Systematic Entomology (1982) 7 , 367-463
A taxonomic revision of the North American species of Stigmella (Lepidoptera: Nepticulidae)
PHILIP J. NEWTON and CHRISTOPHER WILKINSON Department of Biological Sciences, Portsmouth Polytechnic, Portsmouth, England, and Department of Animal Systematics and Zoogeography, Vrije Universiteit, Amsterdam, The Netherlands ABSTRACT. A revision of the fifty-one species of the genus Stigmella from North America is presented. Thirty-one species are recorded only from the U.S.A., seventeen species and one subspecies from the U.S.A. and Canada and three species and one subspecies are known only from Canada. Five species are new: Stigmella inconspicuella (U.S.A., California); S.heteromelis (California); S.sclerostylota (Arkansas); S.longisacca (California) and S.plumosetaeella (Arizona). Life histories, mine characteristics and larval habits are given where known. These are collated from field study, material examined, breeding records and the literature. A list of host plants is also provided. A neotype is designated for Nepticula prunifoliella Clemens, previously considered to be possibly dipterous. Although most Stigmella species have different genital morphology, some pairs of species do not. In these pairs, however, external features and characteristics of the mine often differ - some pairs also differ in distribution or season. The taxonomic implications are discussed in the individual treatments - not all the problems, however, can be resolved without further rearing studies and collection. Species-groups have been designated on the basis of morphology. Some species are, however, structurally intermediate between species-groups and others may resemble Ectoedemia Busck in aspects of the male genitalia while their female genitalia, external features and venation are typical of Stigmella. Introduction
The family Nepticulidae is a cosmopolitan group comprising some of the smallest Lepidoptera known to science. Members of the family are exclusively herbivorous: larvae are known to mine leaves, produce galls on petioles and small twigs, or tunnel in the bark or fruits of the host-plant. Their small size, and consequent problems of collection and study, has resulted in the family being one of the poorest collected and investigated groups in the world. On a world basis Stigmella is probably not only the largest but also the most widespread Correspondence: Dr P. J. Newton, Citrus and Sub-tropical Fruit Research Institute, Nelspruit, Republic of South Africa.
genus in the Nepticulidae, occurring worldwide apart from in those extremes of climate where its plant hosts do not survive. Why the genus is so extensive and comparatively uniform is not clearly understood. The present study of North American Stigmella is complementary to the work of Wilkinson & Scoble (1 979) on the Nepticulidae of Canada. The species of Nepticulidae of the U.S.A. and Canada are regarded as falling into eight genera: Stigmella Schrank; Ectoedemia Busck (Wilkinson & Newton, 1981); Fomoria Beirne and Microcalyptris Braun (Wilkinson, 1979); Obrussa Braun and Glaucolepis Braun (Wilkinson & Scoble, 1979); Manoneura Davis (1 979) and Artaversala Davis (1978).
0307-6970/82/1000-0367 $02.00 0 1982 Blackwell Scientific Publications 25
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Thirty-one species of Stigmella recorded from the U.S.A. alone are described here. Diagnoses, and descriptions where necessary, are provided also for seventeen species recorded here from the U.S.A. and, previously, from Canada (Wilkinson & Scoble, 1979); a further three species and one subspecies so far recorded only in Canada are also included. In all, accounts of fifty-one species of Stigmella are given. Speciesgroups are adopted as in Wilkinson & Scoble (1979) and Johansson (1971). However, some species are structurally intermediate between species-groups. Other species resemble Ectoedemia in aspects of the male genital structure though female genital structures, external features and venation recommend their inclusion in Stigmella. The methods used are similar to those of Wilkinson & Scoble (1979) and Wilkinson & Newton (1981). Genital morphology is used extensively to provide indications of relationships and diagnostic characters. A thorough comparative morphological study of the various nepticulid genera is necessary, however, not only t o ascertain homologies within the group but also with other families; some of these problems have been discussed in earlier papers (Wilkinson & Newton, 1981; Scoble, 1980a, 1980b). Schematic genitalia figures are provided here (Figs. 7 and 8) t o indicate the terminology used for the various genital structures. Genitalia figures are each marked with a scale line representing 0.1 mm unless otherwise indicated; the scale of the adult illustrations may be estimated from the range of wing measurements in the descriptions. A list of the host plants of North American Stigmella species is appended as an aid to collectors (Appendix). References t o the catalogues of Dyar (1903) and McDunnough (1939) are not included for individual species. Lists of material examined include only those specimens not already listed by Wilkinson & Scoble (1979). Thus the material examined (including type-material) for this paper includes specimens listed here plus those listed by Wilkinson & Scoble (1979).
Abbreviations AFB: Dr Annette F. Braun, late of Ohio,
U.S.A. ANS: Academy of Natural Sciences of Philadelphia, Pennsylvania, U.S .A. BM NH: British Museum (Natural History), London, England. CNC: Canadian National Collection, Ottawa, Ontario, Canada. CW: the second author. DFF: Department of Fisheries and Forestry, Fiton, New Brunswick, Canada. FIS: Forest Insect Survey of Canada, Saulte Ste Marie (Prentice et al., 1965). MCZ: Museum of Comparative Zoology, Cambridge, Massachusetts, U.S.A. MJS: Malcolm J. Scoble (genitalia preparations). PJN : the senior author. SOO: Canada Department of Forestry, Saulte Ste Marie, Canada. UBC: University of British Columbia, Canada. USNM: United States National Museum of Natural History (Smithsonian Institution), Washington D.C., U.S.A. Historical background
From a nomenclatorial point of view there has been much discussion regarding the correct name for this genus. Stigmella is the older name but there have been attempts to reject it as a nomen nudum (e.g. Busck, 1913: 103; Braun 1917: 163) or a nomen dubium (e.g. Borkowski, 1972: 700; Lempke, 1976: 12) in favour of Nepticula. These arguments, together with the further implications that Stigmella is a nomen oblitum (e.g. Johansson 1971: 241), are discussed by Wilkinson (1978: 13-22). As he confirms, however, Stigmella should be retained as the valid name as to discard it would contravene the rules of the International Code of Zoological Nomenclature. Work on Stigmella in North America began in the late nineteenth century with accounts of several species from the immature stages. Clemens (1 862a, b) described fifteen species, but his habit of naming and describing them from the mine and host-plant only has been much criticized ; furthermore, he apparently preserved no type material of his Nepticulidae. Chambers later described fifteen species from captured or bred specimens (1873, 187Sa,b, 1878a, 1880). He sometimes recognized similarities between his mines and host-plants and those of Clemens; nevertheless, he continued to describe his species as new, not knowing the extent of variation to expect for each taxon. The systematics of the genus would be in a state of some confusion now
A taxonomic revision of Stigmella were it not for the subsequent rearing of many of the species by Braun (1909-30): there now remain only two or three unresolved names. As a result of Braun’s collecting and rearing, two Clemens names were synonymized with species he had described earlier in the same paper, and five Chambers species were found t o be synonymous with those described earlier by Clemens. Apart from these, Braun described twenty-five species presently combined with Stigmella. Most of these were obtained from reared material, so contributing t o our knowledge of life-histories. However, Braun rarely looked at the genitalia: following their examination five of Braun’s species were synonymized by Wilkinson & Scoble (1979). The study of genital morphology led Beirne (1945) t o propose the division of Stigmella sensu strict0 into two groups, Stigmella and Nepticula. The significance of this division has been discussed by Wilkinson ( 1 978). Beirne’s proposal was not followed by most subsequent workers. The a priori use of host-plant relationships, not only t o indicate affinity but also for identification of species, has been prevalent in the past but is not recommended. There are distinct dangers in the philosophy: if different host-plants then ips0 facta different species of leaf-miner. This was formerly the belief in the case of the species-group corylifoliella, for example, and was corrected by Wilkinson & Scoble (1 979) after genital studies. However, in general, genital differences correlate with major groupings in classification by hostplants.
Further contributions t o our knowledge of the North American species have been made by several authors. Most notable amongst these are the extensive studies of single species by Forbes & Leonard ( 1 930) and Jones (1 933). One new species described here was collected by Jones and Braun in 1929. It is labelled ‘Stigmella photiniae’ but the name has never appeared in the published literature: it is given a new name here t o avoid confusion. It is, however, interesting t o note the use of the generic name Stigmella which was uncommon amongst contemporary North American writers. One exception was Grossbeck (1917: 145) who even earlier combined myricafoliella and condaliafoliella with Stigmella. One bibliographic problem, the publication cited here as Clemens (1 862a), warrants some explanation. The publication itself bears the date November 1861 but reference t o Brown (1964: 307) indicates that the Journal was not presented t o the society’s library until 13 January 1862. No questions of priority appear t o be involved. Key to North American genera of Nepticulidae M of forewing coalescing with Cu for part of its length and M or M + Cu passing obliquely t o R to form a cross-vein; R, and R, o f forewing as separate branches (Figs. 2 , 4 , 5 and 6) . . . . . 2 - M of forewing not coalescing with Cu but with R from base, not forming cross-vein; R, and R, o f forewing coincident or R not branched (Figs. 1 and3) . . . . . . . . . . . . . . . . . . . . . . . 5 2 M o f forewing two-branched (Figs. 5 and 6) . 3 - M of forewing not branched (Figs. 2 and 4) . . 4 3 Male genitalia with large dorsal process on valve projecting transversely t o reach opposite side of 1
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FIG. 1. Stigmella slingerlandella d , wing venation.
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M FIG. 2 . Ectoedemia similella P, wing venation.
genital capsule; saccus convex. Female genitalia with sclerotized antrum on ductus; signa comprising two ovate, reticulate patches. Hindwing with M single-branched (Fig. 5 ) . . Obntssa Braun Male genitalia with valve without dorsal process; saccus concave, bilobed. Female genitalia without sclerites on ductus; signa comprising two linear patches of pectinations. Hindwing with M twobranched (Fig. 6) . . . . . . . . Glaucolepis Braun Male genitalia with uncus in the form of a bridge; valves with large dorsal spine. Female genitalia with funicular sclerotization proximal t o genital aperture, ductus simple in other respects; signum comprising two linear reticulate patches. M + Cu of forewing forming cross-vein, Cu becoming obsolete (Fig. 4) . . . . . . . . . Fomoria Beirne Male genitalia with uncus membranous; valves without dorsal spine. Female genitalia not usually with sclerites proximal t o genital aperture, frequently with sclerotized double ring on ductus proximal t o the bursa; signum comprising two ovate reticulate patches. M only passing to R t o form a cross-vein, Cu reaching a point beyond middle of wing (Fig. 2 ) . . . Ectoedemia Busck Hindwing with R and M fusing in basal half t o form a common trunk (Figs. 1 and 3) . . . . . 6 Hindwing with R and M as separate branches . 7 Male genitalia with U-shaped vinculum; gnathos with short in the shape of a ‘U’, ‘M’ or ‘W’, anterior and posterior processes. Female genitalia with simple ductus; signum,if present, comprising ridges or spicules, R , of forewing separate from R s + M at one-third the distance from the base (Fig. 1) . . . . . . . . . . . . . Stigmella Schrank Male genitalia with ring-shaped vinculum; gnathos horn-shaped medially, with large anteriorlydirected lateral processes associated with the vinculum. Female genitalia usually with complex sclerotization of the ductus; signum comprising two ovate or linear reticulate patches. R , of forewing separate from Rs + M at the base (Fig. 3) Microcalyptris Braun Male genitalia with gnathos complex and closely associated with bridge-like sclerite believed t o represent the uncus; saccus weakly convex ;valves tapering apically, with dorsal spine. [Female
genitalia not known.] R , of forewing coincident with R 1 + 3 . . . . . . . . . . . . Manoneura Davis - Male genitalia with gnathos as simple transverse sclerite with short lateral arms: pseuduncus apparently membranous;saccus tapering markedly anteriorly; valves deeply excavated apically, without dorsal spine. Female genitalia with coiled vesicle on ductus; bursa copulatrix without signum. R of forewing not branched A rtaversala Davis
Stigmella Schrank Stigmella Schrank, 1802: 169. Type species by subsequent designation (Walsingham, 1907 : 1008), Phalaena (Tinea) anomalella Goeze, 1783: 168. Stigmella Schrank; Wilkinson, 1978: 13 [Detailed taxonomic history of the nomenclature and more complete bibliography] ; Wilkinson & Scoble, 1979: 12 [Revision of Nepticulidae of Canada]. Nepticula Heyden, 1843: 208. Type species by subsequent designation (Tutt, 1899: 184), Tinea aurella Fabricius, 1775: 666. (Synonymized by Walsingham, 1907: 1008.) Johanssonia Borkowski, 1972: 702. Type species by original designation and monotypy, Nepticula acetosae Stainton, 1854: 303. (Synonymized by Karsholt & Nielsen, 1976: 17, 81, but retained as separate subgenus.) External features: d0. Palps extending just beyond front of head capsule; antenna approximately half length of forewing. Venation as in Fig. l . Forewing: M coalescing with R from base and anastomosing to a point
A taxonomic revision of Stigmella beyond middle of wing; M single-branched; Cu arising separately, not reaching further than three-quarters length of wing; R 1 separate from Rs + M at approximately one-third distance from base; R4 coincident with R5; A not meeting Cu. Hindwing: M and R both single-branched, forming a common trunk in basal half. Forewing: elongate-ovate; fringe with diffuse margin at apex, or, in exceptional cases, well defined by apical band of wing scales; other markings, if present, as one or two fasciae or patches. Hindwing: narrow and lanceolate, half t o two-thirds width of forewing. Legs: proximal pair of spurs on hind tibia before the middle. Male genitalia. Vinculum U-shaped, except in one species where it is ring-shaped. Tegumen strap-like, articulating or sometimes fusing with dorsal arms of vinculum. Uncus bilobed, articulating with tegumen and vinculum laterally, sclerotized marginally. Gnathos Ushaped, M-shaped or W-shaped with dorsolateral arms arising from transverse ventral plate; in three species, medial arms fusing t o form a central process, Saccus bilobed. Valves usually quadrate and frequently bifurcate distally to form a style. Transtilla: either Ushaped or W-shaped ;transverse bars sometimes separate. Juxta, if present, weakly sclerotized. Aedeagus: simple, most variable in length between species; vesica usually adorned with small denticles orientated in a ridge and with a plate of minute papillae, rarely with anellar spines or plates. sc
Female genitalia. Apophyses generally shorter than ductus bursae. Ductus with accessory sac sometimes denticulate. Bursa copulatrix large and variously pectinate; signum usually absent, but if present weakly sclerotized and comprising ridge-like plates or spicules. Larvae. Exclusively leaf-miners, usually forming linear tracts, occasionally terminating in blotches; only six true blotch-makers are known at present. Remarks For a differential diagnosis of Stigmella and other North American genera of Nepticulidae, see Wilkinson & Scoble (1 979). Check-list of North American Stigmella
STZGMELLA Schrank, 1802 NEPTICULA Heyden, 1843 crutuegifoliellu (Clemens, 1862a) scintilluns (Braun, 1917) pomivorellu (Packard, 1870) chulybeiu (Braun, 1914) scinunellu Wilkinson & Scoble, 1979 purpurutella (Braun, 19 17) stigmaciefla Wilkinson & Scoble, 1979 tueniolu (Braun, 1925) prunifoliellu (Clemens, 1862a) bifusciella (Clemens, 1862b) serotinaeella (Chambers, 1873) ceunothi (Braun, 1910) intermedia (Braun, 19 17) rhoifoliella (Braun, 19 12) R,
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FIG. 4.Fomoria hypericella d , wing venation.
rhamnicola (Braun, 1916) diffasciae (Braun, 1910) inconspicuella sp.n. gossypii (Forbes & Leonard, 1930) heteromelis sp.n. cerea (Braun, 19 1 7) rosaefoliella (Clemens, 1862a) r.rosaefoliella (Clemens, 1862a) r.pectocatena Wilkinson & Scoble, 1979 slingerlandella (Kearfott, 1908) villosella (Clemens, 1862a) dallasiana (Frey & Boll, 1876) apicialbella (Chambers, 1873) leucostigma (Braun, 19 12) fuscotibiella (Clemens, 1862b)
ciliaefuscella (Chambers, 1873) discolorella (Braun, 19 12) populetorurn (Frey & Boll, 1878) aromella Wilkinson & Scoble, 1979 pallida (Braun, 19 12) saginella (Clemens, 1862a) quercicastanella (Chambers, 1873) fuscocapitella (Chambers, 1873) latifasciella'(Chambers, in Hayden, 1878a) macrocarpae (Freeman, 196 7) nigriverticella (Chambers, 1875a) maculosella (Chambers, 1880) castaneaefoliella (Chambers, 18 7 5 a) flavipedellu (Braun, 1914) sclerostylota sp.n.
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FIG. 5 . Obrussa sericopeza 9 , wing venation.
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sc
FIG. 6. Glaucolepis saccharella d, wing venation.
corylifoliella (Clemens, 1862a) virginiella (Clemens, 1862a) minimella (Chambers, 1873) opulifoliellu (Braun, 19 14) paludicola (Braun, 1917) exasperuta (Braun, 1930) ostryaefoliella (Clemens, 1862a) caryaefoliella (Clemens, 1862a) obscurella (Braun, 1912) myricafoliella (Busck, 1900) juglandifoliella (Clemens, 1862a) longisacca sp.n. unifasciella (Chambers, 1875a) condaliafoliella (Busck, 1900) ti6iella (Braun, 1912) quercipulchella (Chambers, in Hayden, 1878a) terminella (Braun, 1914) variella (Braun, 1910) altella (Braun, 19 14) procrastinella (Braun, 1927) alba Wilkinson & Scoble, 1979 braunella (Jones, 1933) argentifasciella (Braun, 1912) plumosetaeella sp .n. resplendensella (Chambers, 187Sa) belfrageella (Chambers, 1875b) amelanchierella (Clemens, 1862a) (generic position uncertain) The following species previously assigned to ‘Nepticula’ have been transferred to Ectoedemia Busck by Wilkinson ( 1 98 1).
canadensis (Braun, 19 17) grandisella (Chambers, 1880) anguinella (Clernens, 1862a) platea (Clemens, 1862a) Key to North American Stigmella, species and species-groups* 1
Vertex pale ochreous or buff
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2
.... 4 Forewing pale buff, without markings . . pallida Forewing dark brown, markings present . . . . 3
- Vertex usually dark ochreous o r brown 2 -
3 Forewing with basal patch of creamy white scales. Female with pectinations of bursa covering distal two-thirds cerea - Forewing without basal patch of white scales. Female with pectinations of bursa restricted t o medial t h u d heteromelis 4 Male with uncus single-lobed 5 - Male with uncus bilobed 6 5 Male with cuiller rather pointed apically. Female with a patch of heavily sclerotized spines on bursa or accessory sac species-group procrustinella - Male with cuiller not pointed apically. Female without patch of spines on bursa o r accessory sac
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braunella
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6 Male with gnathos M-shaped 7 - Male with gnathos U-shaped 18 7 Male with aedeagus approximately twice length of genital capsule. Female with accessory sac markedly spiral species-group quercipulchella - Male with aedeagus usually less than twice length
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*Three species - resplendensella, belfrageella and arnelanchierella - have been omitted from the key because there is either very limited material or none in existence on which t o base a diagnosis or, indeed, confirm identity.
374
Philip J. Newton and Christopher Wilkinson tegumen
. ,
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style
lateral arm dorso lateral arm transverse ventral date
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lateral arm transti Ila
transverse bar ventral arm
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FIG. 7. Schematic drawing of male genitalia of a Stigmella.
of genital capsule. Female with accessory sac not markedly spiral . . . . . . . . . . . . . . . . . . 8 Iemale with accessory sac spinose o r granulate 9 or granulate ~~~~l~ with accessory sac
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13
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Male with transtilla fused. Eemale with accessory sac usually larger than bursa . . . . . . . . . . 10 Male with transtilla separate. Female with accessory sac smaller than bursa . . . . . . . . . . . 12 10 Male with valve markedly divided into style and cuiller; valve with plumose setae along inner surface . . . . . . . . . . . . . . . .plumose taeella - Male with valve weakly divided into style and cuil1er;valve without plumose setae . . . . . . 11 11 Male with cuiller hook4ike apically. Female with accessory sac with stout, single spines apicialbella - Male with cuiller tapering apically. Female with accessory sac with weakly sclerotized, double spines . . . . . . . . . . . . . . . . . . . villosella 12 Male with vinculum and S ~ C C U S narrow; lateral aims of gnathos usually closely juxtaposed. Female with signum comprising a U-shaped hand or two rows of spinose ridges species-group saginella - Male with vinculum and saccus very hroad; lateral arms o f gnathos widely spaced. Female without signum . . . . . . . . . . . . . unifasciella 13 Female with signum comprising a Ushaped band o f pectinations . . . . species-group fuscotibiellu - Female without signum . . . . . . . . . . . . . 14 14 Forewing with two complete, whitish fasciae . 1 5
Forewing without two complete fasciae: markings, if present, a single fascia or spots or patches 16
. . . . . . . . . . . . . . rliamnicola Thorax brown . . . . . . . . . . . . . . diffusciae
1 5 Thorax white
16 Male with transtilla continuous; transverse bars of
transtilla very broad; uncus quadrate, with small, pointed, lateral lobes . . . . . . . . inconspicuella - Male with transtilia sometimes separate; transverse bars of transtilla narrow; uncus with broad lobes. . . . . . . . . . . . . . . . . . . . . . . . 17 1 7 Male with gnathos with posteriorly-directed processes reaching beyond tegumen; valve tapering apically . . . . . . species-group rosaefoliella - Male with gnathos with posteriorly-directed processes n o t usually reaching beyond tegumen; valve quadrate . . . .species-group crafaegifoliella 18 Male with tegumen fused with vinculum dorsally; abdomen with lunulate patches of specialized scales ventrally on segments V1, VI1 and VIII argen tifasciella - Male with tegumen articulating with vinculum dorsally; abdomen without patches of specialized scales. . . . . . . . . . . . . . . . . . . . . . . . 19 19 Male with spinose membrane associated with uncus and gnathos . . . . . . . . . . . . . . . . 2 0 - Male without spinose membrane associated with uncus and gnathos . . . . . . . . . . . . . . . . 2 1 2 0 Male with aedeagus as long as capsule; vesica with many cornuti orientated in a ridge. Female with accessory sac simple and membranous gossypii
A taxonomic revision of Stigmella
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- Male with aedeagus shorter than capsule; vesica
-
with few cornuti, scattered over its surface. Female with accessory sac usually with a patch of spines . . . . . . . . species-group prunifoliella 2 1 Male with aedeagus with a pair of palmate plates apically. Female with a pair of ovate signa species-group curylifulietla - Male with aedeagus without palmate sclerites apically. Female without signum or, if present, single. . . . . . . . . . . , . . . . . . . . . . . . 2 2 22 Male with markedly lobed saccus; valve tapering apically, not divided. Female with plate-like sclerotization of the bursa. . . . condaliafoliella
Wilkinson & Scoble (1979) described this species-group which includes the following:
Male with weakly lobed saccus; valve quadrate, divided into style and cuiller. Female with scallopshaped pectinations of t h e bursa . . . . . . tiliella
Species-group CRA TAEGIFOLIELLA
crataegifoliella, scintillans, pomivorella, chaly beia, scinanella and stigmaciella ; to which are now added t h e North American species taeniola and purpuratella. The group
FIG. 8. Schematic drawing o f female genitalia of a Stigmella.
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Philip J. N e w t o n and Christopher Wilkinson
FIG. 9. Stigmella crataegifoliella d , genitalia: (a) genital capsule; (b) aedeagus
is characterized by the form of the uncus, gnathos, valve and transtilla in the male genitalia, and in the simple bursa copulatrix in the female. All Species in the group mine Rosaceae. Key to species Forewing with extensive ochreous scales crataegifoliella Forewing without ochreous scales, ground colour brownish . . . . . . . . . . . . . . . . . 2 Forewing without markings, generally lacking iridescence . . . . . . . . . . . . . . . . . . . . 3 Forewing with fasciae or patches, strongly 4 iridescent . . . . . . . . . . . . . . . . . . . . . Male with lateral arms of vinculum with swollen knobs dorsally. Larva on Pyrus . . . . . chalyheia Male with lateral arms of vinculum lacking swollen knobs. Larva o n Malus o r Crataegus pomivorella Tufts on front of head and vertex dark brown sclntillans Tufts on front of head and vertex orangeochreous. . . . . . . . . . . . . . . . . . . . . . 5 Male with uncus with ear-like lateral lobes, valve with cuiller pointed apically, aedeagus larger than capsule . . . . . . . . . . . . . . . . 6 Male with uncus with pointed lateral lobes, valve with cuiller not pointed apically, aedeagus shorter than genital capsule . . . . . . . . . . . 7 Male with forewing shining predominantly red and purple, female purple and gold. Larva on Malus scinanella
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- Male with forewing shining purple, female gold. 7 -
[Larva not known] . . . . . . . . . purpuratella Larva on Crataegus, making a stigmatonome stigmaciella Larva on Amelanchier. making an 0phistie;. matonome . . . . . . . . . . . . . . . . . . taeniola
Stigmella crataegifoliella (Clemens) (Figs. 9 ~
~
- \
and 1U) Nepticula crataegifoliella Clemens, 1862a: 8 3 [description of mine and larva] ; Braun, 1914: 17 [description of adult]. Stigmella crataegifoliella (Clemens); Wilkinson & Scoble, 1979: 30 [redescription, neotype designated]. DESCRIPTION. See Wilkinson & Scoble (1979). BIOLOGY. Voltinism. Braun (1917: 194) stated that larvae may be found in early July and the beginning of September and was of the opinion that there are two generations per year. The material examined confirms this, adults being taken in May and August in Ohio; specimens taken in February by Lewis were probably ‘forced’ to emerge earlier than is normal by artificial conditions in the laboratory. DIAGNOSIS. Easily distinguished externally by the extensive ochreous scales of the forewings; male and female genitalia (Figs. 9 and
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the frass pattern is characteristic and stigmaciella which forms a blotch mine. DISTRIBUTION. U.S.A. : Ohio, Pennsylvania, Kentucky (Chambers). CANADA: Ontario. Material examined See Wilkinson & Scoble (1979). U.S.A.: Ohio, Remington, 1 mine, o n Crataegus sp., 24.viii. 1916 (Braun) (ANS); Pennsylvania, West Chester, 6 mines, o n Crataegus sp., 1965 (Lewis) (CNC). CANADA, Ontario, Simcoe, 7 mines, o n Crataegus sp., 4.viii. 1959 (Lewis) (CNC).
Stigmella scintillans (Braun) Nepticula scintillans Braun, 1917: 167. Stigmella scintillans (Braun); Wilkinson & Scoble, 1979: 33 [ redescription].
DESCRIPTION. See Wilkinson & Scoble (1979). BIOLOGY. Voltinism. The material examined here shows that there are two generations per year; the second generation larvae found in July and early August normally overwinter but Braun (1917: 168) noted that a third generation may sometimes be collected in late August.
-FIG. 10. Stigmella crataegifoliella 9 , genitalia.
l o ) , however, are not easily separated from those of their nearest relatives, scintillans and pomivorella. The form of the gnathos, the length of the capsule, the variable width of the aedeagus, and the spiculate nature of the accessory sac in pomivorella, are the only observed differences. The flattened lobes of the uncus separate these three species from other members of the group in which the lobes are ear-like or pointed laterally. The broad ophionome produced by larvae of crataegifoliella distinguishes the species from other Crataegus feeders, scintillans in which
DIAGNOSIS. The external features differ from those of all other members of the group in the dark colour of the head tufts; the ground colour of the forewing differs in the other brightly shining species, stigmaciella and scinanellu. The genitalia are indistinguishable from those of cratuegifoliella except in the degree of sclerotization of the pectinations of the bursa; in all other respects the diagnosis of the former species serves t o separate the genitalia of scintillans from those of other members of the group. The contorted nature of the mine, in which the frass is distributed in arcs across its breadth, separates this species from crataegifoliella which produces a broad ophionome, and stigmaciella which forms a blotch, in Crataegus. DISTRIBUTION. U.S.A. : Ohio, Michigan. CANADA: Ontario. Material examined See Wilkinson & Scoble (1979). U.S.A.: Ohio, Anderson’s Ferry, 4 mines, 1 cocoon, on Crataegus mollis, 22.vi. 1909 (Braun) (ANS). CANADA: Ontario, Ottawa, 4 mines,
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Philip J. Newton and Christopher Wilkinson
FIG. 1 1 . Stigmella pomivorella d , genitalia: (a) genital capsule; (b) aedeagus.
on Crataegus sp., 4.viii. 1955 (Freeman), 12.viii. 1955 (Lewis) (CNC).
members of the crataegifoliella group in the ground colour and degree of iridescence.
