Research Article |
Corresponding author: Erik J. van Nieukerken ( nieukerken@naturalis.nl ) Academic editor: Donald Lafontaine
© 2020 Erik J. van Nieukerken, Charles S. Eiseman.
This is an open access article distributed under the terms of the Creative Commons Attribution License (CC BY 4.0), which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited.
Citation:
van Nieukerken EJ, Eiseman CS (2020) Splitting the leafmining shield-bearer moth genus Antispila Hübner (Lepidoptera, Heliozelidae): North American species with reduced venation placed in Aspilanta new genus, with a review of heliozelid morphology. ZooKeys 957: 105-161. https://doi.org/10.3897/zookeys.957.53908
|
The new genus Aspilanta gen. n. is described to harbour Nearctic heliozelid moths with reduced venation, previously placed in Antispila Hübner, 1825, with type species Antispila oinophylla van Nieukerken & Wagner, 2012. The erection of this genus has become possible now that monophyly has been supported by a recent phylotranscriptomics analysis. Six species are combined in this genus: Aspilanta oinophylla (van Nieukerken & Wagner, 2012), comb. n., A. hydrangaeella (Chambers, 1874), comb. n., A. ampelopsifoliella (Chambers, 1874), comb. n., A. voraginella (Braun, 1927), comb. n., A. argentifera (Braun, 1927), comb. n., A. viticordifoliella (Clemens, 1860), comb. n. and two candidate species are recognised. DNA barcode COI sequences of Malaise trapped specimens suggest a rich fauna of Aspilanta in Central America. All are leafminers, with Vitaceae as main host family, and single species feeding respectively on Hydrangeaceae and Myricaceae. The species are briefly diagnosed, and data on biology, DNA barcodes and distribution are provided. To place the genus in context, a review of heliozelid morphology and phylogeny is presented and a key to Nearctic genera is given. The genus is confined to North and Central America, possibly also occurring in South America. Aspilanta oinophylla is also an invasive species on grapevine in Italy. The genus is sister to Coptodisca Walsingham, 1895. Another species is removed from Antispila: Heliozela eugeniella (Busck, 1900), comb. n., feeding on Eugenia (Myrtaceae), from Florida.
Canada, DNA barcodes, grapevine pest, Heliozela, phylogeny, United States, Vitaceae
The taxonomy of the Lepidoptera family Heliozelidae, the shield bearer moths, has recently received considerable attention due to the discovery of several previously unknown pest species, attacking grapevines and walnuts (van
It soon became apparent that the relatively large genus Antispila Hübner, 1825, was a catch-all genus for species similarly patterned with a fascia before the middle of the forewing and a pair of opposing spots (the meaning of the Greek anti-spila / ἀντί-σπίλα) in the distal half. Several African and Asian species in Antispila with a reduced venation were transferred to the previously monotypic genus Holocacista Walsingham & Durrant, 1909 (van
To place this description in a wider context, we review the taxonomic history of the genus Antispila sensu lato and review knowledge on the morphology and phylogeny of the family Heliozelidae, partly based on the unpublished thesis of the late Ebbe
In an attempt to divide the large genus Tinea into smaller entities, at the time used for almost all small microlepidoptera,
The first to restrict the genus to heliozelid species was
In North America, Clemens discovered Antispila leafmines on “Nyssa multiflora” (black gum, now known as Nyssa sylvatica, Nyssaceae), but at first did not know where the reared moths belonged and suggested in a letter to Stainton on 15 May 1859 the new name Diacopia. This name has unintentionally, but validly been published as Diacopia Clemens, 1872, when Stainton published their correspondence much later (
A formal selection of Antispila stadtmuellerella as the type species was made by
Until recently, the validity of the genus Antispila was rarely challenged; only the small Mediterranean species A. rivillei (Stainton, 1855) was moved to the genus Holocacista, when the different venation was noted by
The presence of species with a reduced venation was already noted by
When describing the new Antispila oinophylla¸ the non-monophyly of North American Antispila became apparent once again (van
As most species treated here were already discussed in the paper describing Antispila oinophylla, we re-use several photographs from van
All examined and observed material, including parasitoids and all available barcoding data are listed in the occurrence dataset, published with GBIF (DOI: https://www.doi.org/10.15468/79mu7z). In the text we omit all material that was already listed by van
Under the heading “Distribution” we list countries, states and provinces; an asterisk denotes a new record.
