte yonAcer (Lepidoptera: Nepticulidae). Folia ent. hung. XXYll. Suppl., 213-219, 4 figs., 4 tables. Dyar, H.G. 1903. A list of North American Lepidoptera and key to the literature of this order of insects. Bzl/. U.S. natn. Mus. xix+723 pp. Washington. Eastop, V.F. 1973. Deductions from the present day host plants of aphids and related insects. pp. l5'7-178 irz van Emden, H.F. (Ed.), Insects/Plant Relationships. Symp. R. ent. Soc. Lond., No.6,4 figs. Ehrlich, P.R. and P.H. Raven. 1964. and plants: A study in co-evolution. Evolution (Lancaster. Pa.) 18: 586-608. Emmet, A.M. 1911. Notes on some British Nepticulidae. Entomologist's Rec. J. Var. E3:240-248. 1976. Nepticulidae. pp. 11l-267 ln Heath, J. (Ed.), The Moths and Butterflies of Great Britain, and Ireland, 1, figs. 56-73, maps 19-113, pis. 1-9, 11-12. Eyer, J.R. 1924. The comparative morphology of the male genitalia of the primitive Leprdoptera. Ann. ent. Soc. Am. 17:2'75-328. 14 ols. Fabricius, J.C. 1'7'75. Systema Eniomologiae. 832 pp. Fletcher, T.B. 1929. A list of the generic names and for Microlepidoptera. Mem. Dept. Agric. India. Calcutta ent. Ser. ll: ix+244 pp. Fraenkel, G. S. I 959. The raison d'etre of secondary plant substances. Science, N .Y . 129: 1466-1470, 1 fig. Freeman, T.N. 1953. The spruce budworm, Choristoneura fumiferana (Clem.) and an allied new species on pine (Lepidoptera:Tortricidae). Can. Ent. E5(4): l2l-12'7,22 figs. 1962. A new species of Nepticula v. Heyd. on Birch (Lepidoptera: Nepticulidae). Ibid. 94(5): 522-523,2 figs. 1965. A lepidopterous leaf-mine from the Tertiary Period. Ibid.97(10): 1069-1070, 2 figs. 1961 . A new species of Nepticula on Bur Oak in Ontario (Nepticulidae). J. Res. Lepid. 6(l): 19-21,2 figs. Frey, H. and Boll, J. 1878. Tineen aus Texas. Stettin. ent. Ztg 39:249-280. Gijze, J.A.E. 1783. Entomologische Beytrlge zu des Ritter Linn6 Swijlften Ausgabe des Natursystems. xx*178 pp. Leipzig. Hampson, G.F. 1918. Some small families of the lepidoptera which are not included in the key to the families in the catalogue of Lepidoptera Phalaenae, A list of the families and subfamilies of the Lepidoptera with their types and a key to the families.- Novit. zool. 25:366-394. Hering, E.M. 1951. Biology of the Leaf Miners, iv*420 pp., 2 pls., 180 figs. 's-Gravenhage. Herrich-Schdffer, G.A.W. 1855. Systematische Bearbeitung der Schmetterlinge von Europa 5. 394 pp. Heyden, C. von 1843. Amtlicher Bericht der Versammlung der Naturfoncher zu Mainz, p. 208. Hickey, L.J. and R.W. Hodges. 1915. Lepidopteran leaf mine from the Early Eocene Wind River Formation of northwestem Wyoming. Science, N.Y. 189: 718-720,2 ftgs. Hosie, R.C. 1969. Native Trees of Canada. 7th ed., 380 pp. Imms, A.D. 1951 . A General Textbook of Entomology, 9th (rev.) ed., x+886 pp., 609 figs. London' International Code of Zoological Nomenclature. l9g. Adopted by the XVth int. Congr. ZooL,2nded., xx+ 176 pp. London. Jayewickreme, S.H. \940. A comparative study of the larval morphology of leaf-mining Lepidoptera in Britain. Trans. R. ent. Sot:. Lond. 90(4):63-105, 12 figs. Johansson, R. 19'71. Notes on Nepticulidae (Lepidoptera). l. A revision of theNepticula ruficapitella group. Ent. scand. 2:241-262,52 frgs. Kearfott,W.D. 1908. NewNorthAmericanTortricidaeandTineina. JlN.Y.ent. Soc. 16(3):167-188, 1 pl. Kirby, W.F. 1897 . Lloyds Natural History. A handbook of the Order Lepidoptera. 5. xii 332 pp., 32 pls. Edward Lloyd, London. Klimesch, J. 1948. Zur Frage der verwandtschaflichen Beziehungen einiger Stigmella-Arten auf Grund des Baues des Mannl. Kopulationsapparates. (Lep., Stigmellidae). Z. wien. ent. Ges. 33:49-82,62 figs. 1951. Zur Kenntnis der Genitalmorphologie einiger Nepticula-Arten (Lep. Nepticulidae). Z. wien. ent. Ges. 36:4-9, 7 figs. 1953. Die europaischenTrifurcula-und Ectoedemia-Arten (Lep., Nepticulidae). Z. wien. ent. Ges. 38: 160-170, 19l-196,22figs. Kloet, G.S. andW.D. Hincks. 1972. ACheckListof Britishlnsects;Lepidoptera l1(2). 2nd(rev.)ed., viii+153 pp. Royal Ent. Soc. London. Klots, A.B. 1956. Lepidoptera. pp.97-l1l in Tuxen, S.L. (Ed.), Taxonomists glossary of Genitaliaof Insects. 12 figs. Copenhagen. I 18 MEMOIRS OF THE ENTOMOLOGICAL SOCIETY OF CANADA