Stigmella pomivorella (Packard) (Fig. 1 1)
DISTRIBUTION. U.S.A.: New York, Washington, Massachusetts (Packard). CANADA: Nova Scotia, Ontario, British Columbia.
Micropteryx pomivorella Packard, 1870: 237. Nepticula pornivorella (Packard);Busck, 1901: 52 [ T y p e ] . Stigmella pomivorella (Packard); Wilkinson & Scoble, 1979: 33 [ redescription] . DESCRIPTION. See Wilkinson & Scoble 1979); regarded as being represented by two forms. DIAGNOSIS. Males of the western form differ from those taken in the east in the broad aedeagus relative t o the genital capsule, and in the cornuti which are confined to a medial patch; the cornuti are widespread in the eastern form. This species differs from its nearest relative, chalybeiu, in the absence of the swollen knobs on the lateral arms of the vinculum (Fig. 1 1) whereas t h e female genitalia are indistinguishable from those of crutaegifoliellu. The external features differ from other
Material examined See Wilkinson & Scoble ( 1 979).
Stigmellu chalybeia (Braun) (Figs, 12, 13, 84 and 99) Nepticula chalybeia Braun, 1914: 2 0 ; Braun, 1917: 190 [redescription] ; Braun, in Forbes, 1923: 93 [ redescription] . Stigmella chalybeia (Braun); Wilkinson & Scoble, 1979: 35 [diagnosis, lectotype designated]. DESCRIPTION. External features: 0”. Palps greyish; antenna dark brown with metallic reflections; tuft on front of head ochreous, vertex paler; eye-caps shining white; collar off-white. Thorax dark brown with strong metallic grey reflections, abdomen pale and shining metallic grey beneath. Forewing:
A taxonomic revision of Stigmella
ground colour of fringe and dorsal surface brown with purplish grey lustre, strongly iridescent only when light at suitable angle. Hindwing: ground colour and fringe brownish grey, with metallic grey reflections. Legs brown above, paling o n tarsi, leaden grey on lower surface. 0 (Fig. 99). As d except for a pair of ventromedial fenestrae o n fourth abdominal segment. Wing measurements. d, 3 G 4 . 2 mm (5). 9, 3.4-4.4 mm (8). Lectotype, 3.8 mm. Genitalia. d (Fig. 12). Uncus bilobed, with each lobe broad and rather flattened. Gnathos: as in Fig. 12(a); transverse ventral plate broad; posteriorly-directed lateral arms widely spaced, short and broad; dorsolateral arms broad with short posteriorly- and anteriorlydirected processes. Tegumen a broad arcuate plate. Vinculum: lateral arms broad, bifurcate dorsally and articulating with tegumen, with posterior branches expanded and fusing dorsally as in Fig. 12(a); ventral plate broad. Saccus bilobed, approximately half width of
319
ventral plate. Valve with long blunted style reaching just beyond uncus and with a broad constricted cuiller. Transtilla: lateral arms broad and arcuate; transverse bars broad and continuous forming a horizontal bar at right angle t o lateral arms. Aedeagus: narrow and regular in width, approximately equal in length t o genital capsule; vesica with many denticulate cornuti and with globular plate of minute papillae. 9 (Fig. 13). Ductus bursae long and broad. Accessory sac large with reticulate lobe proximally and with granular duct arising distally. Bursa copulatrix: large covered with long chains of pectinations as in Fig. 13(b); signum absent. Anterior apophyses long and narrow, slightly arcuate. Posterior apophyses long and straight, reaching beyond apophyses anteriores. BIOLOGY. Egg. Laid o n lower surface of leaf. Host. Pyrus sp.: specimens examined here are bred from larvae mining wild pear and cultivated pear. Braun (1 9 17: 191) suggested
FIG. 12. Stigmella chalybeia d , genitalia: (a) genital capsule; (b) aedeagus.
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Philip J. Newton and Christopher Wilkinson
that the species is confined to pear as a food plant since apple trees, adjacent t o pear trees covered with mines, are entirely free from them. Mine. Upper-surface ophionome, beginning as short, linear tract gradually increasing in breadth; frass deposited as broad central line (Fig. 84). Larva. Emerges on upper surface of leaf. Cocoon. Small, obovoid, greenish brown. Voltinism. Braun (1914: 21) reported that mines may be collected in June, July and late August and that there are three generations per year. The material examined in this study represents the two later broods; the first cannot be confirmed.
DIAGNOSIS. Differentiated from other members of the group by the uniform colour of the forewing and from pomivorella b y the generally smaller alar expanse, the paler ground colour and absence of purple reflections on t h e wings. The swollen lateral arms of the vinculum separate chalybeia from pomivorella and it differs from scinanella and purpuratella in the nature of the uncus. The female genitalia are indistinguishable from those of pomivorella. DISCUSSION, This species is closely related to pomivorella in external features and genitalia but is apparently specific in its choice of host plant. DISTRIBUTION. U.S.A.: Ohio, New Jersey. CANADA: Ontario. Material examined See Wilkinson & Scoble (1979). U.S.A.: Ohio, Cincinnati, 3 d d , 399, 3 mines, o n Pyrus communis, 8.vii. 1913, 13-18.vii. 1914 (Braun), 1 d, 4 9 0 , 2 mines, o n cultivated pear, 15-31 .vii. 1916, 3.viii. 1916 (Braun), 1 d, 30.viii. 1919 (Braun) (ANS); New Jersey, Hadden Heights, 1 9 , 2.viii. 1931 (Bottimer) (USNM).
Stigmella scinanella Wilkinson & Scoble (Figs. 14 and 15) Stigmella scinanella Wilkinson & Scoble, 1979: 36. DESCRIPTION. See Wilkinson & Scoble ( 1979).
DIAGNOSIS. Closely related to purpuratefla but the ground colour of t h e forewing is darker and the degree of iridescence much greater in scinanella. The large ear-like lobes of the uncus and the distribution of t h e cornuti distinguish scinanella males from stigmaciella and taeniola: the male genitalia of scinanella and purpuratella are, however, indistinguishable (see Figs. 14 and 15).
0.02 rnm FIG. 1 3 . Stigmella chalybeia 0,genitalia: (a) genitalia; (b) enlarged detail of pectinations on bursa copulatrix.
DISCUSSION. See taeniola. DISTRIBUTION. CANADA: Ontario. Material examined See Wilkinson & Scoble (1979). CANADA, Ontario: Ottawa, 1 mine, o n Malus sp., 4.viii. 1955 (Freeman); Normandale, 6 mines, on Malus sp., 2 6 . 6 . 1 9 5 6 , 1 5 . v l i . 1 9 5 7 (Freeman and Lewis) (CNC).
A taxonomic revision of Stigmella
381
-
PIG. 14. Stigmella scinanella d , genitalia: (a) genital capsule; (b) aedeagus.
Stigmella purpuratella (Braun) c0mb.n. (Figs. 16 and 100) Nepticula purpuratella Braun, 19 17: 176; Braun, in Forbes, 1923: 88 [redescription]. DESCRIPTION. External features: S (Fig. 100). Palps whitish; antenna dark brown; tuft o n front of head orange-ochreous, vertex ochreous; eye-caps shining white ; collar offwhite, sometimes pale ochreous. Thorax and abdomen dark brown with deep bronze reflections. Forewing: ground colour of dorsal surface dark brown, basal third with green, red and mainly gold reflections, apical twothirds shining deep purple; fringe grey-brown, shining grey; single postmedial fascia, shining silver. Hindwing: ground colour and fringe greyish brown, shining metallic grey. Legs pale grey-brown, shining metallic grey. 0 . As in d except for forewing with predominantly gold reflections on the distal twothirds. Wing measurements. 9, 4 . 4 m m (1). Holotype, 4.2 mm. Genitalia. d (Fig. 16). Uncus bilobed, each
lobe ear-like and with lateral papillae. Gnathos: broadly H-shaped as in Fig. 16(a); posteriorlydirected lateral arms broad, not reaching uncus; dorsolateral arms with very short posteriorly-directed processes and long anteriorly-directed processes. Tegumen an ovate plate with serrate posterior margin. Vinculum: lateral arms tapering markedly with club-like swellings dorsally at point of articulation with tegumen ;ventral plate broad. Saccus narrow and deeply bilobed, each lobe as long as broad a t base. Valve divided into broad pointed style and cuiller with triangular process terminally. Transtilla: W-shaped as in Fig. 16(a); lateral arms narrow; ventral arms triangular; transverse bars tapering markedly and fusing medially. Juxta a quadrate membranous plate, broadening posteriorly. Aedeagus: flask-shaped, one-and-a-half times length of genital capsule; vesica with many small cornuti orientated in a ridge and with a globular plate of minute papillae. 0 . Not known (1 specimen, abdomen missing).
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Philip J. Newton and Christopher Wilkinsl711 CNC 3467; DFF). Paratype P, Pennsylvania, Allegheny County, 5 .vi. 1 9 10 (Marloffl (ANS).
Stigmella stigmaciella Wilkinson & Scoble Stigmella stigmaciella Wilkinson & Scoble, 1979: 38.
1
._."
.
I
DESCRIPTION. See Wilkinson & Scoble (1979). DIAGNOSIS. The ground colour is darker and there are more red and purple reflections on the forewing in stigmaciella than in taeniola. The male genitalia differ from those of scinanella and purpuratella in the lobes of the uncus, which are small and widely separated, and in the aedeagus which is shorter than the genital capsule in stigmaciella. Both male and female genitalia of stigmaciella are indistinguishable from those of taeniola except for the terminally digitate setae in the male of the latter. DISTRIBUTION. CANADA: British Columbia.
Material examined See Wilkinson & Scoble (1 979). CANADA: British Columbia, Vernon, 6 mines, on Crataegus sp., 3.vi. 1957 (Freeman and Lewis) (CNC).
.
.,..
- -
..-.
/
/'
/
.... '.
...
FIG. 1 5 . Stigmella scinanella 0, genitalia.
BIOLOGY. Immature stages and host unknown. DIAGNOSIS. The paler and less iridescent forewings differentiate this species from scinanella which is otherwise indistinguishable. The large ear-like lobes of the uncus in purpuratella differ from those of stigmaciella and taeniala, in which the uncus is rather pointed laterally, DISCUSSION. See taeniola. DISTRIBUTION. U.S. A . : Pennsylvania.
Type material examined Holotype d , U.S.A., Pennsylvania, Pittsburgh, 30.v. 1906 (Engsl) (genitalia slide no.
Stigmella taeniola (Braun) c0mb.n. (Figs. 17, 1 8 , 8 5 and 101) Nepticula taeniola Braun, 1925: 226. DESCRIPTION. External features: d (Fig. 101). Palps greyish; antenna dark greyish brown; tufts o n front of head and vertex ochreous, becoming whitish laterally on the vertex, eye-caps shining white; collar offwhite. Thorax dark brown with bronze reflections. Abdomen dark grey-brown, shining metallic grey on both surfaces. Forewing: ground colour of dorsal surface dark greyish brown, basal half shining gold, distal half more golden bronze; fringe pale greyish brown, shining metallic grey; single postmedial fascia, shining silver. Hindwing: ground colour and fringe pale grey-brown, shining silver. Legs brownish grey, shining silver. 0. As d except for forewing with predominantly purple reflections distally. Wing measurements. 0, 5.2mm (1). Holotype, 5.2 mm. Genitalia. d (Fig. 17). Uncus very broad with pair of widely separated, lateral papillae. Gnathos: transverse ventral plate narrow with
A taxonomic revision of Stigmella
383
I
FIG. 16.Stigmella purpuratella d , genitalia: (a) genital capsule; (b) aedeagus.
medial expansion; posteriorlydirected lateral arms long and narrow; dorsolateral arms with broad, triangular, posteriorly-directed processes. Tegumen a broad transverse plate. Vinculum: lateral arms narrow, fusing with tegumen at dorsal extremities; ventral plate narrow. Saccus broader than ventral plate, markedly bilobed. Valve: very large and quadrate, with broadly pointed style not reaching beyond uncus; style bears terminally digitate scales as in Fig. 17(c). Transtilla: lateral arms broad; transverse bars regular in width and at right angles to lateral arms, meeting but not fusing. Aedeagus: almost as broad as long, shorter than genital capsule; vesica with many large cornuti evenly distributed and with a globular plate of minute papillae. 26
9 (Fig. 18). Ductus bursae short. Accessory sac markedly folded and with simple duct arising proximally. Bursa copulatrix : covered with long chains of weakly sclerotized pectinations as in Fig. 18(b); signum absent. Anterior apophyses broad. Posterior apophyses narrow and reaching just beyond the apophyses anteriores. BIOLOGY. Egg. Laid on lower surface of leaf. Host. Amelanchier alnifolia. Braun ( 1 9 2 5 : 226) reported that untenanted mines, identical t o those produced by this species, were observed on leaves of Crataegus sp. in Idaho. Mine. Upper surface ophistigmatonome : begins as a short, contorted linear mine which enlarges into a funnel and then a large blotch; frass deposited centrally in all regions of mine, densely packed (Fig. 8 5 ) .
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Philip J. Newton and Christopher Wilkinson
Larva. Emerges o n upper surface of leaf. Cocoon. Pale brownish.
DIAGNOSIS. Differs from stigmaciella in the weaker reflections of the forewings and the paler ground colour. The lobes of the uncus, which are small and widely separated, and the aedeagus, which is shorter than the genital capsule, separate taeniola from scinanella and purpuratella. See Table 1 for other diagnostic features. DISCUSSION. The preceding three species (scinanella, purpuratella and stigmaciella) are closely related t o each other and to taeniola. The external features are similar in all respects apart from the major colour of iridescence of the distal area of the forewing, which is most variable in females. The male genitalia are of two forms and, together with the type of mine produced and the host plant, provide the major diagnostic characters; female genitalia are more difficult t o separate. Wilkinson & Scoble (1979) stated that scinanella lacks the markedly folded accessory sac in the genitalia, which is present in stigmaciella and taeniola;
however, there is some evidence of a weakly folded accessory sac in one preparation of scinanella (see Fig. 15). The distribution of t h e most significant attributes for the four taxa is summarized in Table 1. The differences in the mines provide the most sound evidence for the separate identities of taeniola and stigmaciella. Sscinanella and purpuratella are also regarded as separate species because the differences in external features correlate with those between taeniola and stigmaciella : host and mine characteristics are not known for purpuratella however. DISTRIBUTION. U.S.A.: Utah; Idaho (Braun). Type material examined Holotype d, U.S.A., Utah, Cache County, 9.vii. 1924 (Braun) (genitalia slide no. 23 PJN; ANS). Paratype 0 , dataas holotype,9.vii. 1924 (Braun) (genitalia slide no. 24 PJN; ANS). Further material examined U.S.A., Utah, Cache County, Logan Canyon, 1 mine, on Amelanchier alnifolia, 21 [ o r 261 .vi. 1924 (Braun) (ANS).
TABLE 1. Distribution of key characters amongst the species scinunelfn. purpuratella, stigmaciellu and taeniola. Major colour of iridescence in distal portion of forewing
d genitalia
Q genitalia
Host plant
Mine
scinanella
d: red-purple Q: purple or gold
1 . Uncus with earlike lobes 2. Aedeagus longer than capsule 3. Cuiller pointed 4. Transtilla fused
With weakly folded accessory sac
Malus
Ophionome Ontario (Canada)
purpuratella
d: purple 0 : gold
1. Uncus with ear-
Not known
Not known
Not known
Pennsylvania ((1 .S.A .)
Crutaegus
Stigmatonome
British Columbia (Canada)
A melanchier (Crataegus)
Ophistigmatonome
Utah (Idaho) (U.S.A.)
stigmaciella
d: gold
0: gold or purple
t ae ni o1u
d : gold 0 : purple
like lobes 2. Aedeagus longer than capsule 4. 3. Cuiller pointed Transtilla fused 1. Uncus with
With markedly pointed lobes 2. Aedeagus shorter folded than capsule accessory sac 3. Cuiller not pointed 4. Transtilla separate 1. Uncus with
2.
3. 4. 5.
pointed lobes Aedeagus shorter than capsule Cuiller not pointed Transtilla separate Setae digitate
With markedly folded accessory sac
Locality
A taxonomic revision of Stigmella
385
style.
Species-group PRUNIFOLIELLA
This group includes prunifoliella, intermedia, ceanothi and rhoifoliella and is characterized by external features, particularly the forewing markings, and t h e form of t h e uncus, gnathos and associated membrane in the male genitalia. Key to species Tufts o n front of head and vertex ochreous. Male with saccus weakly biolobed; uncus with papillate processes laterally. Female with accessory sac spinose.. . . . . . . . . . . . . . . . . . . . . . 2 Tufts on front of head and vertex dark brown. Male with saccus markedly bilobed; uncus pointed laterally. Female with accessory sac lacking spines 3 Cornuti of male long and thin; transtilla slender. Female with bursa heavily pectinate prunifoliella Cornuti of male short and s t o u t ; transtilla broad. Female with bursa weakly pectinate . . ceanorhi Forewing with sub-basal fascia dusted grey in appearance. Male with cornuti stout and relatively few in number . . . . . . . . . . . inrermedia Forewing without greyish sub-basal fascia. Male with numerous, slender cornuti . . . . rhoifoliella ~
~
Stigmella prunifoliella (Clemens) c0mb.n. (Fig. 19)
Nepticula prunifoliella Clemens, 1862a: 84 [description of mine] . Nepticula bifasciella Clemens, 1862b: 133 [description of adult] ; Braun, 1917: 170 [redescription and account of rearing work]. Syn.n. Stigmella bifasciella (Clemens); Wilkinson & Scoble, 1979: 59 [redescription]. Nepticula serotinaeella Chambers, 1873 : 126 [description of adult, mine and larva]. [Synonymized with bifasciella by Braun, 1917: 1701. DESCRIPTION. See Wilkinson & Scoble (1979). DIAGNOSIS. Differs from the closely related ceanothi in the cornuti of the vesica in the male genitalia, which are longer and thinner in prunifoliella; the transverse bars of the transtilla are stouter in ceanothi. The females of the t w o species are similar in all respects apart from t h e possible difference that the pectinations of the bursa copulatrix are more weakly sclerotized in ceanothi; the sample is very small, however, and this may not prove t o be a real species difference. The mines
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Philip J. Newton and Christopher Wilkinson
produced by larvae of prunifoliella differ from those produced by slingerlandella larvae o n Prunus sp., which make a true terminal blotch rather than a blotch formed by confluence of a linear tract. For other diagnostic features see Wilkinson & Scoble (1979), under bifasciella, and Fig. 19 here. DISCUSSION. The original description of this species is most unsatisfactory and has not been accepted by later workers. Clemens (1 862a: 84) described ‘Nepticula’ prunifoliella from untenanted mines on wild cherry, Prunus serotina, but introduced doubts as to identity by suggesting that the mines might be the product of dipteran larvae. Clemens (1 862b: 133) later described the species from a single adult, caught at light, which he named bifasciella; Chambers (1 873 : 126) also described it from adult, mine and larva with the name serotinaeella. Comparing the adult of serotinaeetta with the description of bifasciella and the mine with the description of prunifoliella, Chambers (1873; 1878b: 158) suggested that the names may be synonymous but did not feel able t o adopt either of those proposed by Clemens because of slight differences in the characters recorded. Clemens ( 1 86 2a) considered the mine of prunifoliella as ‘more or less blotchy from the beginning’ whereas Chambers observed that serotinaeella larvae appeared t o make a highly convoluted linear tract. Braun (1917: 170) was the first t o rear several specimens of the species and found that in many cases the mine is contorted to such a degree that a blotch is formed by confluence of the feeding tract. Thus she considered the evidence sufficient t o regard serotinaeella, bifasciella and prunifoliella as the same species. While proposing that serotinaeella become the junior synonym of bifasciella, she rejected the oldest name, prunifoliella, on the grounds of it being a nomen nudum. Unfortunately, however, prunifoliella remains an available name and cannot be ignored. Clemens was uncertain whether his prunifoliella was a dipteran. Thus, it is necessary to exclude the possibility that his mine could have been dipteran or, indeed, referable t o Hymenoptera or Coleoptera. There is indeed an agromyzid mine of Prunus serotina cur-
rently called Phytomiza persicae Frick. Having compared Clemens’ description with illustrations and description of persicae and bifasciella mines it seems, however, that prunifoliella must be referred to Stigmella. There is known no other species with which the description fits and there is little doubt that prunifoliella is thus the senior name for bifasciella. In order t o establish once and for all the position of prunifoliella a neotype has been designated. This neotype is also the holotype of Nepticula bifasciella Clemens.
DISTRIBUTION. U.S.A.: Pennsylvania, Ohio, New York, Kentucky. CANADA: Ontario. Type material examined Here designated, Neotype d = holotype of
c------(
0.02mm
u FIG. 18. Srigmella taeniola 0 , genitalia: (a) genitalia; (b) enlarged detail of pectinations on bursa copulatrix.
A taxonomic revision of Stigmella bifasciella, U.S.A., at light, 11 .viii (Clemens) (genitalia slide no. CNC 3465 ; ANS). Further material examined See Wilkinson & Scoble (1979). U.S.A.: Ohio, Cincinnati, 2 6 4 9 9 9 , 3 mines, o n Prunus serotina, 30.vi. 1909-13.v. 1922 (Braun), Adams County, 3 mines, on Prunus sp., 6 . x . 1927 (Braun), Salem Pike, 1 mine on Prunus sp., 26 .vii. 1913 (Braun);Kentucky, Magoffin County, 1 ex., on Prunus sp., 7.vi. 1932 (Braun) (ANS); Illinois, Putnam County, 11 Sd, 7 ?9,22.vi. 1956-23.vii. 1970 (Glenn);Massachusetts, Barnstable, 6 dd, 4 99, 2 ex., 6.vii.1952-18.vii.1958, 1 ex., 9.vii. ?1973 (Kimball) (USNM); Pennsylvania, West Chester, 12 mines, on Prunus pensylvanica, 1 7 . x . 1965 (CNC). CANADA: Ontario, Normandale, 10 mines, o n Prunus sp., 1 .vii. 1956; Ottawa, 1 mine, o n Prunus nigra, 3.viii. 1964 (Lewis)(CNC).
Stigmella ceanothi (Braun) c0rnb.n. (Figs. 20, 2 1 , 8 6 and 102) Nepticula ceanothi Braun, 1910: 1 7 2 ; Braun, 1917: 170 [redescription].
DESCRIPTION. External features: d? (Fig. 102). Palps greyish, antenna very dark brownish grey; tuft on front of head orange-ochreous,
387
vertex yellow; eyecaps shining white, sometimes with dark scales distally; collar dark grey-brown. Thorax dark grey-brown. Abdomen dark brown above, shining metallic grey beneath. Forewing: ground colour of dorsal surface dark brown with golden bronze reflections, when viewed at certain angle dusted with green and purplish reflections u p t o first fascia; fringe greyish brown, shining metallic grey; t w o narrow, shining silver fasciae, antemedial and postmedial, the latter sometimes extending toward apex marginally. Hindwing: ground colour and fringe greyish brown, shining metallic grey. Legs dark brown, shining metallic grey on lower surface. Wing measurements: d , 4.2-4.8 mm (6). 9,3.2-6.6 mm (5). Lectotype, 4.2 mm. Genitalia. d (Fig. 20). Uncus bilobed, in the form of a bridge, each lobe with small papillae laterally. Gnathos: broadly U-shaped; transverse ventral plate broad, convex medially; posteriorlydirected lateral arms very broad, n o t reaching beyond uncus; dorsolateral arms very broad with short posteriorlydirected processes. Tegumen a narrow, straplike plate with a spinose membrane ventral to uncus and tegumen as in Fig. 20(a). Vinculum: lateral arms very narrow, articulating with tegumen at dorsal extremities; ventral
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Philip J. Newton and Christopher Wilkinson
I-IG. 20. Stigmella ceanothi 15,genitalia: (a) genital capsule; (b) aedeagus; ( c ) valve; (d) detail of setae on style.
plate with very slight convex expansion medially o n posterior margin. Saccus less than half width of ventral plate, bilobed. Valve: narrow and tapering, extending just beyond uncus; bearing small, terminally digitate scales as in Fig. 20(d). Transtilla: lateral arms short; ventral arms long and tapering, reaching beyond ventral plate; transverse bars short and broad, meeting but not fusing. Juxta a triangular, membranous plate, more heavily sclerotized at margins, vertex articulating with posterior margin of ventral plate. Aedeagus: regular in width, three-quarters length of genital capsule; vesica with many denticulate cornuti, evenly distributed, and with a lunulate plate of minute papillae. 0 (Fig. 21). Ductus bursae short and narrow. Accessory sac very large, adorned with denticles distally and with accessory duct arising distally, serrate and laminate as in Fig. 21. Bursa copulatrix: very large, covered with
short chains of pectinations; signum absent. Anterior apophyses broad and long, slightly arcuate. Posterior apophyses with complex basal lobes as in Fig. 21, very long and straight, reaching beyond apophyses anteriores. BIOLOGY. Egg. Laid o n upper surface of leaf. Host. Ceanothus divaricatus. Five of the specimens examined, collected by H. H. Kelfer in 1926, were bred from a hybrid Ceanothus; we can find n o trace of the host material in order to confirm Kelfer's determination. Mine. Narrow, much contorted, linear, o n upper surface; approximately 3 cm in length, almost filled with evenly deposited frass (Fig. 86). Larva. Emerges from upper surface of leaf. Cocoon. Elliptical, reddish brown. Voltinism, Imagines have been taken in March, mid-April, mid-May and September, while the single recorded larva was taken in mid-February. It would appear that there are
A taxonomic revision of Stigmella two and possibly three broods per year; further collection and rearing is necessary before this may be confirmed. DIAGNOSIS. Differs from its nearest relative, prunifoliella, in that the cornuti are more numerous and smaller; the transverse bars of the transtilla are stouter in ceanothi. Both prunifoliella and ceanothi may be distinguished from intermedia and rhoifoliella by the paler tufts o n the head and the two strongly silver fasciae; the male genitalia differ in the form of the uncus, gnathos, valves and saccus, and
389
females differ in the spinose nature of the accessory sac. DISCUSSION. This species is notable for the wide range of the alar expanse shown, particularly in females. The three specimens labelled ‘Sept., Placer Co., Cal.’ are regarded as conspecific with the type material but differ in the following external features: t h e tufts on the front of the head and vertex are paler, as is the collar; the ground colour of the forewing is also paler and the iridescence more yellow-
FIG. 21. Stigmella ceanothi 0 , genitalia.
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Philip J. Newton and Christopher Wilkinson
I
I FIG. 22.
SrigmeIIu intermedia d , genitalia: (a)
gold with the absence of any purple reflections. Other specimens from Placer County have much darker external features. The atypical specimens have emergence dates in September, whereas others collected in the area emerged in March and April. Thus there may be seasonal variation in the external features of this species. Despite the very close similarity of this species to prunifoliella it is still thought to represent a separate species: the difference in host plants and the slight, but consistent morphological differences between the two species support this separation. DISTRIBUTION. U.S.A.: California. Type material examined Here designated, Lectotype d, U.S.A., California, Dutch Flat, Placer County, o n Ceanothus divaricatus, 18.iv. 1909 (Braun) (genitalia slide no. 12 PJN; ANS). Paralectotypes 2 dd, data as lectotype, 1 4 + 16 .ii. 1909
genital capsule; (b) aedeagus.
(Pilate) (genitalia slide nos. USNM 16207, USNM 16209; ANS). Further material examined U.S.A.: California, San Francisco, 1 d, 1 9 , bred from hybrid Ceanothus, 6 . iii. 1926 (Kelfer) (ANS), 1 d, 22 .v.? (Barber), 1 d , 2 99, bred from hybrid Ceanothus, 5 - 1 0 . 5 . 1 9 2 6 (Kelfer) (DFF), Placer County, 1 d, 299, 'Sept' n o other data (USNM), Dutch Flat, 1 mine, on Ceanothus divaricatus, 20.S. 1909 (Braun) (ANS).