Leafmines were photographed by CSE in the field using a Nikon D50 digital camera and a Nikon AF Micro Nikkor 105 mm lens. The camera’s built-in flash was used for photographs with reflected light; the sun was used for transmitted light. All other photographs by CSE were taken using a Canon EOS Rebel XSi SLR digital camera, MP–E 65 mm macro lens, and Macro Twin Lite MT–24EX flash unit. For transmitted light in leaf mine photographs, one of the flash heads was removed and aimed at the camera while the leaf was held tightly against it.
Leaves containing larvae were placed in sealed, 9– or 15–dram plastic vials, which were stored away from direct sunlight and checked daily (when possible) for emerging insects. After pupal cases were formed, extraneous leaf material was removed from the vial to minimise mould and a moistened, crumpled piece of toilet paper was added to prevent desiccation. Overwintering pupal cases were stored in a refrigerator at 1–3 °C from 6 November 2013 to 25 February 2014, 1 November 2014 to 1 March 2015, 1 November 2015 to 1 March 2016, 20 October 2017 to 1 April 2018, and 17 October 2019 to 21 February 2020. Adult moths were chilled and photographed as described by
Morphological terminology follows van
Host plant names follow the Catalogue of Life (
We list parasitoids that we reared from Aspilanta larvae. Those reared by EvN were routinely stored in ethanol 96–100%, and later DNA barcoding was performed (dataset DS-PARASP, doi: https://www.doi.org/10.5883/DS-PARASP). Chalcidoidea were morphologically identified by Sandrine Ulenberg, Braconidae were tentatively identified by EvN on the basis of DNA barcodes; all are stored in
Abbreviations for depositories:
CSEC Personal collection of C.S. Eiseman.
Antispila: auct. partim, nec Hübner, 1825.
Antispila ampelopsifoliella group; van
Antispila
“group II”;
Antispila oinophylla van Nieukerken & Wagner, 2012: 38, by present designation.
Very small moths, wingspan between 4.0 and 6.2 mm, with a forewing pattern of metallic–silvery markings, comprising an oblique fascia at ¼, a postmedial pair of spots (one costal and one dorsal), and usually a small apical spot (only absent in A. viticordifoliella); fringe line more or less distinct. Males never with androconial scales or hair-pencils. Antennae with only 16–20 segments. Antispila species in North America never have an apical spot. Aspilanta species are diagnosed by the reduced venation (Figs
Adults (Figs
Head (Figs
Thorax. Vestiture of appressed lamellar scales, either concolourous with ground colour of forewings or more metallic and similar to vestiture of head. Foreleg (Fig.
Wings. Male retinaculum a series of 7–12 hook-shaped bristles (Fig.
Venation (Figs
Wing pattern (Figs
Pregenital abdomen. Abdominal sclerites weakly sclerotised. Anterior sternum II subtriangular, free.
Male genitalia (Figs
Female genitalia (Figs
Larva (Figs
Host plants. Most species feed on Vitaceae, one each on Hydrangeaceae and Myricaceae.
Eggs are inserted in leaf tissue, often near a vein or leaf margin. All species construct leafmines, either starting as a narrow linear mine and later widening into a blotch, or sometimes starting almost immediately as a blotch mine. All frass is deposited in the mine, initially filling or irregularly scattered in the linear portion, later often scattered in the blotch or pushed by the larva to one side. During the penultimate (fourth) instar an oval section is cut out from both epidermal layers, forming a portable case or “shield”. This shield (Figs
North America; DNA barcodes suggest a rich fauna in Central America: Mexico, Honduras, Costa Rica, likely also elsewhere in the Neotropics (see under Composition).
The name Aspilanta is an anagram of Antispila, where one “i” was replaced by an “a”. The gender of the name is to be regarded as feminine.
In the checklist below we provide the original genus in brackets, type locality, and the hostplant of the types. The species are listed according to the position in the recent phylogenetic analyses (
We also include the candidate species Aspilanta “Vitis1_USA” (van
The Neotropic species Antispila trypherantis Meyrick, 1916 (Guyana), A. pentalitha Meyrick, 1916 (Guyana) and A. cyclosema Meyrick, 1921 (Brazil) may also belong in Aspilanta, based on their original descriptions that cite the presence of an apical dot, but without examination of types, we refrain from recombination here. Also the Patagonian group of species, associated with Nothofagus (Nothofagaceae), for which Nielsen in his unpublished thesis proposed the name “Neospila”, could belong in Aspilanta on the basis of the very similar externals (Fig.