Lindquist, O.H. 1962. A biological study of a new leaf-miner on birch, Nepticula tindquisti Freeman (Lepidoptera: Nepticulidae), in Ontario. Can. Ent.94(5):524-530, 9 figs. Lindquist, O.H. and A.A. Harnden. 1910. A biological study of Neptlcukt macrocnrpae (Lepidoptera: Nepticulidae) on oak in Ontario. Ibid. 102(10): 129O-1293, 4 figs. McDunnough, N.H. (misspelled McDonnough). 1911. On the nomenclature of the male genitalia in Lepidoptera. Can. Ent.43(6): 181-189, 3 figs. 1939. Check List of Lepidoptera of Canada and the United States of America. II. Microlepidop- tera. Mem. sth. Calif. Acad. Sci. 2(l). Opler, P. 19'73. Fossil lepidopterous leaf-mines demonstrate the age of some insect-plant relationships. Science, N.Y.l79: 1321-1322. 1 fie. Packard, A.S. 1870. New of little kriown injurious insects. Rep. Mass. Bd Agric. 17:231-238. 1889. Guide to the Study of Insects, 9th ed., xii+715 pp., 15 pls., 670 figs. New York and Boston. Petersen, W. 1930. Die Blattminierer-Gattungen Lithot:olletis und Nepticula (Lep.). Teil Il: Nepticula Z. Stettin. ent. Ztg 9l: l-82,3 pls., l6 figs. Prentice, R.M. et al. 1965. Forest Lepidoptera of Canada. Recorded by the Forest Insect Sur- vey. Microlepidoptera. 4: 546-84O, 181 figs. Schrank, F. von P. 1802. Fauna Boica. 2(2). 412 pp. Niimberg. Snodgrass, R.E 1935. Principles of Insect Morphology, lst ed., x*667 pp, 319 figs. New York and London. 1951. A revised interpretation of the external reproductive organs of male insects. Smithson. misc. Collns 135(.6). iii+60 pp., l5 figs. Baltimore. Southwood, T.R.E. 1972. The insectplant relationship an evolutionary perspective. In van Emden, H.F. (Ed.), Insect-Plant Relationships. Symp. R. ent.- Soc.lond. No. 6: 3-30. 4 figs. Spuler, A. 1910. Die Schmetterlinge Europas. 2, 525 pp.,239 figs. Stuttgarr. Stainton, H.T. 1854a. Insecta Britannica. Lepidoptera: Tineina. viii+313 pp., l0 pls. London. . in Gray, J.E. 1854b. List of British in the collection of the British Museum XVI: Lepidoptera. London. 199 pp. 1855. The Natural History ofthe Tineina. l. xii+338 pp., 8 pls. John van Voorst, London. Svensson, I. 1966. NewandconfusedspeciesofMicrolepidoptera. Opusc. Ent.3l(3): 183-202,4p1s. Thomteinson, A.J. 1960. Host selection in phytophagous insects. A. Rev. ent.5: 193-218. Trdgirdh, I. 1913. Contributions towards comparative morphology of the trophi of the lepidopterous leaf-miners. Ark. Zool.8(9): l-48, 67 figs. Tutt, J.W. 1899. A Natural History of the British Lepidoptera. 1: 1-560. Swan Sonnenschein, London. Walsingham, T. 1880. On some new and little known species of Tineidae. Ibid.11-93,2 pls. - (under Rt. Hon. Lord). 1907. Microlepidoptera ofTenerife. Proc. zool. Soc. Lond.910- - r028. Whalley,P.E.S. 1974. ScentdispersalmechanismsinthegenusStrlglinaGuenle,withadescriptionofa new species (Lepidoptera, Thyrididae). J. Ent. (8) 43(1): 121-128. Wilkinson, C. 1978. On the StigmellalNepticula: Lepidoptera controversy. Tijdschr. Ent. l2l(.2): 13-22, pls, l-2. Zeller, P.C. 1839. Versuch einer naturgembssen Eintheilung des Schaben, Tinea. Isis, Jena 32: 161 -220. 1848. Die Gattungen der mit Augendeckeln versehenen blattminierenden Schaben. Linn. ent. 3: 248-343. W1LKINSON AND SCOBLE: THE NEPTICULIDAE OF CANADA t19

on Prunus Pr-nrr 1 . Larval mines of rosaefoliella group. I , S. rosaefoliella on Rosa. 2, S . slingerlandella x 3, S. fuscotibielta oo Salix. 4, S. populetorum on Populus trichocarpa. 5, S. populetorum on Populus canaaenslS. t20 MEMOIRS OF THE ENTOMOLOGICAL SOCIETY OF CANADA