Stigmella intermedia (Braun) (Fig. 22) Nepticula intermedia Braun, 1917: 171. Stigmella intermedia (Braun); Wilkinson & Scoble, 1979: 6 2 [redescription]. DESCRIPTION. (1 979).
See Wilkinson & Scoble
DIAGNOSIS. Differs from the very closely related rhoifoliella which has a basal patch of
A taxonomic revision of Stigmella gold reflections o n the forewing and lacks t h e sub-basal dusted fascia found in intermedia. The colour of the tufts o n the head and t h e single silver fascia differentiate this species from prunifoliella and ceanothi. The male genitalia of intermedia (Fig. 22) differ slightly from those of rhoifoliella in that there are probably fewer, stouter cornuti o n the vesica; t h e transverse bars of the transtilla may be stouter in rhoifoliella. The female genitalia of this species are indistinguishable from those of rhoifoliella. DISTRIBUTION. U.S.A.: Ohio, Arkansas, Kentucky (Braun). CANADA: Ontario. Material examined See Wilkinson & Scoble (1979). U.S.A.: ‘333u’ [ n o further d a t a ] , 1 9 , 1 ex., 1 6 + 20.iv. 1890 (MurfeEdt)(USNM); Ohio, College Hill Woods, 1 mine, o n Rhus typhina, 22.viii. 1908 (Braun), Rimington, 1 mine, on Rhus arornatica, 24.viii. 1916 (Braun) (ANS).
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CANADA, Ontario, Ottawa, 5 mines, on Rhus typhina, 8 . i x . 1955 (Freeman) (CNC).
Stigmella rhoifoliella (Braun) comb .n. (Figs.2 3 , 24,87 and 103) Nepticula rhoifoliella Braun, 19 12 1 93 ;Braun, 1917: 1 7 3 [redescription] ; Braun, in Forbes, 1923 : 87 [ redescription] . DESCRIPTION. External features. d (Fig. 103). Palps greyish; antenna dark brown; tufts o n front of head and vertex dark brown; eyecaps shining white; collar dark brown with purple reflections, sometimes pale. Thorax brown with purple reflections. Abdomen dark brown to black with scattered metallic reflections above, shining metallic grey beneath. Forewing: ground colour of dorsal surface dark brown with deep reddish bronze reflections when viewed at certain angles; fringe grey, pale off-white at apex, shining metallic grey; shining gold patch at basal third, costal margin shining purple, followed by purple
FIG. 2 3 . Sfigmella rhoifoliello d, genitalia: (a) genital capsule; (b) aedeagus.
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Philip J. Newton and Christopher Wilkinson
lustre before postmedial silver fascia, shining deep purple beyond. Hindwing: ground colour fringe pale grey brown, shining metallic grey. Legs brownish grey, shining silver on lower suf ac e . 9. As d except for pair of ventromedial fenestrae on fourth abdominal segment. Wing measurements. d, 3.2-4.2 mm (6). 0 , 3.6-3.8 mm (2). Lectotype, 4.0 mm. Genitalia. 6 (Fig. 23). Uncus bilobed, in the form of a bridge, each lobe broad and pointed. Gnathos: broadly U-shaped as in Fig. 23(a); transverse ventral plate narrow; posteriorly-directed lateral arms widely spaced, broad basally and tapering; dorsolateral arms folded and with short anteriorly-directed processes. Tegumen a narrow strap-like plate with weakly serrate posterior margin and with spinose membrane lying ventrally t o uncus and tegumen as in Fig. 23(a). Vinculum with
FIG. 24. Stigmella rhoifoliella 0 , genitalia.
broad lateral arms tapering markedly and articulating with tegumen at dorsal extremities. Saccus deeply bilobed. Valve broad basally, tapering markedly and reaching beyond uncus, inner surface concave. Transtilla: lateral arms very long; transverse bars narrow, pointed and at right-angle t o lateral arms, meeting but not fusing. Juxta a Yshaped plate, heavily sclerotized basally and along margins. Aedeagus: narrow and regular in width, half length of genital capsule; vesica with cornuti as many small denticles with larger spines posteriorly and with a bearded rectangular plate of minute papillae, 9 (Fig. 24). Ductus bursae long and narrow. Accessory sac small with reticulate patch medially and with duct arising proximally. Bursa copulatrix: large and covered with weakly sclerotized chains of pectinations; signum absent. Anterior apophyses broad basally, tapering, slightly arcuate. Posterior apophyses straight and of medium width, reaching beyond apophyses anteriores. BIOLOGY. The following details are collated from the literature and from Braun’s original breeding records. Egg. Laid o n upper surface of leaf, Host. Rhus toxicodendron (poison ivy). Mine. Much contorted, linear, o n upper surface of leaf; frass dispersed, in most places, throughout entire breadth of mine, granular (Fig. 87). 1,arva. Emerges o n upper surface of leaf, Cocoon. Reddish ochreous. Voltinism. Braun (1917: 173) reported that late instar larvae may be found in June, late July and September and suggested that there are three generations per year. The material examined here confirms that there are at least two generations, imagines being collected in June and particularly August. DIAGNOSIS. The external features differ from those of intermedia in the slightly paler forewing ground colour, the presence of the iridescent basal patch of gold found in rhoifoliella, and in the absence of the subbasal dusted fascia. The colour of the tufts on the head and the single silver fascia distinguish this species from prunifoliella and ceanothi. The male genitalia differ from those of intermedia in that the cornuti are smaller and more numerous and the transverse bars of the transtilla are stouter in rhoifoliella.
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fa ) I + fb) FIG. 2 5 . Stigmella rhamnicola d, genitalia summer generation: (a) genital capsule; (b) aedeagus. DISCUSSION. The relationship between rhoifoliella and intermedia is extremely close: they are regarded as sympatric even though the actual sites of collection in Ohio differ; the host plant differs only at the species level; there are slight but consistent morphological differences; specimens of rhoifoliella were found in August (with the exception of a single male) whereas intermedia has been collected largely from May to late July. It is possible, therefore, that what are here regarded as two species may prove to be a single species showing seasonal variation, or that the observed differences are due t o the effect of the different host. DISTRIBUTION- U.S.A.: (Braun).
Ohio, Kentucky
Type material examined Here designated, Lectotype d, U.S.A., Ohio, Cincinnati, o n Rhus toxicodendron,
23 .viii. 19 1 1 (Braun) (genitalia slide no. 36 PJN; ANS). Paralectotypes 4dd, 2 9 9 , 1 ex., data as lectotype, 22 + 23.viii.1911 (Braun) (ANS). Further material examined U.S.A.: Ohio, Cincinnati, 1 d, 9.vi. 1907 (Braun), 1 d, on Rhus toxicodendron, 16.viii. 1916 (Braun) (ANS). Of the following species, rhamnicola, diffasciae, gossypii and heteromelis possess several characters typical of the prunifoliella group, but differ in aspects of the male genitalia. The external features are similar to the prunifoliella group in the possession of two fasciae but in rhamnicola., diffasciae and "gossypii the proximal fascia is more off-white or yellow and the distal fascia silver.
Stigmella rhamnicola (Braun) c0mb.n. (Figs. 25-27,88,104 and 105)
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Nepticula rhamnella Braun, 1912: 96 [junior homonym of Nepticula rhamnella HerrichSchaffer, 1860: 601. Nepticula rhamnicola Braun,l916: 5 5 [replacement name] ; Braun, 1917: 172 [redescript i o n ] ; Braun, in Forbes, 1923: 8 7 [ redescription] DESCRIPTION. There are t w o infraspecific forms, the summer and winter generations. External features. Summer generation: d. Palps greyish; antenna dark brown; tufts on front of head and vertex ochreous; eye-caps shining white; collar off-white. Thorax offwhite with scattered silver reflections. Abdomen brownish grey above, paler and shining ~
metallic grey beneath. Forewing: ground colour of dorsal surface brown with purple and bronze reflections becoming predominantly bronze distally; fringe well defined at apex by a row of wing scales, brownish, shining white at apex; t w o fasciae, antemedial creamy white, shining yellow, postmedial silver and weakly concave. Hindwing: ground colour and fringe brownish grey, shining silver. Legs dark brown with various reflections above, shining metallic grey below, tarsi annulate with paler areas. 0 (Fig. 105). As d except for a pair of ventromedial fenestrae on fourth abdominal segment.
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Vinculum with narrow lateral arms expanding slightly dorsally and fusing with tegumen at dorsal extremities. Saccus very narrow, bilobed, each lobe as broad at base as ventral plate and as long as broad, lobes widely spaced. Valve with broad, bifid style, not reaching uncus, cuiller constricted. Transtilla: lateral arms long and broad; transverse bars continuous and at right-angles to lateral arms. Aedeagus: regular in width, just greater in length than genital capsule; vesica with cornuti as many short, stout denticles, evenly distributed, with a discrete patch at the phallotreme and with a patch of two or three very large, heavily sclerotized spines medially, also with a bearded rectangular plate of minute papillae. Winter generation (Fig. 26). As summer generation but: uncus slightly convex with lateral flattened lobes. Gnathos: as in Fig. 26(a); transverse ventral plate with medial convex expansion. Tegumen an arcuate, narrow strap. Vinculum with lateral arms tapering markedly t o point of articulation with tegumen at dorsal extremities. Aedeagus: regular in width, one-and-a-half times length
I
W
FIG. 2 7 . Stigmella rhamnicola 4, genitalia: (a) genitalia; (b) and (c) enlarged details of bursae; (b) pectinations of summer generation; (c) pectinations of winter generation.
Winter generation: d9 (Fig. 104). As summer generation but tufts o n front of head and vertex dark brown; collar dark brown. Wing measurements. d: (summer generation) 4.2-4.8 mm (3), (winter generation) 4.45.6 mm (1 1). 0 : (summer generation) 4.85.0mm (3), (winter generation) 4.6-5.6 mm (7). Lectotype, 4.6 mm. Genitalia. d. Summer generation (Fig. 25). Uncus bilobed, each lobe flattened and widely separated.Gnathos: as in Fig. 25(a): transverse ventral plate very narrow; posteriorlydirected lateral arms very broad basally, tapering markedly and stout; dorsolateral arms broad with short, stout posteriorly-directed processes. Tegumen a broad, horizontal bar.
.
.
la I
I
0.5 mm
I
I
0.05mm
I
FIG. 28. Stigmella specimens 1 and 2 , d abdomen (ventral) showing: (a) pairs of scent scale patches o n segments V , VI and VII; (b) enlarged detail of the scales.
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FIG. 29. Stigmella specimens 1 and 2, d genitalia: (a) genital capsule; (b) aedeagus.
of genital capsu1e;vesica with cornuti as many short, stout denticles, evenly distributed and with a bearded, rectangular plate of minute papillae. 0 . Summer generation (Fig. 27). Ductus bursae short and broad. Accessory sac very broad and tapering with duct arising distally. Bursa copulatrix: large and covered with weakly sclerotized chains of pectinations as in Fig. 27(b); signum absent. Anterior apophyses broad basally, tapering markedly and slightly arcuate. Posterior apophyses with basal lobe spinose, straight and narrow, reaching beyond apophyses anteriores. Winter generation (Fig. 27). As summer generation but bursa copulatrix covered with heavily sclerotized chains of pectinations as in Fig. 27(c). BIOLOGY. Egg. Laid o n lower surface of leaf. Host. Rhamnus lanceolata. Mine. Much contorted, linear initially, not visible from upper surface; later lengthening
and straightening, crossing t o upper surface, broadening markedly; frass deposited as continuous black line initially, becoming a granular arcuate line in later stages (Fig. 88). Larva. Emerges o n upper surface of leaf. Cocoon. Tawny. Voltinism. Mines have been collected in early July and October and are most abundant in October according to Braun (1917: 172). The material examined in this study includes t w o specimens which may represent the third generation that Braun mentions. DIAGNOSIS. The external features of both generations of rhamnicola differ from those of all other Stigmella listed here with t w o fasciae, except gossypii, in that the thorax is white. Summer and winter generations are best separated by the colour of the t u f t and collar of the head, the cornuti of the vesica in the male genitalia, and possibly by t h e degree of sclerotization of the pectinations of the bursa copulatrix in the female. The genitalia
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diffasciae are indistinguishable from those of the winter generation of rhamnicoh and both show slight but consistent differences from the male of the summer generation. The external features, mine and host plant of diffasciae differ from those of rhamnicola but are closely related. It is not unlikely that these three taxa represent different species but, until such time that rhamnicola may be bred through successive generations, we regard them as representing two species, o n e of which shows marked seasonal variation. There are two badly worn male specimens, collected in April 1916, which possess the genital characters of the summer generation but aspects of their external features are nearer to the winter generation; these specimens may represent a third generation.
of both generations differ from members of t h e prunifoliella group in the size of the aedeagus, the form of the uncus, and the continuous transverse bars of the transtilla in t h e male, and, in the females the spinose bases t o the apophyses posteriores. DISCUSSION. One male specimen, in the USNM, collected in 1916 bears a ‘cotype’ label in Braun’s handwriting; this specimen has not type status. The original description of rhamnicola was based o n imagines which emerged in late July and early August (the summer generation). Braun (1917: 172) later described the species as having two forms, the winter and summer generations, based o n additional material emerging in May (the winter generation). Braun also noted the similarity to the following species, diffQSCiQe.Examination of the genitalia of these two species has shown that the three taxa are very similar in form; t h e genitalia of
DISTRIBUTION. U.S.A.: Ohio. Type material examined Here designated, lectotype d, U.S.A., Ohio,
-
FIG. 30. Srigmellu inconspicuella d , genitalia: (a) genital capsule; (b) aedeagus.
398
Philip J. Newton and Christopher Wilkinson head and vertex dark ochreous; eyecaps shining white; collar off-white. Thorax and abdomen very dark brown with bronze reflections, abdomen shining metallic grey beneath. Forewing: ground colour of dorsal surface dark brown with bronze reflections towards the apex; fringe well defined at apex by a row of wing scales, greyish brown, shining silver when light at a suitable angle; two fasciae, antemedial creamy white, postmedial shining silver, narrow and extending t o apex at margins. Hindwing: ground colour and fringe greyish buff, shining silver. Legs brownish, shining metallic grey o n lower surface. 9 (Fig. 107). As d except for a pair of ventromedial fenestrae on fourth abdominal segment . Wing measurements. 9 , 5.4-5.8 mm (3). Lectotype, 5.6 mm. Genitalia. d (Fig. 26). Indistinguishable from the winter generation of Stigmella rhamnicola. 9 (Fig. 27). Indistinguishable from the winter generation of Stigmella rhamnicola.
0.02rnrn 1’’iG. 31. Stigmella inconspicuella 9 , genitalia: (a) genitalia; (b) enlarged detail of pectinations on
bursa copulatrix.
Cincinnati, bred from Rhamnus lanceolata, 26.vii. 1912 (Braun) (genitalia slide no. 63 PJN; ANS). Paralectotypes 3 dd, 2 99, data as 1 .viii. 1912 lectotype, 28 + 29.vii. 1912, (Braun)(genitalia slide nos. 64-67 PJN; ANSI. Further material examined U.S.A.: Ohio, Cincinnati, 8 dd, 5 99, bred from Rhamnus lanceolata, 14.iv. 1916-10.v. 1920 (Braun), Miami River, 1 mine,2cocoons, o n Rhamnus lanceolata, 5. x. 19 1 1,8 .vii. 19 12, 1 7 . x . 1915 (Braun) (ANS), 3 dd, 3 99, bred from Rhamnus lanceolata, 14-2O.v. 19 16 (Braun) (USNM), 1 9 , bred from Rhamnus lanceolata, 14.v. 1916 (Braun) (MCZ).
Stigmella diffasciae (Braun) c0mb.n. (Figs. 89 and 107) Nepticula diffasciae Braun, 1910: 172; Braun, 1917: 171. DESCRIPTION. External features. d. Palps grey; antenna dark brown; tufts o n front of
BIOLOGY. Egg. Laid on upper surface of leaf in three examples and on lower surface in the fourth. Host. Braun does not name the host in the original description but her breeding records say ‘the other species of Ceanothus’. Mine. Long, narrow, linear, o n upper surface of leaf. Tract contorted, turning several times in a small area. Frass deposited evenly throughout breadth of mine (Fig. 89). Larva. Emerges o n upper surface of leaf. Cocoon. Yellowish green. Voltinism. The relatively small sample of this species indicates a single generation per year; larvae collected in March produce adults in May. DIAGNOSIS. The external features differ from the summer and winter generations of rhamnicola in that the thorax is brown and the forewings darker in diffasciae. Male and female genitalia are indistinguishable from those of the winter generation of rhamnicola and in this respect the diagnosis for that form serves to differentiate diffasciae from related species. The mine of diffasciae differs from that of rhamnicola in its relatively even breadth and in the distribution of the frass. DISCUSSION. As discussed in the treatment
A taxonomic revision o f Stigmella
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I
FIG. 32. StigmelIa gossypii d, genitalia: (a) genital capsule; (b) aedeagus.
of Stigmella rhamnicola, the two species show a very complex relationship: emergence dates are similar for this species and the winter generation of rhamnicola; o n the basis of present records they are allopatric, differ in host plant and mine characteristics. DISTRIBUTION. U.S.A.: California. Type material examined Here designated, Lectotype d, U.S.A., California, Dutch Flat, Placer County, bred from Ceanothus sp., 8 . v . 1909 (Braun) (genitalia slide no. 1 PJN; ANS). Paralectotypes 16, 3 9 9 , 1 ex., data as lectotype, 8-26.v. 1909 (Braun) (genitalia slide nos. 2-3 PJN, USNM 16205-16206; ANS). Further material examined U.S.A., California, Dutch Flat, Placer County, 4 mines, 2 cocoons, o n Ceanothus sp., 9.iii. 1909 (Braun) (ANSI. Specimens 1 and 2 (Figs. 28 and 29)
Two male specimens show similarities to rhamnicola and diffasciae in external features, 21
genital morphology and host plant. Although there are features by which these specimens may be separated from these species and from other members of the genus listed here, they are insufficient to regard the specimens as representing a new species. DESCRIPTION. External features. d. Worn specimens allow a tentative description only - palps buff; antenna dark brown; tuft on front of head ochreous, vertex darker; eyecaps shining white; collar off-white. Thorax and abdomen greyish with scattered silver reflections. Abdomen with three pairs of scale patches o n ventral surface of segments V, VI and VII, each scale heavily sclerotized and flattened distally as in Fig. 28. Forewing: ground colour of dorsal surface brown with weak bronze lustre; fringe greyish; two fasciae, antemedial broad and creamy white, postmedial silver. Hindwing: ground colour and fringe greyish brown, shining metallic grey. Legs brown with scattered paler areas. Wing measurements. d, 3.8-4.8 mm (2).
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Genitalia. d (Fig. 29). Uncus very large and bilobed, each lobe flattened. Gnathos: as in Fig. 29(a); transverse ventral plate narrow; posteriorly-directed lateral arms short and broad ;dorsolateral arms with long posteriorlydirected processes. Tegumen a broad horizontal strap. Vinculum: lateral arms broad, fusing with tegumen at dorsal extremities; ventral plate broad. Saccus narrow, bilobed with each lobe broader than ventral plate at base and as long as broad. Valve divided into long, broad style and broad constricted cuiller. Transtilla continuous, in the form of a large U-shaped bar reaching beyond the saccus. Aedeagus: broad and ovate approximately equal t o length of genital capsule; vesica with cornuti and
many small denticles evenly distributed with scattered large denticles, and with a lunulate plate of minute papillae. BIOLOGY. Egg. Laid o n lower surface of leaf. Host. Rhamnus caroliniana and R.californica. Mine. On Rhamnus californica an upper surface, small and much contorted linear mine; frass deposited in arcs almost the breadth of the mine. Braun's rearing notes suggest that the mine o n Rhamnus caroliniana is longer with the frass scattered across its breadth. Larva. Emerges o n upper surface of leaf. Voltinism. The Ohio specimen was bred from a larva collected in November, the adult emerged the following May. The California specimen was reared from a larva taken in July, and emerged the following month. DIAGNOSIS. The genitalia of these two specimens differ from those of rhamnicola and diffasciae males in the arcuate form of the transtilla and the size and lobing of the uncus; the paired scale patches of the abdomen are not found in either species and are easily distinguished from the single pockets in aromella. DISCUSSION. These two specimens closely resemble rhamnicola and diffasciae, the most significant difference being the presence of the abdominal patches in the males. These scale patches are assumed t o be scent producing or distributing organs. The specimen from California differs from the other in the form of the uncus, which is much larger, and the choice of host, Rhamnus californica as. opposed t o R.caroliniana. DISTRIBUTION. U.S.A.: Ohio, California.
Material examined Specimen 1 , d, U.S.A., Ohio, Adams County, o n Rhamnus caroliniana, 5 .v. 1941 (Braun) (ANS). Specimen 2, d, U.S.A., California, Yosemite National Park, o n Rhamnus californica, 2 0 . v l i . 1 9 1 5 (Braun), 1 mine, same data, 26.vii. 1915 (Braun) (ANS).
Stigmella inconspicuella sp.n. (Figs. 30, 3 1 and 106) DESCRIPTION. External features. 6. Palps greyish; antenna dark grey-brown; tufts on FIG. 33. Stigmella gossypii 0 , genitalia.
front of head and vertex ochreous, vertex
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FIG. 34. Stigmella heteromelis d , genitalia: (a) genital capsule; (b) aedeagus; ( c ) valve.
usually brown distally; eyecaps shining white; collar off-white. Thorax and abdomen dark grey-brown with bronze and purple reflections above, abdomen shining metallic grey beneath. Forewing: ground colour of dorsal surface dark brown with bronze reflections, lustrous
grey when viewed at certain angles; fringe greyish brown, shining metallic grey. Hindwing: ground colour and fringe grey-brown, shining metallic grey. Legs greyish brown with scattered ochreous patches, shining metallic grey o n lower surface.
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0 (Fig. 106). As d except for a pair of ventromedial fenestrae o n fourth abdominal segment. Wing measurements. d, 5.4 mm (1). 0 , 4.8 mm (1). Holotype, 5.2 mm. Genitalia. d (Fig. 30). Uncus very broad
and quadrate, with small lobes laterally. Gnathos: as in Fig. 30(a); transverse ventral plate broad; posteriorly-directed lateral arms closely juxtaposed at base, broad and tapering; dorsolateral arms with short posteriorlydirected processes and long anteriorly-directed
on bursa
A taxonomic revision o f Stigmella processes. Tegumen a broad arcuate plate with serrate posterior margin. Vinculum: lateral arms broadening at point of articulation with tegumen at dorsal extremities; ventral plate broad. Saccus very narrow, markedly bilobed, each lobe longer than broad at base. Valve with broad cuiller and short pointed style, not reaching uncus. Transtilla: lateral arms long and narrow; transverse bars continuous, very broad and at right-angles to lateral arms. Aedeagus: flask-shaped, as long as genital capsule; vesica with cornuti as long, thin denticles orientated in a ridge posteriorly and with a bearded rectangular plate of minute papillae. 9 (Fig. 3 1). Ductus bursae long. Accessory sac small and membranous with broad, spiral duct arising proximally. Bursa copulatrix: small and covered with long, weakly sclerotized chains of pectinations; signum absent. Anterior
403
apophyses very broad basally, tapering markedly. Posterior apophyses straight and narrow, extending beyond apophyses anteriores. BIOLOGY. Host. Ceanothus arboreus. Cocoon. Spun o n lower surface of leaf, obovoid, buff. DIAGNOSIS. The bronze reflections of the forewings distinguish inconspicuella from chalybeia, with a blue-grey lustre, and pomivorella, with deep purple reflections. The male genitalia are similar to those of chalybeia, rhamnicola and diffasciae but the quadrate uncus, the more concave saccus and the broad transtilla differentiate inconspicuella. The broad accessory duct in inconspicuella separates the female from chalybeia and pomivorella. DISCUSSION. This species is externally similar t o chalybeia and pomivorella, but is placed here because the genitalia more closely resemble those of rhamnicola and diffasciae (which also mines Ceanothus). DISTRIBUTION. U.S.A.: California. Type material Holotype 8, U.S.A., California, Berkeley, o n Ceanothus, 15.v. 1929 (Jones) (genitalia slide no. 16269; USNM). Paratypes 1 d, 1 9 , 1 ex., same locality, o n Ceanothus arboreus, 10 + 12.v. 1929 (Braun) (genitalia slide nos 87 PJN, USNM 16213; ANS). Material excluded from type-series 1 cocoon, o n Ceanothus arboreus, n o further data (Braun) (ANS). Specimen 3
(b) M
002 m m FIG. 3 6 . Stigmella cerea 9 , genitalia: (a) genitalia; (b) enlarged detail of pectinations on bursa copulalrix.
There is a single d with genitalia indistinguishable from those of inconspicuella except that the uncus is papillate and the valves broader in relation t o the genital capsule. The specimen is damaged so its external similarity t o inconspicuella is uncertain. The label data d o not make clear whether this specimen was reared from a larva mining cotton; from a pupa in cotton leaf-litter or was collected as an imagine which had alighted on cotton. Material examined d, U.S.A., Florida, Miami, in cotton leaves, 20.iv. 1933 (Rainwater) (genitalia slide no. 16292; USNM).
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fa)
t
1
FIG. 31. Stigmella rosaefoliella pectocatena d , genitalia: (a) genital capsule; (b) aedeagus.
Stigmellagossypii (Forbes & Leonard) c0rnb.n. (Figs. 3 2 , 3 3 and 108) Nepticula gossypii Forbes & Leonard, 1930: 149. DESCRIPTION. External features. 6. Palps whitish; antenna dark brown; tufts on front of head and vertex ochreous; eyecaps shining creamy white; collar white. Thorax grey with metallic reflections. Abdomen dark brown, shining metallic grey beneath. Forewing: ground colour of dorsal surface dark brown with purple and silver reflections; fringe well defined at apex by row of wing scales, greyish, shining metallic grey; two fasciae, antemedial shining gold, broadening on posterior margin, postmedial shining silver, narrower. Hindwing: ground colour and fringe grey, shining metallic grey. Legs brown, shining metallic grey on lower surface. 9 (Fig. 108). As male except for a pair of ventromedial fenestrae o n fourth abdominal segment. Wing measurements. d, 3.0 mm (1). 0 , 3.03.2 mm (5). Holotype, 3.0 mm. Genitalia. d (Fig. 32). Uncus very large and quadrate with two lateral papillae. Gnathos: broadly U-shaped as in Fig. 32(a); transverse ventral plate narrow; posteriorlydirected lateral arms long and narrow, widely spaced ;dorsolateral arms with short anteriorlydirected processes. Tegumen a narrow,
horizontal bar with serrate posterior margin and with a spinose membrane lying between lateral arms of gnathos as in Fig. 32(a). Vinculum: lateral arms bifurcate distally, articulating with tegumen and with posterior branches fusing dorsal t o tegumen as in Fig. 32(a); ventral plate with weakly concave posterior margin. Saccus narrow, deeply bilobed. Valve tapering markedly and bifid distally, just reaching uncus, weakly concave along inner margin. Transtilla: lateral arms short and narrow; ventral arms stout; transverse bars short and stout, meeting but not fusing. Juxta a quadrate, membranous plate. Aedeagus: flask-shaped, approximately equal in length t o genital capsule; vesica with cornuti as many small denticles orientated in a small ridge posteriorly and with a lunulate plate of minute papillae. 0 (Fig. 33). Ductus bursae short and broad. Accessory sac large with a reticulate patch at vestibule and with duct arising distally. Bursa copulatrix: large, covered with weakly sclerotized chains of pectinations; signum absent. Anterior apophyses long and narrow. Posterior apophyses straight and narrow, reaching just beyond apophyses anteriores. BIOLOGY. The following notes are collated from the comprehensive account of the lifehistory by Forbes and Leonard. Egg. Laid on lower surface of leaf and, as far as observed, in the fork of small leafveins. Host. The mines are found in great numbers on G.6erbadense (sea island cotton) at the type locality. Forbes and Leonard were unsuccessful in their attempts t o record the species in other localities in Puerto Rico and on the other major cotton plant, G.hirsutum (tree cotton), apart from a single untenanted mine. Mine. Slender, sinuous, on lower surface but visible from upper surface of leaf. In first half of mine frass deposited in arcuate waves and is granular while in latter half it is a single, continuous line, Several mines on a single leaf are observed t o cross and anastomose frequently. Larva. Green, emerging o n lower surface of leaf. Cocoon. Oval, flattened, white; o n lower surface of leaf. Voltinism. Forbes and Leonard reported
A taxonomic revision of Stigmella that tenanted mines were only located in March and early April, imagines emerging approximately 10 days later. They noted that there appeared to be a single ‘active’ mining period in these months, indicating that there is probably one generation per year. DIAGNOSIS. Differs from rhamnicofa and diffasciae in the purple reflections of the forewings and the golden rather than white proximal fascia in gossypii. The males differ from those of rhamnicola and diffasciae particularly in the form of the uncus and aedeagus, but also t o a lesser extent in the gnathos, bifurcate style of the valve and the two halves of the transtilla which meet b u t d o not fuse in gossypii. DISCUSSION. This species is morphologically related to rhamnicola and diffasciae and to members of the prunifoliella group, but is much smaller. DISTRIBUTION. U.S.A.: Puerto Rico, Florida. Type material examined Holotype 9, U.S.A., Puerto Rico, Juana Diaz, o n Gossypium sp., 13.iv. 1930(leonard) (genitalia slide no. DRD 3 100; USNM). Paratypes 6 ex., data as holotype, April 1930 (Leonard) (genitalia slide no. 9 DRD 3101; USNM).