In the Museum of Comparative Zoology (Cambridge, MA), there is a series of externally similar moths of unknown provenance, ex coll. Dietz, allegedly reared from poison ivy, (Toxicodendron radicans, Anacardiaceae) with the manuscript name “Antispila rhoifoliella” (handwritten label: on Rhus radicans. Coll 9.7.[18]99, many mines empty, larva pale, green frass line; head + 1 dark brown. [word crossed out] mine begins with a fine tract along edge of leaf, expands & frass collects along edge of mine.). These specimens have emergence dates of 1–20 June 1900 (written as “19C”). As we have never seen such mines on poison ivy we cannot exclude the possibility that Virginia creeper was mistaken for poison ivy, as both often grow together; in this case the series would likely represent Aspilanta ampelopsifoliella. We thus ignore this information until these specimens have been examined in more detail.
1 | Forewing colour pattern comprising silvery white spots and bands on a dark background | 2 |
– | Forewing basally uniform silvery to leaden grey (exceptionally with paler longitudinal streaks), distal part comprising a pattern of a yellow to orange background, with three silver strigulae and various amounts of black | Coptodisca |
2 | Forewing with an apical silver spot in addition to a fascia at ¼ and two opposite spots at 2/3 | Aspilanta (most species) |
– | Forewing without apical spot | 3 |
3 | Forewing with a pattern of a fascia at ¼ and two opposite spots at 2/3 | 4 |
– | Forewing with only two spots on dorsum, or a spot at ¼ and a narrow fascia at 2/3 | Heliozela |
4 | Antenna with white tip | Aspilanta viticordifoliella [note: there may be an Antispila species with a similar white tip (see under A. viticordifoliella). In that case checking wing venation is necessary for the correct identification] |
– | Antenna without white tip | Antispila |
Aspilanta oinophylla (van Nieukerken & Wagner, 2012): 38, comb n. (Antispila) USA, Georgia, Murray Co., Vitis aestivalis var. aestivalis [type species]
Aspilanta hydrangaeella (Chambers, 1874a): 170, comb n. (Antispila) USA, Kentucky, Covington, Hydrangea arborescens L.
Aspilanta ampelopsifoliella (Chambers, 1874a): 168, comb n. (Antispila) USA, Kentucky, Covington, Parthenocissus quinquefolia (L.) Planchon
Aspilanta voraginella (Braun, 1927b): 191, comb n. (Antispila) USA, Utah, Washington Co., Vitis arizonica Engelm.
Aspilanta argentifera (Braun, 1927a): 56, comb n. (Antispila) Canada, Ontario, Sparrow Lake., collected as adult without definite host association.
Aspilanta viticordifoliella (Clemens, 1860): 209, comb n. (Antispila) USA, Pennsylvania, Easton, Vitis vulpina L.
Aspilanta “Vitis1_USA” van Nieukerken & Wagner, 2012 (in Antispila) USA, Florida, Connecticut, Vitis aestivalis Michx.
Aspilanta “Vitis.arizonica_USA” USA, Arizona, Vitis arizonica Engelm.