Plern 2. Larval mines of crataegifoliella group. 1, S. crataegifoliella on Crataegus. 2, S. scintillans on Crataegus. 3, S. scinanella on Malus. 4, S. stigmaciella on Crataegus. WILKINSON AND SCOBLE: THE NEPTICULTDAE OF CANADA t2l

Purr 3. Larval mines of saginella group. 1, S. saginella on Quercus alba.2, S. castaneaefoliella on Castanea denlata.3, S. latifasciella onQuercus palustis. 4, S. castaneaefoliella onCastanea dentata. t22 MEMOIRS OF THE ENTOMOLOGICAL SOCIETY OF CANADA

Pr-erE 4. Larval mines of corylfolielta group 1, S. corylifuliella on Corylus 2, S. corylifuliella on Vaccinium.3,S ostryaefoliella onCaryaovata.4,S juglandifoliella onJuglans. Larval mines of bifusciella group and quercipulchella group. I, S. bifasciella on Prunus pensylvanica, 2, S. intermedia onRirus ryphina. 3, S. quercipulchella onQuercus. 124 MEMOIRS OF THE ENTOMOLOCICAL SOCTETY OF CANADA

Prlrs 6. Larval mines of Ectoedemia spp. I , E. argyropeza on Populus tremuloides . 2, E. argyropeza on Populus tremuloides . 3, E. clemensella on Platanus occidentalis . 4, E. platanella on Platanus occidentalis . WTLKTNSON AND SCOBLE: THE NEPTICULIDAE OF CANADA r25

Prerr 7. Larvalmines of Ectoedemia spp. (continued)andGlaucolepls sp. l, E.rubfoliellaonRubus.2, E. ulmella onUlmus.3, E. quadrinotata ooCarpinus.4,G. saccharella onAcer rubrum. r26 MEMOIRS OF THE ENTOMOLOCICAL SOCIETY OF CANADA

Prrrn 8. Adult external features of Stigmetla spp. 1, S. slingerlandella. 2, S. fuscotibiella. 3, S. populetorum. 4, S. crataegifuliella. 5, S. scintillans. 6, S. pomivorella. 7 , S. scinanella. WILKINSON AND SCOBLE: THE NEPTICULIDAE OF CANADA r27

Purr 9. Adult extemal features of Stigmella spp. l, S. saginella. 2, S. castaneaefoliella. latifascie lla. 4, S. juglandifuliella. 5, S. bifasc iella. 6, S. intermedia. r28 MEMOIRS OF THE } NTOMOLOCICAL SOCIETY OF CANADA

PrArs 10. Adult external features of Stigmella, Ectoedemia, Obrussa, znd Glaucolepis spp. 1, S. procrastinella.2, S. alba. 1, E. canutus. 4, E. lindquisti. 5, E. rubifoliella. 6, E. quadrinotata.7, O. ochrefasciella, 8, O. sericopeTa. 9, G. saccharella, S{LKINSON AND SCOBLE: THE NEPTICULIDAE OF CANADA 129 Index to Genera, Species, and Subspecies Pace Pacr, alba...... '13 Nepticula .....5,12 anomalella ...... 18 Obrussa ...... 98 argyropeza ... ..ij obscurella .....54 a. argyropeza ...... 80 ochrefascielln ...... 99 a.downesi .....80 opulfoliella ...... 50 aromella ...... 27 ostryaefoliella ...... 54 bifosciella ...... 59 pallidella ...... 7 canutus ...... 81 paludicola .....51 caryoe.loltelld ...... 54,57 pectocatena ...... 18 castaneaefoliella ...... 44 plagicolella ...... 23 chalybeia ...... 35 platanella ...... 89 ciliaefuscella ...... 23 pomivorella ...... 33 clemensella .....86 populella ...... 74 corylifoliella ...... 50 populetorum ...... 26 crataegfoliella ...... 30 procrastinella ...... 70 Dechtiria .....7,'73 quadrinotata ...... 95 discolorella ...... 23 quercicastanella ...... 41 downesi ...... 80 quercipulchella ...... 65 Ectoedemia .....73 rosaefoliella ...... 14 Etainia ...... 98 r.pectocatena ...... 18 exasperata .. . .51 r. rosaefoliella ...... 17 fuscocapitella ...... 41 rubifoliella .....90 fuscotibiella ...... 23 saccharella ....105 Glaucolepis ...... 102 saginell.a ...... 39 immundella ...... 7 scinanella ...... 36 intermedia .....62 scintillans ...... 33 jughndifoliella..... sericopeTa ....101 ...... 57 serotinaeella ...... 59 latifasciella .....47 slingerlandella ...... 19 lindquisti ...... 83 stigmaciella ...... 38 Lyonetia .....77,80,98,101 Stigmella ...... 12 macrocarpae ...... 4'7 ...... 65 maximella terminella .....89 Trfurcula .7 ,73,98 Micropteryx ...... 33 turbidella ...... 78 Microsetia ...... 12 ...... 91 minimella ...... 50 ulmella myricafoliella ...... 5'l variella ...... 67 virginiella ...... 50