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medial fascia white. Hindwing: ground colour and fringe brownish grey, shining silver grey. Legs dark grey, shining metallic grey. Wing measurements. d, 5 . 2 - 5 . 5 mm ( 8 ) . 9, 5.0-6.5 mm (19). Holotype, 5.4 mm. Genitalia. d (Fig. 34). Uncus broad and flattened, weakly bilobed. Gnathos: narrowly U-shaped as in Fig. 34(a); transverse ventral plate broad; posteriorly-directed lateral arms very broad ; dorsolateral arms broad basally with very short posteriorly-directed processes. Tegumen broad and ovate with spinose membrane lying ventrally t o uncus and tegumen as in Fig. 34(a). Vinculum: lateral arms tapering markedly, bifurcate dorsally and articulating with tegumen, with posterior branches dorsal t o tegumen as in Fig. 34(a); ventral plate
Further material examined U.S.A., Florida: Fort Myers, 1 d, 3 99, o n Gossypium sp., no date (Rainwater); Miami, 1 d , 2 9 9 , o n Gossypium sp., 20.iv. 1933 (Rainwater) (USNM).
Stigmella heteromelis sp.n. (Figs. 34, 3 5 and 109) DESCRIPTION. External features. d9 (Fig. 109). Palps brownish grey; antenna very dark brown; tuft o n front of head ochreous, becoming paler and buff laterally o n the vertex; eye-caps shining creamy-white; collar offwhite. Thorax and abdomen very dark brown, abdomen with gold reflections above, shining metallic grey beneath. Forewing: ground colour of dorsal surface dark brown, basal third with gold and silver reflections becoming bronze and purple toward the apex; fringe greyish brown, shining silver at apex; basal patch of iridescence forms a broad, indistinct fascia with extreme base of wing brown, post-
a) w FIG. 3 8 . Stigmella rosaefoliella pectocatena 0 , genitalia: (a) genitalia; (b) and (c) enlarged detail of pectinations o n bursa copulatrix; (b) subordinate subspecies; (c) nominate subspecies.
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FIG. 39. Stigmello slingerlondella d, genitalia: (a) genital capsule; (b) aedeagus.
broad, convex mediaIly. Saccus narrow and bilobed. Valve very broad, bifurcate distally with style pointed and not reaching uncus, cuiller constricted and broadly pointed terminally. Transtilla: W-shaped as in Fig. 34(a); lateral arms long and thin; ventral arms broad and tapering; transverse bars continuous and narrow. Juxta a broad, triangular membranous plate. Aedeagus: very broad and regular, threequarters length of genital capsule; vesica with cornuti as many long, thin denticles and with collar-like plate of minute papillae. 9 (Fig. 35). Ductus bursae short and narrow. Accessory sac large, with reticulate patch at vestibule and with patch of heavily sclerotized denticles lateral t o origin of medial accessory duct. Bursa copulatrix : very large with medial third covered in scallop-shaped chains of pectinations as in Fig. 35(b); signum absent. Anterior apophyses short and tapering. Posterior apophyses straight and narrow, reaching beyond apophyses anteriores.
BIOLOGY. Host. Heteromeles sp., and He terom eles arbu tifolia. Voltinism. There is no indication of the
number of generations per year in the wide range of emergence dates recorded.
DIAGNOSIS. The diffuse nature of the forewing basal patch separates this species from those with t w o fasciae, especially rhamnicola, diffasciae and gossypii. The wing markings are similar to those of infermedia and rhoifoliella, but heteromelis differs from these species in the ochreous tufts o n the head and in t h e larger alar expanse, The genitalia may be easily separated from rhamnicola, diffasciae and gossypii by the form of the valve, uncus and the long cornuti of the vesica in the male, and by the spinose patches of the accessory sac and the band of pectinations on the bursa copulatrix in the female. DISCUSSION. This species is described partly from a series collected by Braun and W. W. Jones. Some were labelled ‘Stigmella photiniae’ by Jones; however, this name has not been found in the literature. Photinia is a junior synonym of Heteromeles - the food plant. This species shows similarity to all those species with t w o fasciae listed here, especially
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in the male genitalia and the choice of host plant. DISTRIBUTION. U.S.A.: California. Type material Holotype d, U.S.A., California, Berkeley, U.C. Campus, o n Heteromeles [ = Photinia] arbutifolia, 2 2 . i. 196 1 (Powell) (USNM). Paratypes 6 dd, 10 99, locality as holotype, 23.i. 1960-22.i. 1961 (Powell);Berkeley, 1 9 , on Heteromeles [= Photinia] , 22.v. 1929 (Jones) (genitalia slide no. 17224; USNM), 9dd, 1199, 6.iii-10.v.1929 (Braun) (genitalia slide nos. 4-9 PJN, USNM 17336; ANS).
Stigmella cerea (Braun) combm. (Figs. 36 and 110) Nepticula cerea Braun, 19 17 : 172 ; Braun, in Forbes, 1923 : 8 7 [ redescription] . DESCRIPTION. External features. 9 (Fig. 110). Palps whitish buff; antenna pale brown; tufts on front of head and vertex pale ochreous; eye-caps shining white; collar off-white. Thorax greyish buff. Abdomen dark brown above, shining metallic grey beneath and with a pair of ventromedial fenestrae o n fourth abdominal segment. Forewing: ground colour of dorsal surface dark brown with bronze reflections; fringe well defined at apex by a row of wing scales, grey, shining white at apex; basal patch of creamy white scales extending half length of t h e wing, broken by two triangular, brown patches at extreme base of each margin in the holotype, followed by single postmedial fascia, shining silver, broadening on dorsal margin. Hindwing: ground colour and fringe greyish brown, shining metallic grey. Legs brownish with scattered grey areas, shining metallic grey. Wing measurements. 9, 3.5 mm (1). Holotype, 3.6 mm. Genitalia. 9 (Fig. 36). Ductus bursae long and very broad with duct arising proximally. Bursa copulatrix : covered with irregular chains of pectinations medially, as in Fig. 36(b), becoming less dense distally; signum absent. Anterior apophyses broad basally, tapering and straight. Posterior apophyses very narrow and straight, reaching just beyond apophyses anteriores. BIOLOGY. Immature stages and host unknown.
FIG. 40. Stigmella slingerlandelia 0 , genitalia: (a) genitalia; (b) enlarged detail of pectinations o n bursa copulatrix.
DIAGNOSIS. The external features are similar to those of heteromelis but this species may be separated by the irregular basal patch o n the forewing. The genitalia also resemble heteromelis but the nature and distribution of t h e pectinations differ, and cerea lacks the large, spiculate accessory sac found in heteromelis. DISCUSSION. In the absence of any male representatives cerea has been placed here because of t h e similarities t o the preceding species. DISTRIBUTION. U.S.A.: Ohio, Pennsylvania.
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Type material examined Holotype 0, U.S.A., Ohio, Cincinnati, 28.viii. 1909 (Braun) (genitalia slide no. 10 PJN; ANS). Paratypes 10, data as h o l o t y p but 15.vi. 1909 (Braun) (ANS); 1 9 , Pennsylvania, Oak Station, Allegheny County, 2 5 . v . 1910 (Marloff) (MCZ). Species-group ROSAEFOLIELLA
The rosaefoliella group, as described by Wilkinson & Scoble (1 979) included the species rosaefoliella, slingerlandella, fuscotibiella, populetorum and aromella. The form of the uncus, gnathos and aedeagus are characteristic of the male genitalia and in this respect the group is homogeneous. The female genitalia, on the-otherhand, are of two types: with a signum comprising a U-shaped band of pectinations, or without. This difference in the female genitalia corresponds to a difference in the family of host plant mined by the larvae: Salicaceae and Rosaceae respectively. F o r these reasons a separate group is reserved for
fuscotibiella, populetorum and aromella in which the females lack a signum. Key to species Male with transtilla fused; gnathos with dorsolateral arms juxtaposed. Larva making an op hist igm at onome . . . . . . . . . slingerlandella Male with transtilla usually not fused; gnathos with dorsolateral arms widely separated. Larva making an ophionome . . . . . . (rosaefoliella)’ Male with small, spiculate cornuti. Female with scallop-shaped pectinations of the bursa r.rosaefoliella Male with large, denticulate cornuti. Female with long chains of pectinations on the bursa r.pectocatena
Stigmella rosuefoliellu (Clemens) Nepticula rosaefoliella Clemens, 1862a: 85 [description of mine and larva] ; Braun, 19 12: 8 9 [description of adult] . Stigmella rosaefoliella (Clemens); Wilkinson & Scoble, 1979: 14 [redescription, neotype designated] DISCUSSION. See Wilkinson & Scoble (1 979).
FIG. 41. Srigrnella villosella d , genitalia: (a) genital capsule; (b) aedeagus.
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I t is regarded as being represented by two subspecies. DIAGNOSIS. Differs from slingerlandella in the darker abdomen and the white, rather than silver, fascia of the forewing in rosaefoliella. The male genitalia differ in the form of t h e uncus and gnathos and in t h e transverse bars of t h e transtilla which are always continuous in slingerlandella. The ophionome of rosaefoliella differs from the ophistigmatonome made b y larvae of slingerlandella.
Stigmella rosaefoliella rosaefoliella (Clemens) Stigmella rosaefoliella rosaefoliella (Clemens); Wilkinson & Scoble, 1979: 17.
DIAGNOSIS. The nominate subspecies may be distinguished from rosaefoliella pectocatena by the scallop-shaped pectinations of the bursa copulatrix in the female (Fig. 38c) and by the spiculate rather than spine-like cornuti of the aedeagus in the male. DISTRIBUTION. U.S.A. : Ohio, Pennsylvania, Arkansas, New York, Michigan, Missouri. CANADA: Ontario. Material examined See Wilkinson & Scoble (1979) (including ‘specimen a’). U.S.A.: Ohio, 3 mines, o n Rosa setigera, 7. vii. 1909, 20. x. 19 1 2 (Braun), no locality data, 1 mine, o n Rosa sp., 25.vi. 1909 (Braun) (ANS); Pennsylvania, West Chester, 7 mines, o n Rosa sp., 1965 (Lewis) (CNC). CANADA, Ontario, Simcoe, 7 mines, on Rosa sp., 1969 (Lewis) (CNC).
Stigmella rosaefoliella pectocatena Wilkinson & Scoble (Figs. 37 and 38) Stigmella rosaefoliella pectocatena Wilkinson & Scoble, 1979: 18.
DIAGNOSIS. Differs from the nominate subspecies in the form of the pectinations of the bursa copulatrix in the female and in the spine-like cornuti of the aedeagus in the male (Figs. 3 7 and 38). Wilkinson & Scoble (1979) also distinguish this subspecies from t h e European species anomalella (Goeze). DISTRIBUTION. CANADA: Ontario. Material examined See Wilkinson & Scoble (1979). Mines of nominate and subordinate subspecies are indistinguishable : material is placed here on the basis of locality alone. CANADA, Ontario,
FIG. 42. Stigmella villosella 9 , genitalia: (a) genitalia; (b) enlarged detail of spines on ductus bursae.
Ottawa, 8 mines, on Rosa sp., 20.ix.1955 (Lewis) (CNC). See also Wilkinson and Scoble ( 1979) for specimen b.
Stigmella slingerlandella (Kearfott) (Figs. 1, 39 and 40) Nepticula slingerlandella Kearfott, 1908: 187. Stigmella slingerlandella (Kearfott); Wilkinson & Scoble, 1979: 1 9 [redescription, lectotype designated].
DESCRIPTION. See Wilkinson & Scoble (1979). This species is closely related to the North American species rosaefoliella and the European plagicolella (Stainton).
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FIG. 43. Stigmelta apicialbella d , genitalia: (a) genital capsule; (b) aedeagus; ( c ) valve.
DIAGNOSIS. Differs from rosaefoliella in the narrower fascia of the forewing. The male genitalia differ in the nature of the papillae of the uncus, the horns of t h e gnathos and t h e transverse bars of the transtilla which d o not always fuse in rosaefoliella. The larva of slingerlandella forms an upper surface ophistigmatonome whereas rosaefoliella mines lack the terminal blotch as do those on Prunus produced by prunifoliella larvae. Wilkinson & Scoble (1 979) also distinguish between this species and plagicolella (Stainton). (See Figs. 3 9 and 40.) DISTRIBUTION. U.S.A.: New York, Michigan, Ohio, ?Florida. CANADA: Ontario. Material examined See Wilkinson & Scoble (1979). U.S.A., Ohio: West Fork Woods, 1 mine, on wild plum, 6 . i x . 1909 (Braun); North of Madeira, 1 mine, on Prunus sp., 30.viii. 1909 (Braun) (ANS). CANADA, Ontario: Elmira, several
mines and larvae in vials, on Prunus sp., 17.viii. 1966 (FIS); Normandale, 4 mines, on Prunus nigra, 3O.viii. 1956 (Freeman and Lewis), 4 mines, o n Prunus serotina, 15.viii. 1957 (Freeman and Lewis); Simcoe, 4 mines, on Prunus serotina, 15.viii. 1966 ( L e w i s ) (CNC). Specimen 84. A single female specimen from Florida may belong here or represent a new allied species. It has t h e following data: Florida, Monroe Co., Key Largo, 2 8 . x . 1972 ( K e m p1.
Stigmella villosella (Ciemens) c0mb.n. (Figs. 4 1 , 4 2 , 9 0 a n a 111) Nepticula villosella Clernens, 1862a: 84 [description of mine and larva] ; Clemens, in Stainton, 1872: 174; Chambers, in Hayden, 1878b: 1 5 8 [listed] ; Braun, 1912: 94 [description of adult]; Braun, 19 17 : 174 [ redescription] ; Braun, in Forbes, 1923: 88 [redescription].
A taxonomic revision of Stigmella Nepticula dallasiana Frey & Boll, 1876: 288 [synonymized by Braun, 19 12: 941 . DESCRIPTION. External features. d? (Fig. 11 1). Palps grey; antenna dark brownish grey; tufts o n front of head and vertex ochreous; eye-caps shining white; collar dark brown with gold reflections. Thorax and abdomen very dark brown, thorax with gold reflections, abdomen with purple reflections above, shining metallic grey beneath. Forewing: ground colour of dorsal surface dark brown,
FIG. 44. Stigmella apicialbella 9, genitalia.
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shining deep gold before fascia, deep purple beyond; fringe brown, shining silver at apex and resembling an apical spot; single postmedial fascia, shining silver, sometimes slightly wider at margins. Hindwing: ground colour and fringe pale greyish brown, shining silver. Legs very dark brown t o black, shining metallic grey on lower surface. Wing measurements. d, 2.8-4.6 mm (7). 9 , 4.0-4.4 mm (4). Neotype, 4.0 mm. Genitalia. d (Fig. 41). Uncus bilobed, each lobe flattened, with a deep notch between. Gnathos: as in Fig. 41(a); transverse ventral plate broad; posteriorly-directed lateral arms very closely juxtaposed basally, narrow and long; dorsolateral arms broad with long posteriorly-directed processes and short, stout anteriorly-directed processes. Tegumen a narrow arcuate strap. Vinculum with lateral arms bifurcate dorsally, articulating with tegumen and posterior branches fusing dorsal t o the tegumen as in Fig. 41(a). Saccus broad, weakly bilobed. Valve narrow with stout, pointed style and cuiller tapering anteriorly. Transtilla: lateral arms broad and long: ventral arms short and pointed; transverse bars fused and narrow. Aedeagus: flaskshaped, longer than capsule; vesica with cornuti as many large denticles orientated in a ridge posteriorly with t w o more heavily scierotized spines apically and with a globular plate of minute papillae. 0 (Fig. 42). Ductus weakly funicular
FIG. 45. Stigmella fuscotibiella d, genitalia: (a) genital capsule; (b) aedeagus.
4 12
Philip J. Newton and Christopher Wilkinson berry (Rubus spp.) but occasionally on wild raspberry (Rubus occiden talis). Mine. Long, very narrow, serpentine tract on upper surface of leaf; frass deposited centrally as a dense black line becoming more diffuse in distal fifth (Fig. 90). Larva. Pale brown in colour, emerging o n the upper surface of t h e leaf. Cocoon. Brown. Voltinism. Braun ( 1 9 12) reported that late instar larvae are found in Cincinnati in the middle of June, latter part of July and October. Imagines have been collected in May and late June t o t h e middle of July, indicating that there are certainly two and possibly three generations per year.
...
FIG. 46. Stigmella fuscotibiella 9 , genitalia.
proximal to the genital aperture, long and broad. Accessory sac very large relative t o the bursa and adorned with small denticles which become more dense distally and with a duct arising distally. Bursa copulatrix: large, covered with irregular chains of pectinations proximally, as in Fig. 42(b), which are not apparent distally; signum absent. Anterior apophyses broad basally, long and tapering. Posterior apophyses very long and straight, reaching beyond apophyses anteriores. BIOLOGY. Egg. Laid on upper o r lower surface of leaf; in the material examined here it is always next to a leaf vein. Host plant. Braun (1912: 95) reported that t h e species is most commonly found on black-
DIAGNOSIS. The external features differ from those of rosaefoliella and slingerlandella in t h e degree of iridescence of the dorsal and ventral aspects, particularly in the apical patch of bright silver and the basal gold reflections o n the forewings of villosella. The flaskshaped aedeagus of the male genitalia and the many small cornuti arranged in a ridge differentiate this species from rosaefoliella and slingerlandella, while the form of the valve differs in apicialbella. The presence of a large, spiculate accessory sac in the female genitalia distinguishes villosella from rosaefoliella and slingerlandella. Females of villosella and apicialhella resemble each other in aspects of the genitalia but differ in their external features. The mine produced by the larvae of villosella differs from others o n Rubus spp., produced by the larvae of Ectoedemia rubifoliella, in the absence of the terminal blotch. DISCUSSION. Clemens named this species from mine and larva only. As Braun (1912: 94) indicated, it was later described under the name dallasiana by Frey & Boll (1876: 228) from a single specimen bred from blackberry in Texas: this holotype has not been traced despite efforts t o locate it among the Frey collection at the BMNH and in the MCZ where Zeller noted that many specimens of the Frey collection were deposited. This species is placed here largely o n the grounds that larvae mine members of the Rosaceae. The form of the aedeagus in the male and the ornamentation of the accessory sac of the bursa copulatrix of the female exclude viflo-
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sella from the rosaefoliella group but relate it closely t o the following species, apicialbella. DISTRIBUTION. U.S.A.: Texas, Ohio, Kentucky (Braun), Pennsylvania (Clemens). Type material examined Here designated, Neotype d, U.S.A., Ohio, Cincinnati, on Rubus sp., 1 2 ,v . 196 1 (Braun) (ANS). Further material examined U.S.A., Ohio, Cincinnati, 8 dd, 4 9 9 , on Rubus sp., 23.vii. 1909-3.v. 1919 (Braun), 2 mines, on Rubus occidentalis, 3.vii. 1909 (Braun) (ANS).
Stigmella apicialbella (Chambers) (Figs. 43, 4 4 , 9 1 and 113) Nepticula apicialbella Chambers, 1873: 127; Chambers, in Hayden, 1878b: 157 [listed] ; Braun, 1914: 21; Braun, 1917: 182 [redescription] ; Braun, in Forbes, 1923: 90 [ redescription] . Stigmella apicialbella (Chambers); Wilkinson & Newton, 1981: 54. Nepticula leucostigma Braun, 1912: 88 [synonymized by Braun, 19 14: 2 1 1 . DESCRIPTION. External features. d? (Fig. 113). Palps whitish; antenna dark brown; tufts on front of head and vertex ochreous; eye-caps shining white; collar white. Thorax and abdomen very dark brown with purplish bronze reflections above, abdomen paler beneath. Forewing: ground colour of dorsal surface very dark brown with bronze lustre before fascia, shining more strongly beyond; fringe greyish brown along margins, shining metallic grey at apex, single postmedial fascia, shining white, convex and oblique followed by small, shining white apical spot. Hindwing: ground colour and fringe brownish grey, shining metallic grey. Legs dark greyish brown with scattered black patches, very pale o n lower surface. Wing measurements. d, 3.6-4.2 mm (4). 9, 3.6-4.8 mm (8). Lectotype, 3.8 mm. Genitalia. d (Fig. 43). Uncus bilobed with deep notch between lobes, each lobe papillate. Gnathos: as in Fig. 43(a); transverse ventral plate broad; posteriorly-directed lateral arms closely juxtaposed, narrow, not reaching uncus; dorsolateral arms with short anteriorlydirected processes. Tegumen arcuate, straplike. Vinculum: lateral arms narrow and
FIG. 47. Stigmella populetorum d, genitalia: (a) genital capsule; (b) aedeagus.
articulating with tegumen at dorsal extremities; ventral plate narrow with medial expansion o n posterior margin. Saccus as wide as ventral plate, deeply lobed, each lobe as long as broad at base. Valve with narrowly pointed style and broad cuiller with distal hook as in Fig. 43(c), not reaching uncus. Transtilla: lateral arms narrow and arcuate; ventral arms reaching saccus; transverse bars fused t o form a continuous arcuate strap. Juxta quadrate, broadening posteriorly. Aedeagus: flaskshaped, equal to length of capsule; vesica with cornuti as many large denticles orientated in a ridge posteriorly and with a globular plate of minute papillae. 9 (Fig. 44). Ductus bursae long. Accessory sac very large and adorned with many small denticles distributed laterally and in a small distal patch, and with a long simple duct arising proximally. Bursa copulatrix : smaller than accessory sac, without pectinations; signum absent. Anterior apophyses long and narrow, arcuate. Posterior apophyses straight and narrow, equal in length to apophyses anteriores. BIOLOGY. Egg. Laid o n lower surface, adjacent t o leaf vein in the single mine examined. Host. The bred specimens examined are from Ulmus fulva (red elm) and Ulmus
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americana (white elm); Braun (1917: 182) also reported specimens from the cork elm, Ulmus thomasi (= racemosa). Mine. Long, linear, o n upper surface, increasing gradually in breadth; pale brown with conspicuous dark line of frass deposited centrally (Fig. 9 1). Larva. Emerges o n upper surface of leaf. Cocoon. Ovoid, dark brown. Voltinism. There are three generations per year. Braun (1917: 182) noted that late instar larvae may be found in mid-June, late July, August and September. Imagines have been collected in late April to mid-May and early July through to August. DIAGNOSIS. This species is differentiated from all other North American Stigmella by the very white fascia and the discrete, white apical spot of the forewing. The male genitalia differ from those of fuscotibiella in the form of the aedeagus, the ridge of small cornuti on the vesica and in the narrower, more widely spaced ventral arms of the transtilla in apicialbella. The bursa copulatrix lacks the U-shaped signum present in fuscotibiella, while the fewer and stouter spicules of t h e accessory sac separate apicialbella from villosella. The mine produced by the larvae of apicialbella differs from others on Ulmus spp. (produced by larvae of
Ectoedemia ulrnella) in the absence of a terminal blotch. DISCUSSION. Chambers, in his description of this species, failed to list the material he examined although four specimens have been labelled by him and given the USNM type no. 523. He also failed to provide collecting dates for the type material but stated that the imago is present in June. Braun (1912: 88) subsequently described this species, from a single specimen, under the name leucostigma; later examination of a larger series (1914: 21) led her to synonymize the two species: genitalia preparations of the t y p e material confirm this synonymy. The form of the female genitalia, in particular the very large spiculate accessory sac, relate this species t o villosella. The male genitalia, however, while possessing some attributes of villosella seem more closely related to those of fuscotibiella in the form of the uncus, gnathos and valve. DISTRIBUTION. U.S.A.: Kentucky, Ohio. Type material examined Here designated, Lectotype d (of apicialbella), U.S.A., Kentucky, no further data [see discussion] (Chambers) (genitalia slide no. CNC 3476; USNM). Paralectotypes 1 d, 299, data as lectotype (Chambers) (genitalia slide nos. CNC 3474, USNM 16276, USNM 17232; USNM). Holotype 0 (of leucostigrna), U.S.A.,
FIG. 48. Srigmeffaaromella d, abdomen (dorsal) showing: ( a ) position of scale pouches; (b) enlargement of scale pouch.
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Key to males Forewing ground colour brown, with a shining silver fascia. Male with style equal in length t o apical process of cuiller . . . . . . . fuscotibiella - Forewing pale, with terminal patch o f brown scales. Male with style projecting well beyond apex of cuiller . . . . . . . . . . . . . . . . . . 2 2 Abdomen of male ochreous; scale patches present on ventral surface . . . . . . . . . . . . aromella - Abdomen of male greyish-brown; without ventral scale patches . . . . . . . . . . . . .populetorum 1
Stigmella fuscotibiella (Clemens) (Figs. 45 and 46) Nepticula fuscotibiella Clemens, 1862b: 133 [description of adult] . Stigmella fuscotibiella (Clemens); Wilkinson & Scoble, 1979: 23 [redescription]. Nepticula ciliaefuscella Chambers, 1873 : 128 [synonymized by Chambers, 1875a: 1171. Nepticula discolorella Braun, 1912: 86 [synonymized by Braun, 19 17: 1851 . DESCRIPTION. See Wilkinson & Scoble (1979). DIAGNOSIS. Differs from the related populetorum and aromella and from apicialbella in external features and in the form of the valve and gnathos of the male genitalia (Fig. 45). The female genitalia of fuscotibiella (Fig. 46) differ from those of apicialbella in the presence of a U-shaped signum in the bursa copulatrix. The mine of fuscotibiella differs from that o n Salix spp. produced by Stigmella
FIG. 49. Stigmella pallida 9 , genitalia.
Ohio, Cincinnati, bred from Ulmus fulva, 1 .viii. 1912 (Braun) (ANS). Further material examined U.S.A., Ohio, Cincinnati, 1 d, 5 99, 1 mine, o n Ulmus fulva, 29.iv. 1905-8.vii. 1916 (Braun) (ANS), 2 d d , on Ulmus fulva, 8.viii. 1913,6.vii. 1914 (Braun) (USNM).
Species-group FUSCOTIBIELLA The following three species are characterized by features of the male genitalia, in which they are similar to members of the rosaefoliella group; the females possess a U-shaped signum o n the bursa which is lacking in the rosaefoliella group. Larvae of the fuscotibiella group mine members of the Salicaceae. 28
FIG. 5 0 . Stigmellu saginella d, genitalia: (a) genital capsule; (b) aedeagus.
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Philip J. Newton and Christopher Wilkinson Hollow, 2 mines, o n Salix nigra, 3 .vii. 1909 (Braun), Madisonville Swamp, 2 mines, on Salix discolor, 9.vii. 1909 (Braun), Spring Grove, 1 mine, o n SaZix sp., 17.viii.1913 (Braun) (ANS). CANADA: Ontario, Normandale, 3 mines, on Salix sp., 16.viii.1957 (Freeman and Lewis); Nova Scotia, Ohio, 9 mines, on Salix sp., 1 0 + 22.vii.1959 (Lewis) (CNC).
Stigmella populetorum (Frey & Boll) (Fig. 47) Nepticula poputetorum Frey & Boll, 1878:
FIG. 5 1. Srigmella saginella P, genitalia.
pallida which is narrower terminally and not blotch-like in appearance. DISTRIBUTION. U.S.A.: Ohio, Pennsylvania, Kentucky (Chambers), Colorado, Maine, Massachusetts. CANADA: Ontario, Nova Scotia. Material examined See Wilkinson & Scoble (1979) (including ‘specimen c’). U.S.A.: Colorado, Univ. o f , 1 ex., viii. 1912 (Cockerell); Massachusetts, Barnstable, I d , 2 9 9 , 1 ex., lO.vii.194927.vi. 1958 (KimbaZZ); Maine, Lincoln, 1 d, 1 9 , 1 ex., 6.vii-2O.viii.?, Augusta, 1 d, 23.v. 1942 (Brower), Bar Harbor, 1 9 , 16.iv. 1945 (Brower) (USNM); Ohio, Still House
276. Stigmella populetorum (Frey & Boll); Wilkinson & Scoble, 1979: 26 [redescription, lectotype designated] . DESCRIPTION. See Wilkinson & Scoble (1979). BIOLOGY. Voltinism. Late instar larvae have been found in mid-June and August; Braun (1 91 7: 195) also reported their collection in t h e latter part of July. Imagines have been taken in March (a single specimen probably advanced b y artificial conditions) and in late June through t o September. While Braun was of t h e opinion that there are several generations per year the data confirm only t w o and suggest the possibility of a third. DIAGNOSIS. The colour of the forewing distinguishes this species from fuscofibiella and apicialbella; the male genitalia may be differentiated by t h e size of the style of the valve which is much larger in populetorum (Fig. 47); females are indistinguishable from those of fuscotibiella in genital morphology. Differs from aromella males in the lack of abdominal scent scale patches and in the colour of the abdomen. DISTRIBUTION. U.S.A.: Texas, Ohio, Kentucky (Braun), California (Braun). CANADA: Ontario, British Columbia. Material examined See Wilkinson & Scoble (1979). U.S.A., Ohio, Big Dutch Creek, 2 mines, o n Populus X canadensis, 20.vi. 191 1 (Braun) (ANS). CANADA: Ontario, Delhi, 5 mines,on Populus deltoides, 12.viii. 1966 (Lewis); Ontario, Simcoe, 3 mines, on Populus nigra italica, 22.viii. 1957 (Freeman and Lewis); British Columbia, Coldstream, 4 mines, on Populus trichocarpa, 14.vi. 1957 (Freeman and Lewis) (CNC).
A taxonomic revision of Stigmella Specimen d
Wilkinson & Scoble (1979) described a single female specimen having genitalia indistinguishable from those of crataegifoliella but with dissimilar external features. Re-examination of the genitalia has shown, however, that the bursa copulatrix has a U-shaped signum comprising heavily sclerotized pectinations and that they are similar to fuscotibiella and populetorum. The external features are as those of populetorum but differ in the absence of brown scales on the pale portion of the forewing and in the head tufts, which are almost white in this specimen. Material examined 9, CANADA, British Columbia, Summerland, 21.vi. 1935 (Gartrell) (al. ex. 4.8mm; genitalia slide no. 3304; CNC).
Stigmella aromella Wilkinson & Scoble (Fig. 48) Stigmella aromella Wilkinson & Scoble, 1979: 27. DESCRIPTION. See Wilkinson & Scoble (1979).
417
BIOLOGY. Voltinism. Late instar larvae have been taken in mid-August and mid-September with imagines being collected in February, March and late August. The data suggest a bivoltine state: emergence of adults from overwintering larvae was probably ‘forced’ by laboratory conditions and the first generation, therefore, may be somewhat later in the year than is apparent from these records. DIAGNOSIS. Differs from populetorum in the colour of the abdomen which is ochreous in aromella and in the scent-scale patches on the ventral surface of the abdomen in the males of aromella. (See Fig. 48.) DISCUSSION. Wilkinson & Scoble (1979) tentatively identified female specimens from St Williams, Ontario, as being aromella and not populetorum on the basis of locality. The single genitalia preparation of a female from St Williams shows that the bursa copulatrix is much larger and that the chains of pectinations are more weakly sclerotized than in specimens from other localities. Both these attributes may prove t o be real species differences in the light of further study.
FIG. 5 2 . Stigrnella nigriverticella d, genitalia: (a) genital capsule; (b) aedeagus; (c) enlarged detail of scales on style.
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Philip J. N e w t o n and Christopher Wilkinson
FIG. 53. Stigrnella nigriuerricellu 9 , genitalia: (a) genitalia;(b) enlarged detail of spines on bursa copulatrix.
DISTRIBUTION. CANADA: Ontario. Material examined See Wilkinson & Scoble (1979). CANADA, Ontario, St Williams, several vials of mines, larvae and pupae, on Populus deltoides, 21 .ix. 1961, o n Populus x canadensis, 25.viii. 196 1 (FIS).
Stigmella pallida (Braun) c0mb.n. (Figs. 49, 92 and 112) Nepticula pallida Braun, 1912: 8 5 ; Braun, 1917: 196 [redescription] ; Braun, in Forbes, 1923: 9 5 [redescription].
DESCRIPTION. External features. 0 (Fig. 112). Palps pale buff; antenna grey-buff; tufts on front of head and vertex ochreous; eye-caps creamy white, lustrous; collar offwhite. Thorax and abdomen buff, scattered purple reflections on thorax, abdomen shining grey beneath. Forewing: ground colour of dorsal surface buff, each scale tipped with yellow-brown producing an irrorate effect, shining silver-grey when light at suitable angle. Hindwing: ground colour and fringe pale grey, shining metallic grey. Legs buff above, yellowish below.
A taxonomic revision of Stigrnella
4 19
Wing measurements. Holotype, 3.8 mm. Genitalia 0 . (Fig. 49). Ductus bursae short and broad with accessory duct arising medially. Bursa copulatrix large, covered with weakly sclerotized chains of pectinations. Anterior apophyses very broad basally, short and arcuate. Posterior apophyses long and straight, reaching beyond apophyses anteriores. BIOLOGY. Egg. Laid o n lower surface adjacent t o midrib of leaf. Host. Salix sp. Mine. Long, linear, o n lower surface, gradually broadening throughout its length; frass deposited centrally and somewhat loosely as a black line (Fig. 92). Larva. Emerges on lower surface of leaf. Cocoon. Oval, reddish brown. DIAGNOSIS. Differs from any other member of the genus listed here in the generally buff appearance of the dorsal and ventral aspects and in darker tips to the scales of the forewings. The forewing also lacks the distal dark patch that is present in the other species with pale coloration, fuscotibiella, populetorum and nigriverticella. The mine of pallida differs from others on Salix s p . produced by fuscotibiella which are broad terminally, sometimes blotch-like. DISCUSSION. Owing to the fact that the male of this species is unknown i t is difficult t o ascertain its true affinities within the genus. It is placed here largely on the basis that the host plant genus is also that of fuscotibiella and that the external features are similar t o fuscotibiella and populetorum. The female genitalia are damaged and it is impossible to distinguish a signum o n the bursa copulatrix. DISTRIBUTION. U.S.A.: Ohio. Type material examined Holotype 9, U.S.A., Ohio, Cedar Point, on Salix sp., 11 .viii. 1909 (Braun) (genitalia slide no. 11 PJN; ANS). Further material examined U.S.A., Ohio, Cedar Point, 1 mine, on Salix sp., 1 .viii. 1909 (Braun) (ANS). Species-group SAGINELLA
Described by Wilkinson & Scoble (1979) to include saginella, latifasciella, and castaneaefoliella t o which are here added flavipedella, sclerostylota and nigriverticella. The male genitalia possess a heavily sclerotized gnathos
/
.,
,
,
,
. .:-.
.
/
FIG. 54. Stigmella castaneaefoliella 9 , genitalia.
and uncus, the valve bears plumulate or digitate scales o n the style or along the inner margin. There is a large, quadrate juxta which is usually marked with opposite V-shaped folds. The group is also characterized by features of the female genitalia: the basal lobes of the apophyses posteriores; the denticulate accessory sac and the signum comprising spinose ridges. The larvae mine leaves of certain Fagaceae. Key to species 1 Female with signum comprising two rows of pectinations
. . . . .. . . . . . . ... ... ..
2
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Philip J. Newton and Christopher Wilkinson
FIG. 5 5 . (a-c) Sfigmella flavipedella d , genitalia: (a) genital capsule; (h) aedeagus; (c) enlarged detail of scales on style. (d, e) Stigmella castaneaefoliella d , genitalia: (d) aedeagus; (e) enlarged detail of scales on style. -
2 -
3 -
4
-
5
-
Female with signum comprising a U-shaped hand of pectinations 3 Forewing with cream scales forming a narrow medial fascia latifasciella Forewing with whitish scales forming a broad medial fascia . . . . . . . . . . . . nigriverticella Forewing irrorate with creamy-white scales saginella Forewing uniformly brown in colour . . . . . 4 Aedeagus of male with relatively large cornuti; valve with plumulate setae. Female with signum heavily sclerotized . . . . . . . .castaneaefoliella Aedeagus with small cornuti; valve with terminally digitate setae. Female with signum relatively weakly sclerotized . . . . . . . . . . 5 Gnathos of male heavily sclerotized; tarsi white sclerosfylota Cnathos of male weakly sclerotized; tarsi brown with pale yellow patches . . . . . . .flavipedella
.................. ..............
Sfigmella saginellu (Clemens) (Figs. 50 and 5 1) Nepticula saginella Clemens, 1862a: 85 [description of mine and larva] ; Clemens, 1865: 146 [description of adult] ; Busck, 1903 : 220 [ Clemens’ ‘type’] .
Stigmella saginella (Clemens); Wilkinson & Scoble, 1979: 39 [redescription, neotype designated] . Nepticula quercicastanella Chambers, 1873 : 127 [synonymized by Braun, 1917: 1951. Nepticula fuscocapitella Chambers, 1873: 128 [synonymized by Braun, 1917: 1951.
DESCRIPTION.
See Wilkinson & Scoble
( 1979).
DIAGNOSIS. Generally paler than other members of the group in the colour of the thorax, abdomen and forewing ground colour: the variously irrorate forewings separate this species from the related latifasciella and nigriverticella in which the cream scales are confined to fasciae and patches on the forewing. The form of the juxta and the style separate saginella males (Fig. 50) from those of sclerostylota which is very similar in other aspects of the genitalia. The larger denticles of the accessory sac and the U-shaped signum
A taxonomic revision of Stigmella separate the female genitalia of saginella (Fig. 5 1) from those latifasciella and nigriverticella which are otherwise indistinguishable from those of flavipedella except in the nature of the pectinations of the bursa. DISCUSSION. We have examined additional material from Massachusetts and Missouri, localities not previously cited.
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DISTRIBUTION. U.S.A.: Ohio, New York, Virginia, Massachusetts, Illinois, Missouri, Pennsylvania (Clemens), Kentucky (Braun), California (Braun). CANADA: Ontario, Quebec. Material examined See Wilkinson & Scoble (1 979) (including ‘specimen e’). U.S.A.: Ohio, Cincinnati,
FIG. 5 6 . Stigmellaflavipedella Q , genitalia: (a) genitalia; (b) enlarged detail of spines on bursa copulatrix; (c) enlarged detail of pectinations on signum.
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Philip J. Newton an d Christopher Wilkinson
3dd, 4 9 9 , 2 ex., 21.vii.1909-5.ix.1919 (Braun), Adams County, 1 mine, on Quercus prinus, no date (Braun), Spring Grove, 1 mine, on Quercus platanoides, 3. ix .19 13 (Braun), Salem Pike, 1 mine, on Quercus platanoides, 26.vii. 1913 (Braun), Mount Healthy, 1 mine, o n Quercus platanoides, 17.vii. 1913 (Braun), Still House Hollow, 2 mines, on Quercus platanoides, 5. x . 19 16 (Braun); Massachusetts, Hyde Park, 1 d, 8.vii. 1910 (Haimbach)(ANS), Barnstable, 1 b, 1 9 , 24.viii. 1952, 26.vi.1949 (Kimball) (MCZ); Illinois, Putnam County, 1 d, 8 99, 31 .viii. 1962-4.vi. 1967 (Glenn) (USNM); New York State, Leroy, 1 d, 14.viii. 1947 (Kimball); Missouri, Independence,
Jackson County, 1 9, 14.vii. 1971 (Heitzmun) (MCZ). CANADA: Ontario, Kinburn, 2 mines, on Quercus alba, 26.vii. 1957 (Lewis);Quebec, Val Tetreau, 2 mines, on Quercus alba, 2 1. ix. 1960 (CNC).
Stigmella latifasciella (Chambers) Nepticula latifasciella Chambers, in Hayden, 1878a: 106. Stigmella latifasciella (Chambers); Wilkinson & Scoble, 1979: 47 [redescription]. Nepticula macrocarpae Freeman, 1967: 19 [synonymized by Wilkinson & Scoble, 1979: 471.
A taxonomic revision of Stigmella DESCRIPTION. (1 979).
423
See Wilkinson & Scoble
BIOLOGY. Voltinism. Probably trivoltine in the U.S.A. DIAGNOSIS. The externals differ from those of saginella and nigriverticella in the distribution of cream scales on the forewing; these scales are confined to a medial position and an apical spot in latifasciella. The genitalia of both males and females are indistinguishable from those of nigriverticella but aspects of the uncus and gnathos, in the male, and the signa, in the female, separate them from castaneaefoliella, flavipedella and sclerostylo ta. DISCUSSION. We have examined additional material from Pennsylvania, a locality not previously cited. There are also 6 dd from Michigan, Ohio and Massachusetts which have a less striking fascia than other males of this species; they are tentatively identified as latifasciella. DISTRIBUTION. U.S.A.: Kentucky, Ohio, Michigan, Virginia, Arkansas, Pennsylvania, New York, Massachusetts, Illinois, Florida. CANADA: Ontario, British Columbia. Material examined See Wilkinson & Scoble (1979). U.S.A.: Pennsylvania, Pittsburgh, 2 90, 28.viii. 1909 (Kahl), Allegheny County, Oak Station, 1 d, 15.v.1913 (Marloffl (MCZ), 1 d, 17.v.1910; Illinois, Putnam County, 18 dd, 20 99, 4 ex., 19.viii. 1962-1O.k. 1974 (Glenn); Florida, Pensacola, 2 d d , 10+ 12.i. 1961 (Hills) (USNM); New York, Munroe County, 1 9 , 30.vi. 1947,19,30.vi. 1948 (Kimball)(MCZ); Ohio, Salem Pike, 1 mine, on Quercus rubra, 26.vii.1913 (Braun), Spring Grove, 1 mine, o n Quercus rubra, 3.ix. 1913 (Braun) (ANS). CANADA: Ontario, Bells Corner, 3 mines, o n Quercus rubra, 23.ix. 1958 (CNC), Orillia, several vials of mines, larvae and pupae, 19.vii. 1965 (FIS). Tentative determinations: U.S.A.: Michigan,Gratiot County, 1 d, 19.viii. 1955 (Nodges) (USNM); Ohio, Cincinnati, 2dd, 29.ix. 1951 (Braun) (ANS); Massachusetts, Barnstable, 3 dd, 26.viii.1962 (Kirnball) (MCZ). Stigmella nigriverticella (Chambers) c0mb.n. (Figs. 52, 5 3 and 114) Nepticula nigriverticella Chambers, 1875a: 118;
FIG. 58. Stigmellu sclerosfyloru 9 , genitalia: (a) genitalia; (b) enlarged detail of pectinations on bursa copulatrix.
Chambers, in Hayden, 1878b: 158 [listed] ; Braun, 1917: 194 [redescription] ; Braun, in Forbes, 1923: 95 [redescription]. Nepticula maculosella Chambers, 1880: 193 [synonymized by Braun, 19 17: 1941 DESCRIPTION. External features. d. Palps creamy white; antenna pale brown; tuft on front of head dark ochreous, vertex brown; eye-caps shining white ; collar off-white. Thorax brown with scattered creamy white scales. Abdomen grey-brown with metallic grey reflections above, shining metallic grey beneath. Forewing: ground colour of dorsal surface brown, irrorate with creamy white scales; fringe greyish white, shining silver at apex; basal patch of dark brown scales on dorsal margin, reaching half way across wing,
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with purple reflections, distal third brown with bronze and purple reflections, small apical spot of creamy white scales. Hindwing: ground colour and fringe grey, shining silver. Legs brown with scattered paler areas, shining metallic grey. ? (Fig. 114). As d except for a pair of ventromedial fenestrae o n fourth abdominal segment. Wing measurements. d, 4.4-5.2 mm (5). 9, 4.6-5.2 mm (3). Holotype, 4.9 mm. Genitalia. d (Fig. 5 2 ) . Uncus markedly bilobed and heavily sclerotized. Gnathos: M-shaped as in Fig. 52(a); transverse ventral plate broad ; posteriorly-directed lateral arms closely juxtaposed at base. Tegumen a broad semicircular plate. Vinculum: lateral arms broad and fused with tegumen at dorsal extremities, inner surface emarginated dorsally; ventral plate broad. Saccus very narrow, weakly bilobed. Valve reaching beyond uncus, narrow with broad, pointed style and slightly constricted cuiller; style bears plumulate scales as in Fig. 52(c), particularly along inner margin. Transtilla: lateral arms short and very broad; ventral arms short and stout; transverse bars long and tapering, meeting but not fusing. Juxta quadrate, bearing two Vshaped folds with vertices adjacent, articulates with base of valves laterally. Aedeagus: flaskshaped, three-quarters length of genital capsule; vesica with cornuti as minute denticles and with lunulate plate of minute papillae.
? (Fig. 53). Ductus bursae long and narrow. Accessory sac covered with small denticles, evenly distributed and with duct arising distally. Bursa copulatrix: covered with broad heavily sclerotized pectinations on striations of the bursa as in Fig. 53(b); signum double, comprising two straight bands of spinose ridges. Anterior apophyses very broad basally, tapering and arcuate distally. Posterior apophyses with a pair of closely associated lobes basally, straight and narrow, reaching beyond apophyses anteriores. BIOLOGY. Immature stages unknown. Host. The specimen described by Chambers (1 875a: 11 8 ) was taken o n the trunk ('hunk', sic) of wild cherry but there is n o other evidence t o verify that this is the true host of nigriverticella. Volfinisrn. Probably trivoltine though the absence of any records of the larvae prevents confirmation of this. DIAGNOSIS. Very close to latifasciella in externals and differentiated only b y the paler colouring of the light scales of the forewing and the relatively broader pale band in nigriverticella: the genitalia of these two species are indistinguishable. Males are separated from those of saginella by t h e rather broader dorsolateral arms of the gnathos, the form of t h e uncus and the setae of the valves; the double signum in the female differentiates the genitalia from other members of the group except latifasciella.
fb) FIG. 5 9 . Stigmella corylifoliella d, genitalia: (a) genital capsule; (b) aedeagus.
A taxonomic revision o f Stigmella
425
from the cited localities especially, suggests otherwise. DISTRIBUTION. U.S.A. : Texas, Cincinnati, Ohio, New York, Arkansas, Pennsylvania, Kentucky (Chambers). Type material examined Holotype 9 (of Nepticula maculosella Chambers), U.S.A., Texas (Chambers) ‘Kentucky’ deleted (genitalia slide no. CNC 3 4 9 1 ; MCZ). Further material examined U.S.A.: Ohio, Cincinnati, 4 dd, 3 99, 15 .v. 1906-1O.v. 1908 (Braun) (ANS); Arkansas, Washington County, 2 dd, 22 + 30.v. 1966 (Hodges) (USNM), 1 d, 2 1 .viii. 1971 (Heitzman); New York, Munroe County, 3 dd, 3 99, 25.vi. 1948-26.vii. 1949 (Kimball); Pennsylvania, Oak Station, Allegheny County, 1 0 , 29.v. 1910 (Marloff)(MCZ).
Stigmella castaneaefoliella (Chambers) (Figs. 5 4 and 5 5 ) Nepticula castaneaefoliella Chambers, 1875a: 117. Stigmella castaneaefoliella (Chambers); Wilkinson & Scoble, 1979: 4 4 [redescription].
FIG. 6 0 . Stigmella corytifoliella Q , genitalia.
DISCUSSION. Chambers (1 880), in describing maculosella, noted its similarity t o nigriverticella and suggested that it might prove t o be the same species, or a ‘variety’. Braun (1 91 7) synonymized t h e two species on the grounds that specimens captured in Cincinnati show a range of variation sufficient t o encompass both. The labelling of the type examined is confused, with both species’ names appearing. The name of the junior synonym, however, is in Chambers’ handwriting and is taken to be correct. The head of t h e holotype is missing. It is unfortunate that nothing is known of t h e biology of this species: we regard it as separate from latifasciella until evidence gained from further collection and rearing of material,
DESCRIPTION. See Wilkinson & Scoble (1979). DIAGNOSIS. The external features differ markedly from the related species, saginella, latifasciella and nigriverticella in the absence of pale markings o n the forewings, but are not easily distinguished from those of flavipedella and sclerostylota. The male genitalia of castaneaefoliella differ from those of flavipedella in that there are plumulate scales on t h e valve (Fig. 55e) and more numerous, larger cornuti (Fig. 55d). In females, the more heavily sclerotized signum and the chainlike pectinations of the bursa of castaneaefoliella (Fig. 54) also separate this species from flavipedella. DISTRIBUTION. U.S.A.: Kentucky, Ohio, Virginia, Illinois, Pennsylvania, New Jersey, Massachusetts, Florida. CANADA: Ontario. Material examined See Wilkinson & Scoble (1979). U.S.A.: Florida, Pensacola, 1 0, 24.iii. 1962 (Hills); Illinois, Putnam County, l O d d , 6 9 9 , 2.vii. 1960-9 .ix. 1974 (Glenn); Massachusetts,
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Philip J. Newton and Christopher Wilkinson
Barnstable, 3 dd, 15-27.viii. 1952 (Kimball) (USNM); Ohio, Adams County, 1 mine, on Castanea sp., 30.vi. 1918(Braun); 1 mine,date uncertain: 5.vii.1911-9.vii.1916 (Braun) (ANS). CANADA. Ontario: Normandale, 2 mines, on Custanea dentata, 1 .vii. 1956; S t Williams, 2 mines, o n Custanea dentata, 17 .vii. 1956 (CNC), several vials of mines,larvae and pupae, on Castaneadentata, 23 .viii.1961 (FIS).
Stigmella jlavipedella (Braun) Figs. 55, 56, 93 and 115) Nepticula jlavipedella Braun, 1914: 1 9 ; Braun, 1917: 191 [redescription] ; Braun, in Forbes, 1923: 94 [redescription]. Stigmella jlavipedella (Braun); Wilkinson & Newton, 1981: 31. DESCRIPTION. External features. 6. Palps whitish; antenna dark brown; tufts on front of head and vertex dark brown; eyecaps shining white; collar off-white. Thorax and abdomen dark brown with purple reflections, abdomen shining metallic grey beneath. Forewing: ground colour of dorsal surface dark brown with purplish grey reflections; fringe pale grey-brown, shining silver. Hind-
wing: ground colour and fringe greyish, shining metallic grey. Legs dark brown above, with yellow patches o n tarsi, shining metallic grey o n lower surface. 0 (Fig. 115). As d except for a pair of ventromedial fenestrae o n fourth abdominal segment. In one ? (paratype) the tufts o n the front of t h e head and t h e vertex are ochreous. Wing measurements. b, 4.0-5.0 m m (8). 9, 3.6-4.8 mm (12). Holotype, 4.0 mm. Genitalia. 6 (Fig. 5 5 ) . Uncus large and papillate. Gnathos: M-shaped as in Fig. 55(a); transverse ventral plate narrow; posteriorlydirected lateral arms closely juxtaposed at base, broad and heavily sclerotized, reaching just beyond uncus; dorsolateral arms very broad with short posteriorly- and long anteriorly-directed processes. Tegumen a broad arcuate plate. Vinculum: lateral arms broad and fused with tegumen at dorsal extremities, inner surface emarginate dorsally; ventral plate broad. Saccus half width of ventral plate, weakly bilobed. Valve: divided into large style, just reaching uncus, and broad cuiller which narrows distally; style bears terminally digitate scales as in Fig.
LXG. 61. Szigmella ostryaefoliella d, genitalia: (a) genital capsule; (b) aedeagus.
A taxonomic revision of Stigmella
427
FIG. 6 2 . Stigmella juglandifoliella d, genitalia: (a) genital capsule; (b) aedeagus.
55(c). Transtilla: lateral arms short and broad; ventral arms triangular; transverse bars tapering, meeting but not fusing. Juxta quadrate, bearing two V-shaped folds with vertices adjacent, articulating with base of valves laterally. Aedeagus: narrow and regular in width, half length of genital capsule; vesica with cornuti as small spicules and with U-shaped collar of minute papillae. 0 (Fig. 56). Ductus bursae long and narrow. Accessory sac broad at origin, tapering markedly and adorned with patches of large denticles which become smaller and more evenly distributed distally (see Fig. 56). Accessory duct arising distally. Bursa copulatrix: large and covered with short, heavily sclerotized chains of pectinations o n striations of the bursa as in Fig. 56(b); signum single, comprising a U-shaped band of spinose ridges as in Fig. 56(c). Anterior apophyses very broad basally. Posterior apophyses with papillate lobes basally, straight and narrow and equal in length t o apophyses anteriores.
BIOLOGY. Egg. Laid o n upper surface of leaf. Host plants. Quercus platanoides (swamp white oak); Quercus patustris (pin oak); Quercus rubra (red oak); Quercus imbricaria (shingle oak). Mine. At first an upper-surface, very narrow linear mine which abruptly enlarges and becomes transparent; then broadens gradually, much contorted in later stages; frass deposited as diffuse central line in early stages, but across entire breadth of mine in later portion (Fig. 93). Larva. Usually green but occasionally purplish (Braun, 1914: 20); vacates mine through lower surface of leaf. Cocoon. Almost white. Voltinism. There are three generations per year: adults have been taken in March, May and early June, and mid-July and August; larvae may be collected in June and late July (Braun, 1917) and in late August and September.
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FIG. 6 3 . Stigmella Iongisucca d, genitalia: (a) genital capsule; (b) aedeagus.
DIAGNOSIS. The externals are virtually indistinguishable from those of castaneaefoliella and sclerostylota : the only consistent difference being the yellow patches o n the tarsi of flavipedella as opposed t o the whitish tarsi of castaneaefoliella and sclerostylota. The male genitalia of flavipedella differ from those of castaneaefoliella in that the scales of t h e valves are terminally digitate and the denticulate cornuti are much smaller and fewer in number in flavipedella, compare Fig. SS(b) and (c) with (d) and (e); the form of the uncus and gnathos differs in sclerostylota. The female genitalia of flavipedella differ from those of castaneaefoliella in the form of the signum, which is less heavily sclerotized, and in the pectinations of t h e bursa, which are more scallop-shaped in flavipedella; t h e striations of the bursa and form of pectinations separate flavipedella from sclerostylota. The mine produced by larvae of flavipedella is generally much broader
and the frass line more diffuse than in mines of castaneaefoliella and saginella.
DISCUSSION. We now believe that the material which Wilkinson &, Scoble (1979: 46) referred t o in the discussion of castaneaefoliella and labelled 'Specimens 1 to 8' is conspecific with flavipedella. The separate treatment Teferred to in that paper was inadvertently omitted. The 0 paratype with the ochreous tufts is that described by Braun (1914: 19), and corresponds to the specimen bred from a mine on Quercus palustris as opposed t o the holotype and paratype which were bred from Q.platanoides. This difference in host plant does n o t appear to be significant since there are several examples of flavipedella from Q.palustris which have brown tufts. There are t w o additional specimens (Specimens 4 and 5, see below) which have indistinguishable external features from t h e aberrant paratype:
A taxonomic revision of Stigmella there is insufficient evidence to know if they represent the same taxon. DISTRIBUTION. U.S.A.: Ohio, Kentucky (Braun). Type material examined Holotype d, U.S.A., Ohio, Cincinnati, on Quercus plutanoides, 1 1.vii. 19 13 (Braun) (genitalia slide no. 43 PJN; ANS). Paratype (of uncertain identity) 0, locality as holotype, o n Quercus palustris, 9.viii. 1913 (Braun) (genitalia slide no. USNM 1621 1; ANS). Further material examined U.S.A. Ohio, Cincinnati: 2 d d , 499, 4 mines, on Quercus platanoides, 30. vii. 19 11, 1 .v. 1914-13.v. 1922 (Braun), 2 dd, 3 ??, 1 mine, on Quercus rubra, 5.x.1916,31.v12.vi.1917 (Braun), 366, 4 9 9 , 1 mine, on
429
Quercus palustris, 18 . x .19 16, 10. iii-9 .vi. 1917 (Braun), 2 d d , 1 O,onQuercusimbricaria, 6 .v. 1918 (Braun) (ANS). Specimens 4 and 5 : with external features as flavipedella except that the tufts on the front of the head and vertex are pale ochreous. In this respect they are indistinguishable from the aberrant paratype described above. The genitalia are as follows. Specimen 4 ( 0 ) : virtually indistinguishable from those of flavipedella, the only possible difference being that the spicules of the accessory sac are smaller in this specimen. Specimen 5 (13): differs from those of flavipedella in that the scales of the valves are plumulate and the cornuti larger, as in castaneaefoliella. DISCUSSION. The relationship of these two specimens to each other and t o the paratype of flavipedella described above cannot be established until further material has been collected from the respective localities and from the intermediate ranges. Material examined Specimen 4 , 9 , CANADA, Ontario, Simcoe, on Quercus sp., 12.ix. 1966 (Freeman) (al. ex. 4.2mm; genitalia slide no. 3409; CNC). Specimen 5, d, U.S.A., Florida, Lake Placid, Archbold Biological Station, 16-22.v. 1964 (Hodges) (al. ex. 4.4mm; genitalia slide no. 17423 ;USNM).
Stigmella sclerostylota sp.n. (Figs. 5 7 , 58 and
FIG. 64. Stigmella longisacca 9, genitalia: (a) genitalia; (b) enlarged detail of signum.
94) DESCRIPTION. External features. d. Palps whitish; antenna dark brown; tufts on front of head and vertex dark brown; eye-caps shining white. Thorax and abdomen dark brown with purple reflections, abdomen shining metallic grey beneath. Forewing: ground colour of dorsal surface dark brown with purplish grey reflections; fringe pale grey-brown, shining silver. Hindwing: ground colour and fringe brown. Legs brownish above, with tarsi white. 0. As d except for a pair of ventromedial fenestrae on fourth abdominal segment. Wing measurements. d, 4.0-4.4 mm (2). 9, 4.0 mm (1). Holotype, 4.2 mm. Genitalia. d (Fig. 57). Uncus convex with heavily sclerotized lateral lobes. Gnathos: M-shaped; transverse ventral plate narrow;
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Philip J. Newton and Christopher Wilkinson
I
I
I /
I /
II
I \
\
I
I
I
FIG. 6 5 . Stigmella unifasciella d, genitalia: (a) genital capsule; (b) aedeagus; ( c ) enlarged detail of scales on cuiller.
posteriorly-directed lateral arms heavily sclerotized, reaching just beyond uncus; dorsolateral arms with long anteriorly-directed processes. Tegumen an arcuate, oval plate. Vinculum: lateral arms broad and fused with tegumen at dorsal extremities; ventral plate broad. Saccus very narrow and weakly bilobed. Valve: not reaching uncus, deeply bifurcate to form a broad, bifid style which is heavily sclerotized distally, cuiller constricted ; style bears small, terminally digitate scales as in Fig. 57(d); cuiller bears plumulate scales as in Fig. 57(e), particularly along inner margin. Transtilla: lateral arms short and broad; ventral arms short, tapering distally ; transverse bars long and straight, meeting but n o t fusing. Juxta quadrate, bearing two V-shaped folds with vertices adjacent; articulating
laterally with bases of valves. Aedeagus: narrow and regular in width, threequarters length of genital capsule; vesica with cornuti as comma-shaped plate of minute papillae. 0 (Fig. 58). Ductus bursae short and broad. Accessory sac small, evenly adorned with large denticles and with accessory duct arising distally. Bursa copulatrix: very large and covered with long chains of pectinations as in Fig. 58(b); signum single, comprising a U-shaped band of spinose ridges. Anterior apophyses broad, tapering distally. Posterior apophyses with papillate lobes at base, straight and narrow, extending just beyond apophyses anteriores. BIOLOGY. Egg. Laid o n lower surface of leaf in three cases and on upper surface in five.
A taxonomic revision of Stigmella
43 1
before the number of generations may be accurately determined. DIAGNOSIS. The externals are indistinguishable from those of castaneaefoliella and flavipedella except that the hindwings of sclerostylota males are brownish. It also differs from flavipedella in the paler tarsi. The male genitalia differ from those of castaneaefoliella and flavipedella in the form of the uncus, which is convex, and the processes of the gnathos which are straighter and more heavily sclerotized, in sclerostylota; the heavily sclerotized style in sclerostylota separates the genitalia from those of saginella. The absence of striations on the bursa and the form of the pectinations separates the female genitalia from those of castaneaefoliella, saginella and flavipedella. The mines of sclerostylota increase in breadth more gradually than many produced by flavipedella and differ from those of saginella i n the distribution of the frass. DISCUSSION. This species is very closely related to the other members of the group: in externals it is nearer to castaneaefoliella and flavipedella but rather resembles saginella in the male genitalia and the mine. DISTRIBUTION. CANADA: Ontario.
15,.
d
'I
FIG. 6 6 . Stigmella unifasciella 9 , genitalia: (a) genitalia; (b) enlarged detail of spines on bursa copulatrix.
Host. Quercus alba. Mine. Upper-surface, long, linear, gradually increasing in breadth throughout its length; frass deposited centrally as a rather diffuse line in comparison with that of saginella (Fig. 94j. Larva. Emerges o n upper surface of leaf. Voltinism. Possibly bivoltine with emergence dates in Canada during late April and early May and in the U.S.A. during mid-June and early July: the Canadian material may, however, have been 'forced' by laboratory conditions and further collection is required 29
U.S.A.:
Arkansas.
Type material Holotype d, U.S.A., Arkansas, Washington County, Devil's Den Street Park, 17.vi. 1966 (Hodges) (genitalia slide no 16260; USNM). Paratypes. 1 d, data as holotype, l0,vii. 1966 (Hodges) Cgenitalia slide no. 17263; USNM). CANADA, Ontario: Normandale, 1 d, on Quercus alba, 31 .iv. 1962 (Freeman & L e w i s ) ; Simcoe, 1 ?, on Quercus alba, 5 . v . 1967 (Freeman) (genitalia slide nos 2953, 3410; CNC). Material excluded f r o m type-series CANADA. Ontario: Normandale, 3 mines, on Quercus alba, 8.ix.1961 (Freeman & L e w i s ) ; Sirncoe, 5 mines, on Quercus alba, 1 7 . 6 . 1 9 6 6 (Lewis) (CNC). Species-group COR YLIFOLIELLA
Includes corylifoliella, ostryaefoliella, juglandifoliella and myricafoliella, (see Wilkinson & Scoble, 1979); longisacca is added here. The group is well defined by both the male and
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Philip J. Newton and Christopher Wilkinson
female genitalia, particularly the palmate plates of the anellus in the male, and the pair of ovate signa in the female: the females show little variation between species and provide no reliable diagnostic characters. The forewing is marked by a single fascia. The separate identity of one species, myricafoliella, is not certain and therefore it is not included in the key. Key to males Saccus markedly broader than the ventral plate, weakly bilobed . . . . . . . . . . . . . . . . . . 2 Saccus aouroximatelv t o breadth of ventral .. . eaual . plate, bilobed . . . . . . . . . . . . . . . . . . . 3 Valve with dorsal style . . . . . . . . . longisacca Valve without dorsal style . . . . .juglandifoliella Tegumen with serrate posterior margin ; valve long and narrow; eye-caps with fuscous scales corylifoliella
Tepumen with smooth posterior margin; valve short and broad; eye-caps white, without fuscous scales . . . . . . . . . . . . . . . . .ostryaefolie~a
Nepticula corylifoliella Clemens, 1862a: 83 [description of mine] ; Braun, 1912: 92 [description of adult] . Stigmella corylifoliella (Clemens); Wilkinson & Scoble, 1979: 50 [redescription]. Nepticula virginiella Clemens, 1862a: 8 3 [synonymized by Braun, 1917: 1791. Nepticula minimella Chambers, 1873: 127 [synonymized by Braun, 1917: 1791. Nepticula opulifoliella Braun, 19 14: 22 [synonymized by Wilkinson & Scoble, 1979: S O ] . Nepticula paludicola Braun, 1917: 177 [synonymized by Wilkinson & Scoble, 1979: 501. Nepticula exasperata B ~ 1930: ~ ~17 [synonymized by Wilkinson & Scoble, 1979: 501. DESCRIPTION. see Wilkinson & Scoble ( 1979).
DIAGNOSIS. The males differ from those of the fascia is
FIG. 6 7 . Srigmella condaliafoliella d , genitalia: (a) genital capsule; (b) aedeagus.
~
,
A taxonomic revision o f Stigmella
433
broader in corylifolietla than in juglandifoliella and the ground colour of the forewing darker than in longisacca. The male genitalia (Fig. 59) differ from those of juglandifoliella and longisacca in the saccus, which is much shorter, and from ostryaefoliella in the tegumen, which is serrate, and the valve, which is narrower and longer in corylifoliella. The female genitalia of this species (Fig. 60) cannot be reliably separated from those of ostryaefoliella. DISCUSSION. We have examined additional material from California and Maine, localities not previously cited. It seems probable that this species is synonymous with the European betulicola Stainton but this point needs further investigation. DISTRIBUTION. U.S.A.: Ohio, New Jersey, Maine, Michigan, Kentucky, California, Pennsylvania (Braun), Maryland (Braun), North Carolina (Braun). CANADA: Ontario, New Bmnswick, Quebec, British Columbia (Wilkinson & Sco ble). Material examined See Wilkinson & Scoble (1979). U.S.A.: Michigan, Cheboygan County, 1 ex., 21.v. 1943 (Braun); Kentucky, London, 1 ex. ( n o further data); California, Almeda County, 1 ex., vii.1917 (Braun); Maine, Rockport, 2 ex., 20 + 24.vi. 1941 (Hebard); Ohio, Buckeye Lake, 1 mine, on Oxycoccus palustris, n o date (Braun), Adams County, 1 mine, on Vaccinium sp ., 9 . vii. 19 16 (Braun), Spring Grove, 1 mine, o n Corylus sp., 5.vii. 19 12 (Braun), 1 mine, on Opulaster sp., 13.vii. 1913 (Braun) (ANS). CANADA: British Columbia, Field, 2 mines, o n Betula sp., 29.vii. 1915 (Braun); Nova Scotia, Balsam, 1 mine, on Opulastersp., 19.viii. 1912 (Braun) (ANS); Ontario, Simcoe, 2 mines, on Corylus americana, 5.viii. 1959, 11 mines, o n Vaccinium sp., 18.viii. 1957 (Freeman & Lewis), Normandale, 1 2 mines, o n Gaylussacia sp., 11 .ix. 1961 (Freeman & Lewis), 2 mines, on Hamamelis virginiana, 8. ix . 196 1 (Lewis), Quebec Fairy Lake, 2 mines, on A h u s rugasa var. americana, 3 .viii. 1955 (Freeman) (CNC).
Stigmella ostryaefoliella (Clemens) (Fig. 6 1) Nepticula ostryaefoliella Clemens, 1862a: 83. Stigmella ostryaefoliella (Clemens); Wilkinson & Scoble, 1979: 5 4 [redescription].
FIG. 6 8 . Stigmella condaliafoliella 0, genitalia: (a) genitalia; (b) enlarged detail of markings on bursa copulatrix.
Nepticula caryaefoliella Clemens, 1862a: 84 [revised synonymy by Wilkinson & Scoble, 1979: 541. Nepticula obscurella Braun, 1912: 9 5 [synonymized by Wilkinson & Scoble, 1979: 541 DESCRIPTION. See Wilkinson & Scoble ( 1 979). BIOLOGY. Voltinism. Bivoltine in U.S.A. and Canada. DIAGNOSIS. The males differ from those of corylifoliella in the absence of fuscous scales on the eye-caps. The male genitalia differ from those of juglandifoliella and longisacca in the saccus, which is much shorter, and from corylifoliella in the tegumen which is n o t serrate, and the valve, which is shorter and broader in ostryaefoliella (Fig. 61). The
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Philip J. Newton and Christopher Wilkinson
FIG. 69.Stigmella tiliella d, genitalia: (a) genital capsule; (b) aedeagus; ( c ) enlarged detail from vesica.
female genitalia, however, cannot be reliably separated from those of corylifoliella. DISTKIBUTION. U.S.A.: Ohio, New Jersey, New York (Braun), Florida. CANADA: Ontario. Material examined See Wilkinson & Scoble (1979). U.S.A.: Florida, Archbold Bio. Station, Lake Placid, 4 99, 1 .v. 1975 (Heppner) (USNM); Ohio, West Fork Woods, 2 mines, on Carya sp., 19. vi. 1909 (Braun), 2 mines, on Ostrya sp., 8.vii.1909, 6.ix.1909 (Braun), North of Madeira, 2 mines, on Carpinus sp., 30.vKi. 1909 (Braun) (ANS). CANADA: Ontario, Normandale, 2 mines, on Carya ovate, l0.vii. 1956 (Freeman) (CNC).
Stigmella myricafoliella (Busc k) Nepticula myricafoliella Busck, 1900: 238. Stigmella myricafoliella (Busck); Grossbeck, 1917: 145; Wilkinson & Scoble, 1979: 57 [lectotype designated] . DISCUSSION. Possibly a synonym of ostryaeJbliella although its status here remains unchanged for the reasons given by Wilkinson & Scoble (1979). They, however, incorrectly cited myricafoliella as a new combination with Stigmella: they were preceded in this by Grossbeck (1917: 145) who stands alone in his use, at that time, of the name Stigmella. DISTRIBUTION. U.S.A.: Florida. CANADA: Nova Scotia. Material examined See Wilkinson & Scoble (1979). CANADA,
A taxonomic revision of Stigmella Nova Scotia, Tusket, 3 mines, on Myrica sp., 6.viii. 1960 (CNC).
Stigmella juglandifoliella (Clemens) (Fig. 62) Nepticula juglandifoliella Clemens, 1862a: 84 [description of mine and larva] ;Chambers, in Hayden, 1878a: 105 [description of adult, incorrectly cited as ‘n.sp.’] . Stigmella juglandifoliella (Clemens); Wilkinson & Scoble, 1979: 57 [redescription] . DESCRIPTION. See Wilkinson & Scoble (1979).
435
BIOLOGY. Voltinism. The limited material available suggests that there are possibly two generations in Ohio. DIAGNOSIS. Although the ranges overlap slightly, the wing span of juglandifoliella is generally much smaller than that of longisacca. The ground colour of the forewing is darker in juglandifoliella than in longisacca and the fascia more proximal; the fascia is narrower than in corylifoliella. The male genitalia (Fig. 62) differ from those of longisacca in the shorter saccus. The valve lacks the dorsal
FIG. 70. Stigmella tiliella 0, genitalia: (a) genitalia; (b) enlarged detail of bursa copulatrix; ( c ) enlarged detail of signum.
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Philip J. Newton and Christopher Wilkinson
FIG. 7 1 . Stigmella quercipulchella d , genitalia: (a) genital capsule; (b) aedeagus.
style and the transtilla is more club-like in longisacca.
DISTRIBUTION. U.S.A.: Ohio, Pennsylvania (Clemens), Kentucky (Chambers). Material examined See Wilkinson & Scoble (1979). U.S.A., Ohio, Cincinnati, 1 d, 2 mines, on .Juglam sp., 19.viii.1909, l.ix.1909, 12.vii.1912 (Braun) (ANS).
Stigmella longisacca sp.n. (Figs. 63, 6 4 and 117) DESCRIPTION. External jeatures. d? (Fig. 117). Palps greyish; antenna dark grey; tuft o n front of head ochreous, vertex darker; eye-caps shining white; collar greyish brown with metallic grey reflections. Thorax dark
brown with gold and purple reflections. Abdomen grey, shining metallic grey beneath. Forewing: ground colour of dorsal surface dark brown with gold and purple reflections becoming predominantly purple after the fascia; fringe grey-brown, shining metallic grey at apex; single postmedial fascia, shining silver, convex and broadening o n dorsal margin. Hindwing: ground colour and fringe greyish brown, shining metallic grey. Legs dark grey, shining silver. Wing measurements. d, 3.2-4.2 mm (10). 0 , 3.6-4.4 mm (4). Holotype, 3.8 mm. Genitalia. d (Fig. 63). Uncus bifid with lateral papillae. Gnathos: broadly U-shaped as in Fig. 63(a); transverse ventral plate narrow; posteriorlydirected lateral arms very
A taxonomic revision of Stigmella short and narrow; dorsolateral arms broad basally with short, stout posteriorlydirected processes. Tegumen a narrow transverse bar with serrate posterior margin. Vinculum: lateral arms broad and articulating with tegumen at dorsal extremities; ventral plate narrow. Saccus more than 5-times width of ventral plate. Valve: reaching beyond uncus with a short, dorsal style not extending beyond t h e cuiller which is concave along the inner margin. Transtilla: lateral arms very long, reaching the saccus; ventral arms rounded; transverse bars very short and tapering, meeting
437
but not fusing. Aedeagus: flask-shaped, approximately one-third length of genital capsule; vesica with a bearded rectangular plate of minute papillae and anellus comprising a pair of palmate plates. 0 (Fig. 64). Ductus bursae long and narrow. Accessory sac long and narrow with duct arising distally. Bursa copulatrix: large and covered with scallop-shaped chains of pectinations o n striations of t h e bursa; signum double, comprising t w o ovate plates of spinose ridges (Fig. 64b). Anterior apophyses tapering and arcuate. Posterior apophyses
FIG. 7 2 . Stigmella querciputchella Q, genitalia: (a) genitalia; (b) enlarged detail of bursa copulatrix.
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Philip J. Newton and Christopher Wilkinson
straight and narrow, reaching well beyond apophyses anteriores.
BIOLOGY. Host. Juglans sp. and Juglans californ ica. Cocoon. Obovoid, pale brown. Voltiriism. Two broods per year with the possibility of a third since there is a single specimen taken in early December. DIAGNOSIS. Generally larger than the very closely related juglandifoliella and differing in the ground colour of the forewing, which is paler, and the fascia, which is more distal in Iongisacca. The male genitalia are separated from those of juglandifoliella by the larger saccus, the dorsal spine of the valves and the less elongate transverse bars of the transtilla. The female genitalia are virtually indistinguishable from those of corylifoliella and ostryaefoliella, the only possible diagnostic characters being the relatively long apophyses posteriores in longisacca, the scallopshaped pectinations and the slightly unequal signa. DISCUSSION. The species is closely related t o juglandifoliella in both externals and male genitalia: there are no female examples of juglandifoliella available it is likely, however, that the genitalia are indistinguishable from those of longisacca and the other members of the group. The difference in the ~~
ii I
i \'
.-<...--,
FIG. 7 3 . S t i ~ m e l l uspecimen f 0, genitalia showing the spinosr nature of the accessory sac.
relative size of the saccus in longisacca and juglandifoliella may prove t o be geographical variation: other differences, however, support their treatment a5 separate species. DISTRIBUTION, U.S.A.: California.
Type material Holotype d, U.S.A., California, Ventura County, o n Juglrrns sp., 14.vii. 1930 (Barrett) (genitalia slide no. 16285; USNM). Paratypes 9 dd, 2 99, 2 ex,, U.S.A., California, Ventura County, 4-28.iv. 1931 (Barrett), Saticoy, 2 99, on Juglans calijornica, 2 . xii. 1930 (Barretf), Davis, 1 d, 13.viii. 1952 (Schingev) (genitalia slide nos. 16287, 16289, 17215, I723 5, 17236 ; USNM). Material excluded ,from type-series U.S. A,, California, Saticoy ,several cocoons, on Juglans sp., iv. 1931 (Barrett) (USNM).
Stigmella unifasciella (Chambers) (Figs. 6S, 66 and 116)
comb n.
Nepticula unifasciella Chambers, 1875a: 119; Chambers, in Hayden, 1878b: 158 [listed] ; Chambers, 1880: 193; Braun, 1917: 175 [redescription] ; Braun, in Forbes, 1923: 88 [ redescription] . DESCRIPTION. External features. d. Palps greyish; antenna very dark brown; tuft on front of head ochreous, vertex slightly darker; eye-caps shining white; collar dark grey. Thorax dark grey-brown with gold and purple reflections. Abdomen very dark brown t o black and shining grey-brown beneath. Forewing: ground colour of dorsal surface dark brown with gold reflections, basal two-thirds with strong purple reflections along costal margin, blue retlections o n dorsal margin, apical third more brown in ground colour and shining deep purple except for fascia; fringe dark brown, becoming grey distally; single postmedial fascia, shining silver. Hindwing: ground colour dark brown, shining metallic grey; fringe grey, shining metallic grey. Legs dark brown, shining grey o n both surfaces. 9 (Fig. 116). As d except for a pair of ventromedial fenestrae on fourth abdominal segment. Wing measurements. d, 5.6-6.4 mm (4). 0, 6.8-7.4 mm (4). Lectotype, 4.5 mm. Genitalia. d (Fig. 6 5 ) . Uncus papillate, each lobe pointed and sclerotized distally. Gnathos: as in Fig. 65(a); transverse ventral
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Philip J. Newton and Christopher Wilkinson
very large, anterior two-thirds covered with irregular chains of heavy pectinations as in Fig. 66( b); signum absent. Anterior apophyses very long and thin, arcuate. Posterior apophyses straight and narrow, not reaching apophyses anteriores. BIOLOGY. Immature stages unknown. Host. Braun ( 19 17: 175) suggested that the larvae probably mine oak since she collected imagines which were resting on the leaves of red oak this cannot be confirmed in the present study. Voltinism. Possibly bivoltine, with the majority of adults being taken in mid-June in Ohio and a single male o n Plummers’ Island (Maryland) in late July. ~
DIAGNOSIS. The strongly iridescent forewing and the marked change in colour after the fascia separate unifasciella from the other members of the genus in this study with a single fascia, except quercipulchella which has brown tufts on the vertex and a slightly more proximal fascia than in unifasciella. The male genitalia are similar to those of juglandifoliella and longisacca but differ in the absence
of the palmate anellar plates, the presence of the Y-shaped juxta and t h e long ventral arms of the transtilla in unifasciella. The long apophyses and the discrete margin t o the area of pectinations o n the bursa distinguish the female genitalia from the other members of the genus in this study: in particular the granular accessory sac differentiates t h e species from rosaefoliella and the extent of the pectinations differs in heteromelis. DISCUSSION. This species is notable f o r the marked differences in coloration of the forewing basally and terminally, which is also found t o a lesser degree in villosella and quercipulrhella. There are considerable differences in t h e alar expanse between males and females from Ohio and also between these specimens and Chambers’ syntypes from Kentucky: the latter may be due to local population variation. Further material is needed to evaluate these differences. DISTRIBUTION. U.S.A.: Kentucky, Ohio, Maryland, Illinois, Texas (Chambers), Massachusetts. Type material examined Here designated, lectotype 9, U.S.A.,
A taxonomic revision of Stigmella
FIG. 16. Stigmella procrastinella 9 , genitalia.
Kentucky (Chambers) (genitalia slide no. CNC 3488; MCZ). Paralectotype 1 ex., same pin as lectotype (abdomen missing).
Further material examined U.S.A.: Ohio, Cincinnati, 3 dd, 4 99, 1 2 . vi. 1908-13.vi. 1918 (Braun) (ANS); Maryland, Plummers’ Island, 1 d, 28.vii. 1962 (Hodges); Hlinois, Putnam County, 3 d d , 2 9 9 , 3 ex., 1O.vi. 1970-31 .viii. 1976; Massachusetts, Barnstable, 2 8 8 , 21 .vii. 1952, 18.vi.i. 1958 (Kimball) (USNM).
Stigmella condaliafoliella (Busck) (Figs. 6 7 , 6 8 and 119) Nepticula condaliafoliella Busck, 1900: 238; Braun, 19 17: 18 1 [ redescription] . Stigmella condaliafoliella (Busck); Grossbeck, 1917: 145. DESCRIPTION. External features. ds (Fig. 119). Palps whitish; antenna brown; tufts on front of head and vertex pale ochreous;
441
eye-caps shining white; collar white. Thorax dark brown with scattered purple reflections. Abdomen black above, shining silver beneath. Forewing: ground colour of dorsal surface dark brown with purple reflections becoming predominantly bronze beyond fascia; fringe well defined at apex by row of wing scales, brownish, shining silver at apex; single postmedial fascia, shining white, oblique and broadening slightly o n dorsal margin. Hindwing: ground colour and fringe pale brown, shining silver. Legs greyish, shining metallic grey, very pale o n lower surface. Wing measurements. 9 , 2.8 mm (1). Lectotype, 3.1 mm. Genitalia. d (Fig. 67). Uncus bilobed, lobes widely separated and flattened distally. Gnathos: broadly U-shaped as in Fig. 67(a): transverse ventral plate narrow with medial convex expansion; posteriorlydirected lateral arms narrow and arcuate, reaching tegumen. Tegumen a narrow strap-like plate. Vinculum: lateral arms tapering and articulating with tegumen at dorsal extremities ; ventral plate broad. Saccus deeply bilobed, each lobe as long as ventral plate is broad and tapering markedly. Valve broad basally and tapering, reaching beyond uncus, inner surface markedly concave distally. Transtilla: broadly W-shaped as in Fig. 67(a); lateral arms short; ventral arms short and s t o u t ; transverse bars continuous and arcuate. Aedeagus: regular in width, equal t o length of genital capsule; vesica with cornuti as many long, thin denticles and with a rectangular bearded plate of minute papillae. 9 (Fig. 68). Ductus bursae short and narrow. Accessory sac reduced, with a spiral duct arising distally. Bursa copulatrix : large and covered with sclerotized denticulate plates as in Fig. 68(b);signum absent. Anterior apophyses broad basally and tapering distally. Posterior apophyses straight and narrow, reaching just beyond apophyses anteriores. BIOLOGY. Host. Condalia ferrea (black ironwood). Mine. Described by Busck ( 1 900: 238) and Braun (1917: 1 8 1 , 206) as an upper-side, serpentine mine. DIAGNOSIS. The externals differ from those of tiliella in the iridescence of the forewing, which is more bronze and purple, and the
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Philip J. Newton and Christopher Wilkinson
FIG. 77. Stigmella braunel/Q d, genitalia: (a) genital capsule; (b) aedeagus.
fascia, which is more whitish. The male genitalia are similar to those of juglandifoliella and longisacca in the form of the valve and the large ventral plate, but may be easily separated by the absence of the anellar plates, the larger aedeagus and the deeply excavated saccus. The plate-like pectinations of the bursa distinguish the female genitalia from any other member of the genus in this study.
DISCUSSION. This species is similar to rnembers of the corylifoliella group in the externals and the overall form of the genital capsule in the male genitalia: however, the relatively large aedeagus, the absence of anellar plates and the form of the bursa exclude it from that group. DISTRIBUTION. U.S.A.: Florida.
Type material examined Here designated, lectotype 8, U.S.A., Florida, Palm Beach, o n Condalia ftrree, 14.ii. 1900 (Dyar) (genitalia slide no. CNC 3468; USNM). Paralectotypes 10, 1 ex.,
data as lectotype, 1O.ii. 1900 ( D y a r ) (genitalia slide no. CNC 3486 ; USNM).
Stigmella tiliella (Braun) c0mb.n. (Figs. 69, 70,95 and 118) Nepticula tiliella Braun, 1912: 9 0 ; Braun, 1917: 182 [redescription] ; Braun, in Forbes, 1923 : 9 1 [ redescription] . DESCRIPTION. External features. 6. Palps whitish; antenna dark brown; tufts on front of head and vertex ochreous; eye-caps shining white; collar shining white. Thorax and abdomen dark brown with gold reflections above, abdomen paler beneath and shining silver. Forewing: ground colour of dorsal surface brown with bronze and gold reflections becoming predominantly gold beyond fascia and irrorate, with each scale paler at the base; fringe well defined at apex by row of wing scales, greyish, shining silver; single postmedial fascia shining silver, slightly oblique and usually narrower in middle. Hindwing: ground colour very dark brown with purple
A taxonomic revision of Stigmella
reflections; fringe brownish. Legs dark brown with scattered ochreous patches, shining silver o n lower surface. 0 (Fig, 118). As d except for hindwing ground colour and fringe paler and shining metallic grey. A pair of ventromedial fenestrae o n fourth abdominal segment. Wing measurements. d, 3.6 m m (1). 0 , 3.24.0 mm (5). Lectotype, 3.4 mm. Genitalia. d (Fig. 69). Uncus bilobed, each lobe quadrate and heavily sclerotized. Gnathos: broadly U-shaped as in Fig. 69(a): transverse ventral plate very narrow with medial expansion; posteriorly-directed lateral arms horn-like and reaching beyond uncus.
443
Tegumen a broad ovate plate. Vinculum: lateral arms broad with triangular projection at point of articulation with tegumen dorsally; ventral plate broad. Saccus half width of ventral plate, weakly bilobed. Valve very broad with short, pointed style, inner margin of cuiller weakly concave. Transtilla: very short lateral arms; transverse bars fused. Aedeagus: flask-shaped, 1.5 times length of genital capsule; vesica with cornuti as many small denticles, evenly distributed, with several large spines posteriorly and with a comb-like rectangular plate of terminally furcate spines, also with a rectangular plate of minute papillae, see Fig. 69(c).
FIG. 78.Stigmeila braunefla 0,genitalia: (a) genitalia;(b) enlarged detail of markings o n bursa copulatrix.
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Philip J. Newton and Christopher Wilkinson
9 (Fig. 70). Ductus bursae broad and long. Accessory sac large, with a reticulate patch at vestibule and with duct arising proximally. Bursa copulatrix: large and covered with scallopshaped chains of pectinations on striations of the bursa as in Fig. 70(b); signum single comprising an ovate patch of granular, hexagonal cells as in Fig. 70(c). Anterior apophyscs straight and narrow. Posterior apophyses equal in length t o apophyses anteriores. BIOLOGY. Egg. Laid on upper surface of leaf. Host. Tilia arnericana (basswood). Mine. An upper-surface, long, linear mine with tendency toward a spiral form: a striking characteristic is the frequency of very angular turns. Frass deposited irregularly across entire breadth of mine in the first four-fifths
but as a dense, central line in terminal portion (Fig. 95). Larva. Emerges o n upper surface of leaf. Cocoon. Ovate, reddish brown. Voltinism. Bivoltine: adults being found in mid-June and early August or late July. Late instar larvae are found in late August and, according t o Braun (1917: 183), in early July. DIAGNOSIS. The externals are close to members of the corylifoliella group, in particular juglandifoliella, but may be separated b y the slightly more proximal fascia and the silver apical ciliae of the forewings. Both male and female genitalia possess several diagnostic features: the form of the uncus, transtillae and the comb-like cornuti of the male, and in the female the single ovate signum comprising whorls of hexagonal cells. The mine produced by the larvae differs from others
1:IG. 7 9 . Srigmella argentifasciella d , genitalia: (a) genital capsule with detail of spines; (b) aedeagus.
A taxonomic revision o f Stigmella
445
FIG. 80. Stigrnella argentifasciellu d, abdomen (dorsal) showing: (a) patches of heavily sclerotized scales on segments V1, VII and VI11; (b) enlarged detail of scales in situ; (c) scales highly magnified.
found on Tilia americana (produced by argentifasciella) b y the absence of the terminal blotch. DISCUSSION. Although similar in externals
to members of the corylifoliella group, the genitalia of tiliella possess several characters which are not found in the other members of the genus studied here (see diagnosis). DISTRIBUTION. U.S.A.: Ohio, Kentucky (Braun). T y p e material examined Here designated, lectotype 0 , U.S.A., Ohio, Clermont County, on Tilia americana, 12.viii. 1912 (Braun) (genitalia slide no. 50 PJN; ANS). Paralectotypes 2 99, data as lectotype, 11 + 12.viii. 1912 (Braun) (genitalia slide nos. 51 PJN, 5 2 PJN; ANS). Further material examined U.S.A., Ohio, Cincinnati, 2 dd, 3 99, 1 ex., 3 mines, on Tilia americana, 24.viii.1909, 17.vi.1911-29.vii.1916 (Braun) (ANS). Species-group QUERCIPULCHELLA
Described by Wilkinson & Scoble (1979); altella is added here. Contains only three species: these are distinct in aspects of both
male and female genitalia, in particular the hirsute membrane sheathing the extremely large aedeagus and the female accessory sac which is almost twice as large as the bursa and markedly spiral throughout its length. Key to males Saccus weakly lobed; gnathos without anteriorlydirected processes . . . . . . . . . . . . . . altella - Saccus bilobed, each lobe as long as broad; gnathos with broad anteriorly directed processes 2 2 Forewing with weak, whitish reflections; with postmedial fascia and terminal patch; male with purple scales o n the hindwing . . . . . . . variella - Forewing with shining silver reflections; without terminal patch; male hindwing without purple scales. . . . . . . . . . . . . . . . quercipulchella 1
Stigmella quercipulchella (Chambers) (Figs. 7 1 and 72) Nepticula quercipulchella Chambers, in Hayden, 1878a: 105. Stigmella quercipulchella (Chambers); Wilkinson & Scoble, 1979: 6 5 [redescription]. Nepticula terminella Braun, 1914: 23 [ synonymized by Wilkinson & Scoble, 1979: 651.
DESCRIPTION. (1979).
See Wilkinson & Scoble
446
Philip J. Newton and Christopher Wilkinson
BIOLOGY. Host. Quercus palustris, Q.marilandica, Q.rubra, Quercus s p . Mine. Described by Wilkinson & Scoble
( 1 979). Voltiriism. Bivoltine in the U.S.A. : probably bivoltine in Canada also, although the material represents only o n e generation. DIAGNOSIS. Differs from altella and variella in the darker tuft on the vertex and from variellu in the narrower silver fascia. the
absence of the terminal patch o n the forewing and the absence of the purple scales on the hindwing of the males. The genitalia (Figs. 7 1 and 72) are indistinguishable from those of variella, but separated from altella b y the anterior processes of t h e gnathos and the more deeply lobed saccus. S.varielZu exhibits sexual dimorphism which is not shown in quercipulchella and the two taxa mine different hosts which are also allopatric in their
-,
....(
0.02m m
FIG. 81. Stigmella argentifasciella 9 , genitalia: (a) genitalia; (b) enlarged detail of pectinations on bursa copulatrix.
A taxonomic revision of Stigmella
k
447
0.04mm
I
4 FIG. 8 2 . Stigmella plumosetaeella d, genitalia: (a) genital capsule; (b) aedeagus; (c) enlarged detail of setae on style and cuiller.
distribution. The distribution of variella is shown here t o b e wider than previously cited. DISTRIBUTION. U.S.A.: Kentucky, Ohio, Pennsylvania (Braun), Illinois. CANADA: Ontario. Material examined See Wilkinson & Scoble (1979). U.S.A.: Illinois, Putnam County, 7 dd, 4 9 9 , 4 ex., 14 .v. 1962-27. v . 1969 (Glenn);Pennsylvania, Allegheny County, Oak Station, l d , 17.v. 1910 (Marloffl (USNM); Ohio, Brachmann’s Woods, 1 mine, on Quercus palustris, 30.viii. 1 9 1 1 (Braun), Hazelwood, 1 mine, on Quercus marilandica, 27. x . 19 1 2 (Braun), St Thomas, 1 mine, on Quercus rubra, 9 . xi. 1912 (Braun) (ANS). CANADA, Ontario, Bells Corner, 5 mines, o n Quercus rubra, 1 4 . ix .1966 (Freeman & Lewis) (CNC).
Stigmella variella (Braun) Nepticula variella Braun, 1 9 10: 173 ; Opler 1974: 5 , 1 9 [biology]. Stigmella variella (Braun); Wilkinson &, Scoble, 1979: 6 7 [redescription]. 30
DESCRIPTION. See Wilkinson & Scoble (1979). BIOLOGY. Egg. Laid singly on veins often near midrib and usually o n lower surface. Opler (1974: 19) also recorded it on the upper surface. Host. Quercus agrifolia, Q. wistizenii plus hybrids and Q. kelloggii, Mine. Serpentine, in upper epidermis and mesophyll, becoming contorted and almost filled with frass granules, but n o terminal blotch is formed. Larva. Yellow with greyish brown head; its development time ranges from 2 to 4 months with three to eight instars though usually five or six. The larva emerges from either surface of the leaf, t o drop to the ground for pupation. Cocoon. White, darkening with age. Pupa dark brown. Voltinism. Bi- o r trivoltine in California. Completed mines can be found from July to early September and February t o April. DIAGNOSIS. Differs from quercipulchella in the tufts o n the front of the head and vertex,
448
Philip J. N e w t o n and Christopher Wilkinson
which are paler, in the fascia which is broader, and in the purple reflections of the male hindwings. Indistinguishable from quercipulchella in both male and female genitalia and in this respect the diagnosis of quercipulchella serves for this species also. DISCUSSION. Acquisition of a single specimen from Arizona extends the species’ known range beyond California. Biological information, additional t o that given by Wilkinson & Scoble (1979), is provided here, in part from Opler ( 1974: 19). DISTRIBUTION. U.S.A.: California, Arizona. Material examined See Wilkinson & Scoble (1979). U.S.A.: Arizona, Coconino County, 1 0 , 22. vi. 196 1
(Hodges) (ANS); California, Costa County, Antioch Contra, 1 0 , 15.v. 1958 (USNM). There is a single female (Specimen f), reared from ‘tan bark oak’, discussed by Wilkinson & Scoble (1979: 69). This specimen differs from other members of Braun’s type series of variella in the spicular accessory sac, see Fig. 73.
Stigmella altella (Braun) (Figs. 74, 96 and 120) Nepticula altella Braun, 1914: 2 1 ; Braun, 1917: 178 [redescription] ; Braun, in Forbes, 1923: 8 9 [redescription]. Stigmella altella (Braun); Wilkinson & Newton, 1981: 58. DESCRIPTION. External features. d (Fig.
FIG. 83. Srigmella plumosetaeella 9 , genitalia: (a) genitalia; (b) enlarged detail of pectinations on
accessory sac; (c) enlarged detail of spines on bursa copulatrix.
A taxonomic revision o f Stigmella
449
4 FIGS. 84-93. Larval mines of Stigmella spp.: 84, chalybeia on Pyrus c o m m u n i s ; 85, taeniola on Amelanchier alnifolia ; 86, ceanothi on Ceanothus divaricatus; 87, rhoifoliella on R h u s toxicodendron ; 88, rhamnicola on Rhamnus lanceolata; 8 9 , diffasciae on Csanothus sp.; 9 0 , villosella on Rubus sp.; 9 1 , apicialbella on UImus fulva; 9 2 , pallida on Salix sp.; 93,flavipedella on Quercus platanoides.
120). Palps greyish; antenna dark brown; tuft o n front of head ochreous, vertex paler; eyecaps shining white; collar whitish, sometimes pale ochreous. Thorax dark brown with purple reflections. Abdomen dark brown above, paler beneath. Forewing: ground colour of dorsal surface dark brown with bronze and purplish reflections becoming predominantly bronze after the fascia; fringe greyish brown, shining silver at apex ;single postmedial fascia, shining silver and usually broader at the margins. Hindwing: ground colour of dorsal surface brown with strong purple reflections; fringe brownish grey, shining silver grey; lanceshaped scaleless streak in centre of wing, basal t o medial; ventral surface brownish. Legs dark brown, paling t o buff o n tarsi, with scattered silver reflections. Wing measurements. d , 5.6-6.4 mm (13). Holotype, 5.8 mm. Genitalia. d (Fig. 74). Uncus bilobed with lobes widely separated. Gnathos: as in Fig. 74(a); transverse ventral plate narrow; posteriorlydirected lateral arms short and broad; dorsolateral arms very broad, with large posteriorly-directed processes. Tegumen
an ovate plate with serrate posterior margin. Vinculum: lateral arms broad and expanded at point of articulation with tegumen; ventral plate broad, Saccus narrow, weakly bilobed. Valve divided into short style and pointed cuiller, extending just beyond uncus. Transtilla: lateral arms long and arcuate; ventral processes very small; transverse bars fused. Juxta a quadrate, membranous plate, broadening posteriorly. Aedeagus: flaskshaped almost twice length of genital capsule, enclosed posteriorly by hirsute manica as in Fig. 74fb); vesica with cornuti as many large denticles orientated in a ridge posteriorly and with a rectangular plate o f minute papillae. BIOLOGY. Egg. Laid o n lower surface of leaf. Host. Quercus im bricaria ;Quercus palustris. Mine. Much contorted, linear, o n lower surface of leaf, barely visible from upper side; relatively longer than others found on Quercus spp. and narrow throughout its length; frass scattered across entire breadth of mine (Fig. 96). Larva. Emerges o n lower surface of leaf. Pupa. Dark brown.
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Philip J. Newton and Christopher Wilkinson
Voltinism. Univoltine in Ohio, with the mines being collected in October and adults emerging the following May and early June. The single specimen examined from Maine was collected 1 month later than Ohio material: this may represent a later generation although t h e difference in latitude should be taken into account. DIAGNOSIS. The externals are similar to those of quercipulchella but the vertex is ochreous and the forewing without brilliant reflections. The scaleless streak o n the hindwing separates these males from any others in this study. The male genitalia are very similar t o those of quercipulchella and variella in the overaH form and the presence of the manica; altella differs in t h e absence of anteriorlydirected processes o n the gnathos, t h e shorter style, and the more shallow lobing of the saccus. The mines of altella differ from those of quercipulchella in that they are more contorted, narrower and the frass is diffusely deposited. DISCUSSION. This species is closely related t o quercipulchella and variella in both externals and male genitalia; the choice of host plants, as observed, differs only at the species level. Further collection is necessary, particularly t o investigate the biology and the female genital morphology, before the true relationship of altella within the group may be determined.
DISTRIBUTION. U.S.A.: Ohio, Maine. B p e material examined Holotype d, U.S.A., Ohio, Cincinnati, on Quercuspalustris, 11 .v.1913 (Braun) (genitalia slide no. 25 PJN; ANS). Paratypes 2 dd, same data as holotype, 11 .v. 1913 (Braun), 6 dd, Ohio, Cincinnati, 3.vi. 1904-15.v. 1908 (Braun) (genitalia slide nos. 26 PJN, USNM 17333-17334; ANS), 1 d, Ohio, Cincinnati, 20.v. 1905 (Braun) (genitalia slide no. CNC 3485; DFF).
Further material examined U.S.A.: Ohio, Cincinnati, 1 d, 24.v. 1903 (Braun), 1 d, 2 mines: o n Quercus palustris, 1 3 . x . 1 9 1 2 , 15.v.1916 (Braun), 1 d, 1 mine, on Quercus imbricaria, 28.x.1917, 1 .v. 1918 (Braun); Maine, Rockport, 1 d, at light 3.vii. 1941 (Hebard) (ANS). Species-group PROCRASTINELLA
Described by Wilkinson & Scoble ( 1 979) and including t w o species: procrastinella and alha. The group is characterized by the form of the uncus and gnathos in t h e male, and the unique, heavily pectinate patch of spines in the female. Although similar to Ectoedemia in aspects of the male genitalia, .the major generic characters recommend that the species are better placed in Stigmella. The patch of spines in the female, referred t o as the signum by Wilkinson & Scoble (1979), may be part of the bursa copulatrix o r t h e accessory sac; the
94
FIGS. 94-98. Larval mines of Stigmella spp.: 94, sclero$tylota on Quercus alba; 95, tiliella on Tilia americana; 96, altella on Quercus palustris; 9 1 , braunella on Prunus ilicifolia; 98, argentifasciella on Tilia americana.
A taxonomic revision of Stigmella membranous sac t o which they are attached lacks any of the small pectinations usually associated with the bursa in other Species; therefore it is possibly the accessory sac and the bursa is presumably unadorned.
Key to species - Forewing off-white with a medial patch of brown
. . . . .’. . . . . . . . . . . . . . . . d b a fascia, shining silver . . . . . . . . procrustinella
scales.
.
- Forewing greyish brown with a postmedial
Stigmella procrastiaella (Braun) (Figs. 75 and 76) Nepticula procrastinella Braun, 1927: 59. Stigmella procrastinella (Braun); Wilkingon & Scoble, 1979: 70 [redascliption]. DESCRIPTION. See Wilkinson & Scoble (1979). DIAGNOSIS. The form of the u n c w and gnathos of the male (Fig. 75) separate this species from any other in the genus in this study except alba and brauhella : they resemble Ectoedemia. The larger aedeagus, the spinelike cornuti of the vesica and the longer ventral arms of the transtilla in procrastinella separate it from braunella. The heavily sclerotized, spinose ‘signum’ separates the female genitalia (Fig. 7 6 ) from those of any other species studied here. This species differs from alba in the absence of an expanse of off-white scales o n the forewlng. DISTRIBUTION. U.S.A.: Virginia, Mahe. CANADA: Ontario. Material examined See Wilkinson & Scoble (1979). U.S.A., Maine, Kokadjo, 1 d, 1 ? , 7 + 2 1 .vii.? (USNM).
Stigmella alba Wilkinson & Scoble Stigmella alba Wilkinson & Scoble, 1979: 73. DESCRIPTION. See WilkinSon & Scoble (1979). DIAGNOSIS. The off-white thorax and scales of the forewing separate this species from procrastinella. The geriitalih are indistinguishable from those of procrastinella and i n this respect the diagnods of thtit’species serves to separate alba from braunellfi. DISTRIBUTION. U.S.A.: Arizona. British Columbia.
CANADA:
45 1
Material examined See Wilkinson & Scoble (1979).
Stigmella braunella (Jones) (Figs. 77, 78, 97 and 121) Nepticula braunella Jones, 1933: 4 9 ; Rindge, 1947: 25 [ lectotype designated] . Stigmella braunella (Jones); Wilkinson & Scoble, 1979: 13 [footnote]. DESCRIPTION. External features. 6. Palps brownish grey; antenna very dark brown; tuft on‘ front of head dark ochre-brown, vertex pale ochreous; eyecaps shining white; collar off-white. Thorax dark greyish brown with bronze reflections. Abdomen greyish, shining metallic grey. Forewing: ground colour of dorsal surface dark brown with bronze reflections, each scale greyish at base producing an irrorate effect; fringe brownish grey, shining metallic grey; pair of medial streaks marginally, sometimes greatly reduced, shining white. Hindwing: ground colour and fringe brownish grey, shining metallic grey. Legs dark grey, shining metallic grey beneath, tarsi annulate with paler areas. 0 (Fig. 121). As d except for forewing markings which usually meet t o form a biconcave fascia. A pair of ventromedial fenestrae o n fourth abdominal segment. Wing measurements. d, 5.6-6.6 mm (4). 0 , 5.4-6.4 m m ( 7 ) . Genitalia. d (Fig. 77). Uncus with single lobe, tapering markedly and terminally papillate as in Fig. 77(a). Gnathos: an inverted V-shape as in Fig. 77(a); transverse ventral plate produced into large medial expansion; dorsolateral arms broad with long posteriorlydirected processes. Tegumen strap-like with medial expansion. Vinculum: lateral arms narrow, articulating with tegumen at dorsal extremities; ventral plate broad. Saccus markedly bilobed. Valve divided into narrow, pointed style and broad cuiller, not reaching uncus. Transtilla: lateral arms long; ventral arms reduced; transverse bars fused t o form an arcuate strap. Aedeagus: narrow and regular in width, approximately half length of genital capsule; vesica with cornuti as many small denticles and with a rectangular plate of minute spicules. ? (Fig. 78). Ductus bursae short and broad. Accessory sac large with reticulate patch at vestibule and with duct arising distally. Bursa
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Philip J. Newton and Christopher Wilkinson
1:IGS. 99-107. Exterhal features of Stigmellu spp.: 99, chalybeia P ; 1 0 0 , purpuratella d ; 101, taeniola d ; 102, ceanothi 9 ; 103,rhoifoZiella d ; 1 0 4 , rhumnicola 0,winter generation; 1 0 5 , rhumnirola 0 , summer generation; 106, itTcon&icuella 9 ; 1 0 7 , diffasciae 0.
A taxonomic revision of Stigmella copulatrix: covered with small chains of pectinations on scallop-shaped plates as in Fig. 78(b); signum absent. Anterior apophyses tapering markedly from broad base. Posterior apophyses long and straight, reaching beyond apophyses anteriores. BIOLOGY. For a more detailed account of t h e life-history see Jones (1933) which is summarized here. Egg, Usually laid o n lower surface of leaf but occasionally o n upper surface, generally nearer mid-rib than margin. Host. Prunus ilicifolia (islay, evergreen or holly-leaved cherry); and the variety in tegrifolia. Mine. Upper-surface, linear o r serpentine, convoluted terminally and i n this region frequently crossing itself or forming a blotch; gradually increasing in breadth throughout its length. Frass deposited centrally as a continuous line (Fig. 97). Larva. Emerges o n upper surface of leaf. Cocoon. Buff or whitish. Voltinism. There are two broods per year with late instar larvae being encountered throughout the year. DIAGNOSIS. The externals are easily differentiated from the other members of Stigmella in this study by the forewing markings of the males and the darker tufts o n the front of the head but not the vertex. The females are superficially similar t o those of the rosaefoliella group but are larger and lack the reflections of the forewing. The genitalia are similar t o those of procrastinella and alba in the form of t h e gnathos and the uncus, but t h e form of the valve and t h e aedeagus are diagnostic. DISCUSSION. This is a very distinctive species which possesses some male characteristics more typical of Ectoedemia than Stigmella. However, as in the cases of argentifasciella, procrastinella and alba the major generic characters indicate that braunella is correctly placed in Stigmella: braunella is not included 'in the procrastinella group because of the differences in female genitalia. DISTRIBUTION. U.S.A.: California. T y p e material examined Paralectotypes 7 6d, 2 99,3 4 ex., U.S.A., California, Berkeley, 30.iv. 1924-12.iv. 1929
453
(Jones) (genitalia slide nos. 95-97 PJN, USNM 17288- 17289, USNM 1743 1 ; USNM), 1 d, 5 99, same locality,8.iii. 1927-9.iv. 1929 (Jones) (genitalia slide no. T N F 4 2 1 ; CNC). Further material examined U.S.A., California, Berkeley, 1 d, o n Prunus ilicifolia, iv. 1924 (Braun) (ANS), 1 d, 19, on Prunus sp., 2 0 + 22.v. 1929 (Jones) (CNC).
Stigmella argentifasciella (Braun) c0mb.n. Figs. 79-8 1,98 and 122) Nepticula argentifasciella Braun, 1912: 100; Braun, 1917: 166 [redescription] ; Braun, in Forbes, 1923 : 86 [ redescription] . DESCRIPTION. External features. 6. Palps black; antenna very dark brown t o black; t u f t o n front of head ochreous, vertex sometimes darker; eye-caps shining white; collar dark brown. Thorax and abdomen very dark brown, thorax with gold and purple reflections, abdomen with metallic grey reflections above, shining grey beneath. Abdomen with lunulate patches of heavily sclerotized scales dorsally o n segments VI, VII and VIII as in Fig. 80. Forewing: ground colour of dorsal surface dark brown with bronze lustre; fringe brown, shining metallic grey; three fasciae, shining silver, basal extending to one fifth, medial widening o n dorsal margin, terminal variable in extent along dorsal margin. Hindwing: ground colour and fringe greyish brown, shining metallic grey. Legs dark greybrown, paling o n tarsi, shining metallic grey o n both surfaces. 9 (Fig. 122). As 6 but lacking lunulate patches on dorsal surface of abdomen. Wing measurements. d, 4.2-4.6 mm (4). ?,3.6-4.8 mm ( 1 2). Lectotype, 4.2 mm. Genitalia. d (Fig. 79). Uncus flattened with two widely spaced lateral papillae. Gnathos: broadly U-shaped as in Fig. 79(a); transverse ventral plate very narrow; posteriorly-directed lateral arms long and narrow. Vinculum: lateral arms very broad and fusing t o form a complete ring, with the tegumen reduced to a narrow posterior sclerite barely separated from the vinculum; ventral plate broad. Saccus markedly bilobed, each lobe as broad a t base as ventral plate and as long as broad. Valve reaching just beyond uncus, broad and quadrate, with a short pointed style not
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Philip J. Newton and Christopher Wilkinson
110
Is'IGS. 108-115. External features of Stigmella spp.: 108, gossypii 0; 1 0 9 , herevomelis 9 ; 110,cevea 0: I l l , villosella d: 112,pallida 0; 113,apicialbella d ; 114,nigriverticella 9 ; 115,flavipedella 0.
reaching beyond the cuiller, and with distal margin of cuiller papillate as in Fig. 79(a). Transtilla: lateral arms long and broad;ventral arms very short and stout; transverse bars
fused. Juxta a membranous quadrate plate, denticulate as in Fig. 79(a). Aedeagus: flaskshaped, twice length of genital capsule; vesica with cornuti as many short, stout
A taxonomic revision of Stigmella denticles evenly distributed, and with rasp-like plate of minute papillae. 9 (Fig. 81). Ductus bursae short and broad. Accessory sac very large with reticulate patch at vestibule and with large, spiral duct arising distally. Bursa copulatrix: very large and covered with short chains of pectinations as in Fig. 81(b); signum absent. Anterior apophyses narrow. Posterior apophyses equal in breadth and reaching just beyond apophyses anteriores. BIOLOGY. Egg. Upper surface. Host. Tilia americana. Mine. At first within parenchyma of leaf and barely visible o n upper surface; later broadening into a blotch o r broad tract and becoming transparent; frass deposited in sinuous curves (Fig. 98). Larva. Emerges o n lower surface of leaf. Cocoon. Reddish. Voltinism. Braun (1917: 167) was of the opinion that there are two or possibly three generations per year: full-grown larvae being found in June, August and September. The material examined here represents only t w o broods: adults of the third generation, if it exists, should be sought between mid-July and mid-September. DIAGNOSIS. The brilliant reflections of the forewing markings and the abdominal patches in the male separate argentifasciella from the other members of the genus in this study: the form of the valve and vinculum are characteristic of the male genitalia. The mine produced by the larvae of argentifasciella differs from others found o n Tilia americana, produced by tiliella, in the presence of the terminal blotch. DISCUSSION. The form of the vinculum and the degree of fusion with the tegumen resembles Ectoedemia, though the female genitalia and all other generic characters indicate that this species is correctly placed in Stigmella. DISTRIBUTION. (Braun), Illinois.
U.S.A.:
Ohio, Kentucky
Type material examined Here designated, lectotype d, U.S.A., Ohio, Cincinnati, o n Tilia americana, 12.vii. 1912 (Braun) (genitalia slide no. 14 PJN; ANS). Paralectotypes 2 dd, 10 99, 2 ex., data as lectotype, 9-18.vii. 1912 (Braun)(genitalia
455
slide nos. 15- 1 8 PJN; ANS), 2 dd, 2 99, data as lectotype, 9 + 1 2 .vii. 19 12 (Braun) (genitalia slide no. CNC 3481 ; DFF). Further material examined U.S.A.: Ohio, Cincinnati, 1 9 , 31.v.1912 (Braun), 1 mine, o n Tilia arnericana, 22.ix. 1909 (Braun) (ANS); Illinois, Putnam County, 1 d, 3 ex., 9.vi.1964-19.vii. 1965 (Glenn) (USNM).
Stigmella plumosetaeella sp.n. (Figs. 8 2 , 83 and 123) DESCRIPTION. External features. d? (Fig. 123). Palps greyish; antenna pale grey-brown; tuft on front of head pale ochreous, vertex darker; eye-caps shining white; collar white. Thorax and abdomen dark grey-brown, abdomen shining metallic grey beneath. Forewing: ground colour of dorsal surface dark brown with bronze lustre; fringe greyish, shining metallic grey ; single medial fascia, pale yellow and very broad, followed by small apical spot, off-white. Hindwing: ground colour and fringe greyish, shining metallic grey. Legs pale greyish brown, shining metallic grey on lower surface. Wing measurements. 0 , 3.4 mm (1). Holotype, 3.4 mm. Genitalia. d (Fig. 82). Uncus bilobed, each lobe very large and ear-like. Gnathos: H-shaped as in Fig. 82(a): transverse ventral plate broad; posteriorlydirected lateral arms long and tapering; dorsolateral arms broad basally with short posteriorly- and anteriorlydirected processes. Tegumen a broad plate with medial convex expansion. Vinculum: lateral arms broad, tapering markedly to point of articulation with tegumen at dorsal extremities; ventral plate broad. Saccus quadrate, broader than ventral plate. Valve: reaching well beyond uncus and markedly bifurcate distally into narrow style and pointed cuiller; style and dorsal surface of cuiller bear plumose scales as in Fig. 82(c). Transtilla: lateral arms broad and straight; ventral arms short and stout; transverse bars meeting but not fusing. Aedeagus: bulb-shaped, approximately equal to length of genital capsu1e;vesica with cornuti as many small denticles and with a rectangular plate of minute spicules, 0 (Fig. 83). Ductus bursae short and narrow. Accessory sac very large and covered with denticles as in Fig. 83(c), with duct
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Philip J. Newton and Christopher Wilkinson
arising distally. Bursa copulatrix: very small with short chains of pectinations distally as in Fig. 83(b); signum absent. Anterior apophyses short and narrow. Posterior apophyses narrow, reaching well beyond apophyses anteriores. BIOLOGY. Immature stages and host unknown. DIAGNOSIS. The externals are nearest to those of latifasciella and nigriverticella but may be differentiated from them by the ochreous vertex and the intermediate width of the forewing fascia. The genitalia of both male and female show several diagnostic characters: the form of the valves, uncus, aedeagus and the large spiculate accessory sac. IIISCUSSION. Although the external features show similarities to other members of the genus in this study, the genitalia do not indicate any particular relationship to other species represented. IIISTKIBUTION. U.S.A. : Arizona.
Type material Holotype d, U.S.A., Arizona, Superior, 13.vii. 1939 (Braun) (genitalia slide no. 28 PJN; ANS). Paratype 0 , locality as holotype, 12.vii. 1939 (Braun) (genitalia slide no. 29 PJN; ANS). The following three species have not been assigned t o species-groups because the structures of the genitalia are unknown, and insufficient information is available regarding their biology. They have been omitted from the keys for the same reasons.
Stigmella resplendensella (Chambers) cornb .n. Nepticula resplendensella Chambers, 1875a: 118; Chambers, in Hayden, 1878b: 158 [listed] ; Braun, 1917: 175 [redescription, types not examined] ; Braun, in Forbes, 1923: 88 [redescription]. DESCRIPTION. External features. Palps and probably antenna whitish; tuft o n front of head and vertex orange-ochreous; eye-caps probably shining white; collar dark greyish brown. Thorax dark greyish brown. Forewing: ground colour of dorsal surface dark brown with shining bronze, purple and silver reflections. Iridescence and colours depend on angle and intensity of light source. A large
spot extending from postmedial to terminal overlaps the fringe anteriorly and extends backwards for two-thirds width of wing; often shining purple. Hindwing ground colour and fringe greyish brown, shining metallic grey. Legs brown to pale brownish yellow; hind legs with a t least one pair of spurs. Wing measurements. Lectotype, 6 mm. Genitalia. d? Unknown. BIOLOGY. Host. Celtis occidentalis (hackberry tree)? Mine. Unknown. DIAGNOSIS. Differs from unifasciella, with which this species has hitherto been associated, in the presence of an iridescent purple spot located on the forewing which otherwise shines randomly silver and gold. S.unifasciella has a definite silver fascia dividing a basal bronze area from a terminal dark brown area. DISCUSSION. Braun (1917: 175) likened this species to unifasciella and implied t h a t they may even be synonymous but she did not examine any material of resplendensella. C.W. has examined the lectotype and found it to be quite different from t h e types of unifasciella. However, whilst this species is unlikely t o be conspecific with unifasciella, we can say little more about it, having examined only t h e one syntype remaining from t w o originals. This specimen has the head, abdomen and four legs missing. More minor features, n o longer obtainable from the specimen, yet given in the description, come from earlier writings cited above. The affinities of this species therefore remain in doubt. DISTRIBUTION. U.S.A.: Kentucky?
Type material examined. Here designated, lectotype ex., U.S.A., Kentucky, 23.v.?1875 (Chambers) (Type 14954; ANS).
Stigmella belfrageella (Chambers) cornb.n . Nepticula helfrageella Chambers, 1875b: 7 5 ; Chambers, in Hayden, 1878b: 157 [listed] ; Braun, 19 17 : 193 [ redescription, no material examined] . DESCRIPTION. External features. Antenna brown; tuft o n front of head and probably vertex pale yellow; eye-caps white; collar probably grey. Thorax and perhaps abdomen pale grey. Forewing: ground colour of dorsal
A taxonomic revision of Stigmella
122
457
123
FIGS. 1 1 6 - 1 2 3 . External features of Stigmella spp.: 1 1 6 , unifasciella 9 ; 117, longisacca 9 ; 1 1 8 , tiliella 9 ; 119, condaliafoliella 4; 120,altella d ; 1 2 1 , braunella 9 ; 1 2 2 , argentifasciella 9 ; 1 2 3 , plumosetaeella d.
surface pale grey proximally, darkening distally; fuscous spot present o n wing apex. Hindwing and legs most likely pale grey.
DIAGNOSIS. Uncertain; perhaps the fuscous terminal spot on the otherwise grey forewing is diagnostic.
Wing measuremenfs. Type@, 6.25-6.5mm. Genitalia: SQ Unknown. BIOLOGY. The adult appears in April. Host and immature stages unknown.
DISCUSSION. No specimens of this species are known t o exist. We have tried t o locate the type material without success and it would seem that it is no longer extant. There was
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Philip J. Newton and Christopher Wilkinson
probably more than one type specimen since a range of wing measurements was given by Chambers: 2% to more than 3 lines (one line = one twelfth of an inch). The species may be distifict but so little is known about it. Checking of the original description against all species known from Texas (the type-locality) revealed none that exactly correspond. The nearest is perhaps nigriverticella and the closest Ectoedemia is grandisella. Thus the evidence prevents a synonymy and leaves in the literature the unsatisfactory situation of a species name to which no known material can be associated. It is only on the tenuous basis of Chambers calling it ‘Nepticula’ that we place belfrageella in Stigrnella. DISTRIBUTION. U.S.A.: Texas. Material examined None. Type locality: County (Chambers).
Texas,
Bosque
‘Nepticula’amelanchierella Cleme n s Nepticula amelanchierella Clemens, 1862a: 84 [description of mine] ; Clemens, in Stainton, 1872: 173; Packard, 1889: 3 5 6 ; Chambers, in Hayden, 1878b: 157 [listed] ; Braun, 1917: 205; Braun, in Forbes, 1923: 95 [redescription] ; Wilkinson & Newton, 1981: 69. DISCUSSION. This species was described by Clemens (1862a: 84) from mines on Amelanchier (service berry) in June and July. Whilst there have been many references t o this species in the literature, it still remains poorly described and its generic identity doubtful. The original (type) material is not preserved and there are no known bred specimens. Braun ( 19 17: 205) said that untenanted mines have been collected in Ohio, Kentucky and North Carolina. By this she presumably meant ophionomes corresponding with the original description. In her rearing notes (904) for 1915 she reported that all the mines were empty by 9 July in Kentucky. In 1928 she wrote in her notes (1382): ‘Mineral Springs, A d a m Co., 0. ix.7.28. Amelanchier. Long linear transparent mine about 1% mm wide at the end. Frass in a rather broad line through middle, cocoon pale brownish ochreous. Mine does not look just like that of bifasciella [now = prunifoliella] in wild cherry as it
does not have the closely contorted and blotch-like appearance at the beginning.’ In 1930 there is a similar description (1407) except: ‘Frass dispersed in rather broad line at beginning and toward end of mine; more dispersed in middle part, occupying more than half breadth of mine. Cocoon yellow brown.’ The significance of these remarks is that there is n o reference t o the formation of a blotch. The two other species known from Amelnnchier produce ophistigmatonomes with a pronounced blotch. The first, Stigrnella taeniola, is represented by only two specimens bred by Braun from Amelanchier alnifolia. The second is Ectoedemia nyssaefoliella: although normally recorded from Nyssa sylvatica, two female specimens reared by Heinrich from Amelanchier sp. have been tentatively placed as this species (Wilkinson & Newton, 1 9 8 1). Since both these samples were reared from blotch mines on Arnelanchier, and amelanchierella reputedly only makes a serpentine mine, it seems probable that three different species exist. Therefore to propose a synonymy and designate a neotype from known material is probably unwarranted. Unfortunately, there is not enough evidence to know if this species should be placed in Stigrnella o r Ectoedernia. This species is evidently very spasmodic in its appearance, as is Ectoedernia bradfordi Emmet in Britain, which is thought t o be associated with the European service tree (Sorbus torminalis). Acknowledgments
Acknowledgment is d u e t o the following persons and their respective institutes for the loan of the material: Dr D. F. Hardwick, Agriculture Canada, Ottawa; Mr 0. Lindquist, 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; the late Dr A. F. Braun, Cincinnati, Ohio; Dr K. Sattler, British Museum (Natural History), London; and the Director of the Museum of Comparative Zoology, Cambridge, Massachusetts (where some type material is deposited). P.J.N. I am deeply indebted t o Professor
A taxonomic revision of Stigmefla Christopher Wilkinson for his unstinting help and supervision of this project, and to Portsmouth Polytechnic, Hampshire, for providing facilities during my tenure as his research assistant. I express my thanks to Dr Malcolm J . Scoble, Transvaal Museum, Pretoria, for many hours of useful discussion while at Portsmouth, Col. A. M. Emmet, Essex, for his helpful discussions and t o Mr S. N. A. Jacobs, Bromley, Kent, for his meticulous illustrations of adult moths. A large part of this study was included in the project successfully submitted to the Council for National Academic Awards for the degree of Doctor of Philosophy (1980). C.W. My thanks are also due to Mej. R. Kloos, B. J. van Cronenburg and A. Rol, who assisted with the preparation and examination of a large number of additional specimens which were subsequently made available. Mej. Kloos also prepared a number of the illustrations of genitalia. Mej. Silvia Richter kindly typed the manuscript and Dr. Georgina Bryan helped with literature searches. Miss Pamela Gilbert, Librarian, Department of Entomology, British Museum (Natural History), was always willing to give us the benefit of her experience in solving bibliographic problems. References Beirne, B.P. ( 1 9 4 5 ) The male genitalia of the British Stigmellidae (Nepticulidae) (Lep.). Proceedings of the RoyalIrish Academy, B, 5 0 , 1 9 1 - 2 1 8 . Borkowski, A. ( 1 9 7 2 ) Studien an Nepticuliden (Lepidoptera). Teil IV. Bemerkungen Zur Nomencaltur und Systematik der Familie Nepticulidae. Polskie Pismo Entomologiczne, 4 2 , 6 8 9 - 1 0 9 . Braun, A.F. ( 1 9 1 0 ) New species of Tineina from California. Entomological News, 2 1, 17 1-1 7 9 . Braun, A.F. (1912) Notes o n North American species of Nepticula with descriptions of new species (Lepidoptera). Journal of the Cincinnati Society of Natural History, 2 1, 84- 10 1 . Braun, A.F. ( 1 9 1 4 ) Notes on North American species of Nepticula with descriptions of New Species (Lepidoptera). Canadian Entomologist, 4 6 , 1724.
Braun, A.F. ( 1 9 1 6 ) Nepticula rhamnicola nom. nov. Entomological News, 27, 5 6 . Braun, A.F. ( 1 9 1 7 ) Nepticulidae of North America. Transactions of the American Entomological Society, 4 3 , 1 5 5 - 2 0 9 . Braun, A.F. ( 1 9 2 3 ) Superfamily Nepticuloidea Family 5 Nepticulidae. In: Forbes, W.T.M. (ed.) Lepidoptera of New York and Neighbouring States, pp. 7 9 - 9 8 . New York. Braun, A.F. ( 1 9 2 5 ) Microlepidoptera of Northern Utah. Transactions of the American Entomological Society, 51, 1 8 3 - 2 2 6 .
459
Braun, A.F. ( 1 9 2 7 ) New Microlepidoptera from Ontario. Canadian Entomologist, 59, 5 6 - 5 9 . Braun, A.F. ( 1 9 3 0 ) Notes o n new species of Microlepidoptera from the Mineral Springs Region of Adams County, Ohio. Transactions of the American Entomological Society. 5 6 , 1-17. Brown, F.M. ( 1 9 6 4 ) Dates of publication of the various parts of the Proceedings o f the Entomological Society of Philadelphia. Transactions of the American Entomological Society, 8 9 , 3 0 5 308.
Busck, A. ( 1 9 0 0 ) New species of moths of the superfamily Tineina from Florida. Proceedings of the United States National Museum, 2 3 , 2 2 5 - 2 5 4 . Busck, A. ( 1 9 0 1 ) Nepticula pomivorella Packard; alias Micropteryx pomivorella Packard. Canadian Entomologist, 33, 5 2 . Busck, A. ( 1 9 0 3 ) Notes on Brackenridge Clemens' Types of Tineina. Proceedings of the Entomological Society of Washington, 5, 1 8 1 - 2 2 0 . Busck, A. ( 1 9 1 3 ) Tw o Microlepidoptera injurious t o Chestnut. Proceedings of the Entomological Society of Washington. 1 5 , 1 0 2 - 1 0 4 . Chambers, V.T. ( 1 8 7 3 ) Micro-Lepidoptera. Canadian Entomologist, 5, 1 2 4 - 1 2 8 . Chambers, V.T. (1875a) Tineina of the Central United States. Cincinnati Quarterly Journal of Science, 2 , 97- 12 1 . Chambers, V.T. ( 1 8 7 5 b ) Tineina from Texas. Canadian Entomologist, 4, 73-7 5 . Chambers, V.T. (1878a) New Tineina from Texas, and others from more Northern Localities. In: Hayden, F.V. (ed.) Bulletin of the United States Geological and Geographical Survey of the Territories, 4, 7 9 - 106. Chambers, V.T. (1878b) Index to the described Tineina o f the United States and Canada. In: Hayden, F.V. (ed.) Bulletin of the United States Geological and Geographical Survey of the Territories, 4 , 1 2 5 - 16 7. Chambers, V.T. (1880) Descriptions of some New Tineina, with notes o n a f e w old species. Journal of the Cincinnati Society o f Natural History, 2, 179- 194.
Clemens, B. (1862a) Micro-Lepidopterous larvae. Notes o n a f e w species, t h e imagos of which are probably undescribed. Proceedings of the Entomological Society of Philadelphia, 1 ( 1 861), 75-87.
Clemens, B. (1862b) New American Microlepidoptera. Proceedings of the Entomological Society ofPhiladelphia, 1, 1 3 1 - 1 3 7 . Clemens, B. ( 1 8 6 5 ) North American Micro-Lepidoptera. Proceedings of the Entomological Society ofPhiladelphia, 5, 1 3 3 - 1 4 7 . Clemens, B. ( 1 8 7 2 ) In: Sta inton, H.T. (ed.) The Tineina of North America - being a collected edition of his writings o n that group of insects. With notes by the editor. London. Davis, D.R. ( 1 9 7 8 ) New leaf-mining moths of the family Nepticulidae from Florida. Florida Entomologist, 6 1 , 2 0 9 - 2 2 4 . Davis, D.R. ( 1 9 7 9 ) Manoneura, a new name t o replace the generic homonym Oligoneura Davis (Lepidoptera: Nepticulidae). Florida Entomologist, 6 2 , 2 7 6 . Dyar, H.G. ( 1 9 0 3 ) A list of North American Lepidoptera and key t o the literature of this order of Insects. Bulletin of the United States National Museum, 5 2 , 1-723. Fabricius, J.C. ( 1 7 7 5 ) Systema Entomologiae. Forbes, W.T.M. & Leonard, M.D. ( 1 9 3 0 ) A new leaf-miner of Cotton in Porto Rico (Nepticula
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Philip J. Newton and Christopher Witkinson
gossypii msp.). Journal of the Department o f Agriculture, Porto Rico, 14, 149-157. Freeman, T.N. ( 1 9 6 7 ) A new species of Nepticula on Bur Oak in Ontario (Nepticulidae). Journal of Research on the Lepidoptera, 6 . 19-2 1 . b'rey, H. & Boll, J . ( 1 8 7 6 ) Kinige Tineen aus Texas. Stettiner Entomologische Zeitung, 31, 2 0 9 -2 2 8 . Vrey, H. & Boll, J . ( 1 8 7 8 ) Tineen aus Texas. Stettiner Entomologische Zeitung, 39, 2 4 9 - 2 8 0 . Goeze, J.A.E. ( 1 7 8 3 ) Entomologische Beytrage zu des Ritter Linnr Zwolften Ausgabe des Natursystems. Leipzig. Grossbeck, J.A. ( 1 9 1 7 ) 1. Insects of Florida. I V . Lepidoptera. (ed. by F. E. Watson). Bulletin o f the American Museum of Natural History, 37, 1-147.
Herrich-Schaffer, G.A.W. ( 1 8 6 0 ) Nepticula rhamnclla. Correspondenz b f a t t f u r Sammler won Insecten, 1860, 6 0 . Heyden, C. von ( 1 8 4 3 ) Amtlicher Bericht der Versammlung der Naturforscher zu Mainz. Johansson, R. ( 1 9 7 1 ) Notes o n Nepticulidae (Lepdioptera). I . A revision of the Nepticula nificapitella group. Entomologica Scandinavica, 2, 2 4 1 - 2 6 2 .
Jones, W.W. ( 1 9 3 3 ) The description and biology of Nepticula braunella n s p . (Lepidoptera: Nepticulidae), a species of leaf miner o n Pmnus ilicifolia Walp. and the variety integrifolia Sarg. University of California Publications in Entomology, 6 , 49-78. Karsholt, 0. & Schmidt Nielsen, E. ( 1 9 7 6 ) Systematisk fortegnelse over Danmarks sommerfugle. Catalogue of the Lepidoptera of Denmark. Klampenborg. Kearfott, W.D. ( 1 9 0 8 ) New North American Tortricidae and Tineina. Journal of the N e w York Entomological Society. 1 6 , 1 6 7 - 1 8 8 . Kirby, W.F. ( 1 8 9 7 ) LIoyds Natural History. A Handhook of the Order Lepidoptera, Vol. 5. London. Lempke, B.J. ( 1 9 7 6 ) Naamlijst van de Nederlandse Lepidoptera. Uitgave Bibliotheek Koninklijke Nederlandse Natuurhistorische Vereniging, 2 1 , 1-100.
McDunnough, J . ( 1 9 3 9 ) Checklist of the Lepidoptera of Canada and the United States of America. 11. Microlepidoptera. Memoirs of the Southern California Academy of Sciences, 2. 1 - 17 1 . Opler, P.K. ( 1 9 7 4 ) Biology, ecology and host specificity of microlepidoptera associated with Quercus agrifolia (Fagaceae). University o f California Pu hlications in Entomology, 1 5 . Packard, A S . ( 1 8 7 0 ) New o r little known Injurious Insects. Annual Report of the Massachusetts Board of Agriculture, 11. 2 3 7 - 2 3 8 .
Packard, A.S. ( 1 8 8 9 ) Guide to the S t u d y of Insects, 9th edn. New York. Prentice, R.M. e t al. ( 1 9 6 5 ) Forest Lepidoptera of Canada. Recorded by the Forest Insect Survey. Microlepidoptera, 4, 5 4 6 - 8 4 0 . Rindge, F.H. ( 1 9 4 7 ) Designation and distribution of types of Nepticula braunella Jones (Lepidoptera: Nepticulidae). Pan-Pacific En tomologist, 1 3 , 2 5. Schrank, F. von P. (1802) Fauna Boica, 2 , 4 1 2 pp. Nurnberg. Scohle, M.J. (1980a) A new species of Ectoedemia Busck from Botswana with observations on its imaginal skeletal anatomy (Lepidoptera: Nepticulidae). Annals of the Transvaal Museum. 32, 35-54.
Scoble, M.J. ( 1 9 8 0 b ) The genus Niepeltia Stra nd: ta xonomy, and comments o n structure and relationships (Lepidoptera: Nepticulidae). A nnafs of the Transvaal Museum, 32, 1 9 1 - 2 2 9 . Smith, J.B. ( 1 8 9 1 ) List o f the Lepidoptera o f Boreal American Entomological Society, America. Philadelphia. Stainton, H.T. ( 1 8 5 4 ) Insecta Britannica. Lepidoptera: Tineina. London. Tutt, J.W. ( 1 8 9 9 ) A Natural History of the British 1, Swan Sonnenschein, Lepidoptera, Vol. London. Walsingham, T. ( 1 907) Microlepidoptera of Tenerife. Proceedings of the Zoological Society o f L o n d o n , 1901, 9 1 0 - 1028. Wilkinson, C. ( 1 9 7 8 ) On the StigmellalNepticula Controversy. (Lepidoptera). Tijdschrift voor Entomologie, 1 2 1 , 1 3 - 2 2 . Wilkinson, C. ( 1 9 7 9 ) A taxonomic study of the micro-lepidopteran genera Microcalyptris Braun and Fomoria Beirne occurring in the United States of America (Lepidoptera: Nepticulidae). Tijdschrift voor Entomologie, 1 2 2 , 5 9 - 9 0 . Wilkinson, C. ( 1 9 8 1 ) A supplement t o the genus Ectoedemia Busck (Nepticulidae: Lepidoptera) in North America, dealing with some difficult species and also some ne w ones. TijdXhri.fr voor Entomologie, 1 2 4 , 9 3 - 1 1 0 . Wilkinson, C. & Newton, P.J. ( 1 9 8 1 ) The microlepidopteran genus Ectoedemia Busck (Nepticulidae) in North America, Tijdschrift voor En tomologie, 1 24, 2 I -9 2 . Wilkinson, C. & Scoble, M.J. ( 1 9 7 9 ) The Nepticulidae (Lepidoptera) of Canada. Memoirs of the Entomological Society of Canada, 101, 1-129.
Accepted 2 4 November 1981
Appendix: Guide to the host plants of North American Stigmella AInus A.mgasa var. americana
corylifoliella
Amelanchier A . alnifolirIIa Amelanchier sp.
taeniola amelanchierella
Senila B.alba B.lutea (= alleghaniensis)
corylifoliella corylifoliella
Carpinus sp.
ostryaefoliella
Carya C.ova ta Carya sp.
ostryaefoliella ostryaefoliella
A taxonomic revision of Stigmella Castanea C.dentata Castanea sp.
castaneaefoliella ? latifasciella
Ceanothus C.arboreus C.divaricatus Ceanothus sp. Hybrid Ceanothus
inconspicuella ceanothi diffasciae ? ceanothi
Celtis C.occidentalis
? resplendensella
Condalia C.ferrea
condaliafoliella
Corylus C.americana Corylus sp.
corylifoliella corylifoliella
Cra taegus C.mollis Crataegus sp.
crataegifoliella, scintillans crataegifoliella. scintillans, pomivorella, stigmaciella, ? taeniola
Gaylussacia G.baccata
corylifoliella
Gossypium G. barbadense G.hirsu turn Gossypium sp.
gossypii gossypii ? Specimen 3
Hamamelis H . virginiana
corylifoliella
Heteromeles (= Photinia) H.arbu tifolia Hereromeles sp.
he revomelis heteromelis
Juglans J.californica J.nigra Juglans sp .
longisacca juglandifoliella juglandifoliella, longisacca
Malus sp .
pomivorella, scinanella, ? slingerlandella
Myrica M.cerifera Myrica sp.
myricafoliella ostryaefoliella
Opulaster sp .
corylifoliella
Ostrya C. virginiana Ostrya sp.
procrastinella ostryaefoliella
oxycoccus
0.palustris
corylifoliella
Physocarpus P. opulifolius
corylifoliella
Populus P. X canadensis P.canescens P. del to ides P. tric hocarpa Populus sp.
populetorum. aromella populetorum populetorum, aromella populetorum prunifoliella
Prunus P.ilicifolia and var. integrifolia P. nigra P.pensylvanica P.sero tina Prunus sp.
braunella prunifoliella, slingerlandella prunifoliella, slingerlandella prunifoliella, slingerlandella ? nixriverticella
Pyrus P.cornmunis
chalybeia
Quercus Q.agrifo1ia Q.alba Q.coccinea
vaviella saginella, sclerostylota, ? latifasciella ? Specimen e
46 1
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Philip J. Newton and Christopher Wilkinson
Q.imbricaria Q. kelloggii Q. macrocarpa Q.mari1andica Q.palustris Q.pla tanoides Q.pritius Qrubra Tan Bark Oak’ Q,wislizenii Quercus sp .
flavipedella, altella variella saginella, latifasciella quercipulchella latifasciella. flavipedella. quercipulchella, altella saginella. f l a vip edella saginella saginella, lutifasciella, flavipedella, quercipulchella, ? unifasciella Specimen f variella Specimen 4 , ? unifasciella
Rhamnus R.culifornica R.caroliniana R. lanceolata
Specimen 2 Specimen I rhamnicola
Rhus R.aromatica R . toxicodendron R. typhina
intermedia rhoifoliella intermedia
Rosa R.setigera R. virginiana Rosa sp.
rosaefoliella rosaefoliella rosaefoliella rosaefoliella rosaefoliella pectocatena, Specimen a, Specimeri b
Rubus R , occiden talis Rubus sp.
villosella villosellu
Salix S.disc0lor S.nigra Salix sp.
fuscotibiella firsco tibiella pallidu
Shepherdia S.canadensis
Specimen c
Tilia T.am ericana
tiliella, argen tifasciella
Ulmus U.am ericana U.fulva U.thomasi (= racemosa)
apicialbella apicialbella ? apicialbella
Vaccinium s p .
corylifoliella
Index to species-group names
(Synonyms are in italics)
Page
alba Wilkinson & Scoble altella Braun amelanchierella Clemens apicialbella Chambers argentifasciella Braun aromella Wilkinson & Scoble belfrageella Chambers bifasciella Clemens braunella Jones caryaefoliella Clemens castaneaefoliella Chambers ceanothi Braun cerea Braun chalybeia Braun ciliaefuscella Chambers
45 1 448 458 41 3 453 417 456 385 45 1 433 425 3 87 407 3 78 41 5
Page condaliafoliella Busck corylifoliella Clemens crataegifoliella Clemens dallasiana Frey & Boll diffasciae Braun discolorella Braun exasperata Braun flavipedella Braun fuscocapitella Chambers fuscotibiella Clemens gossypii Forbes & Leonard heteromelis s p a . inconspicuella s p n . intermedia Braun juglandifoliella Clemens
441 432 376 41 1 398 415 432 426 420 415 404 405 400 390 43 5
A taxonomic revision o f Stigmella
latifasciella Chambers in Hayden leucostigma Braun longisacca sp.n. macrocarpae Freeman maculosella Chambers minimella Chambers myricafoliella Busck nigriverticella Chambers obscurella Braun opulifoliella Braun ostry aefoliella Clemens pallida Braun puludicolu Braun pectocatena Wilkinson & Scoble plumosetaeella sp.n. pomivorella Packard populetorum Frey & Boll procrastinella Braun prunifoliella Clemens purpuratella Braun
31
Page 422 413 436 422 423 432 434 423 433 432 433 418 432 409 455 378 416 45 1 385 381
quercicastanella Chambers quercipulchella Chambers in Hayden resplendensella Chambers rhamnicola Braun rhoifoliella Braun rosaefoliella Clemens saginella Clemens scinanella Wilkinson & Scoble scintillans Braun sclerostylota sp.n. serotinaeella Chambers slingerlandella Kearfott stigmaciella Wilkinson & Scoble taeniola Braun term in ella Brau n tiliella Braun unifasciella Chambers variella Braun villosella Clemens virginiella Clemens
463 Page 420 445 456 393 391 408 420 380 377 429 385 409 382 382 445 442 438 447 410 432