In order to judge the validity of the morphological characters in Aspilanta, they should be compared with the same characters in other Heliozelidae. There is no single large treatment of the family, but
Adult. Head. All Heliozelidae share the smooth head, with appressed, lamellar scales. This is a strong apomorphy for the family. Antennae in Hoplophanes have ca. 28–30 flagellomeres, Heliozela 15–25, ca. 23 in Antispila, 18 in Antispilina, 12–18 in Holocacista, 12–16 in Coptodisca and 14–18 in Aspilanta. The flagellomeres are much longer than wide in Aspilanta and other genera in the Holocacista group, almost the same in Antispila, but in Heliozela hardly longer than wide; all flagellomeres have two rings of scales. Mandibles are small, but visible in all genera (according to Nielsen absent in the leafmining genera). The proboscis (galea) is more than twice the height of the head in Heliozela and Antispila, almost twice the height in Aspilanta, but hardly longer than the head in Holocacista, Antispilina and Coptodisca. The maxillary palpi are progressively reduced, from four segments in Hoplophanes and Heliozela, through three segments in Antispila and a reduction in the Holocacista group genera into a single segment (Nielsen considered them absent, but we clearly observed a short palp). Labial palpi usually have three segments, but only two in Antispilina ludwigi; the last segment is very long in Aspilanta (3 × second segment) compared to Coptodisca (slightly longer than second segment). Eyes are rather similar in most Heliozelidae, but comparatively large in Hoplophanes (interocular index 1.06, see (
Thorax. The most relevant character in the legs is the presence of an epiphysis on the foretibia, a plesiomorphic condition in Lepidoptera. It is present in Hoplophanes, Tyriozela, and Heliozela, probably also in the other Australian genera, but absent in Antispila and most of the genera in the Holocacista group, but Holocacista rivillei and H. capensis have a reduced epiphysis (van
Forewings in the majority of Heliozelidae are metallic shining. A forewing pattern of spots and fasciae is present in many Heliozelidae, in Antispila, Antispilina, Holocacista and Aspilanta comprising a fascia (or two spots) at one third and an opposite pair of spots at two thirds, somewhat reduced in Ischnocanaba. An apical dot is characteristic for most Aspilanta species, but something similar also occurs in Ischnocanaba and Nielsen’s “Neospila”. Larger terminal spots are known in some Antispila species. Most Heliozela species have the silver spots confined to the dorsal margin, but in some cases one of these reaches the costa, or an extra spot is present on the costa, and a few have a pattern similar to Antispila. Such species were previously incorrectly placed in Antispila (e.g., H. anna (Fletcher, 1920)), like many of the species in Holocacista and Aspilanta. Coptodisca species have a completely different pattern, resembling that of Leucoptera (Lyonetiidae).
Androconial scales on one or both wings are common in Antispila, but absent from most other genera.
Venation. Hoplophanes, Heliozela, and Antispila have the more complete venation, with a closed cell between Rs and CuA, three branches of Rs; in Antispilina, Holocacista, Aspilanta and Coptodisca the venation is reduced, Rs, M and Cu are coalescent from wing base, closing the cell, and ending in three or four branches, rarely two. The label CuA given for the A. oinophylla forewing by van
Male genitalia. These are rather uniform throughout Heliozelidae, with a long and narrow vinculum; bilobed or truncate tegumen (sometimes termed uncus); triangular or elongate valvae, each with a single pectinifer with 8–30 blunt sensilla, often different numbers on each valva; a spear-shaped juxta; phallus very long and narrow, with anellar spines and often large projections at phallotrema. Whereas the male genitalia are diagnostic at species level, few characters can be used at higher phylogenetic level. The recurved appendix of the phallus is probably characteristic for Holocacista, whereas the very long vinculum (more than twice the valva length) separates most species of the Holocacista group from Antispila.
Female genitalia. The oviscapt is spear shaped with many small teeth in Hoplophanes, Tyriozela and Heliozela, whereas in Antispila and the Holocacista group it has just a few cusps at either side. In Antispila there is a distinct interapodemal process between the anterior apophyses, running almost to the caudal tip. In the Holocacista group the process is very short, the caudal part being absent.
Larva. Larvae of the Australian genera are unfortunately hardly known. Leafmining larvae of Heliozela sericiella (
Head. Fourth instar larvae of Heliozela have two pairs of stemmata at either side, those of Antispila three, and Aspilanta two. In Holocacista and Coptodisca probably also two, but difficult to see in our slide mounted specimens. Davis (1978,
Thorax. Legs are absent in most studied larvae, but some Heliozela species have much reduced legs during the 5th, non-feeding instar (
Abdomen. Prolegs are only reported from 4th instar Heliozela larvae on segments 3–6, with 3–8 crochets in single or sometimes double mesoseries. Instead most larvae show either paired or unpaired calli, the position of which may be diagnostic, but is difficult to assess in slide mounted specimens. The integument is covered by slight, often transverse swellings, which may concentrate in the centre of the segments, and in some cases form a row of abdominal plates in species of Antispila. The presence of warts on the 8th larval segment, used for stridulation or drumming (
Cocoon. Cocoons (shields) of leafmining species are prepared from both epidermal layers of the final part of the leafmine, held together by silk, and are very similar in most genera. Antispila has as extra character in that the larva contracts the case with silk, resulting in a lengthwise keel; also in all Antispila species there are silken projections from the anterior (and sometimes the posterior) end of the case; we do not know them from Heliozela, Holocacista or Coptodisca, but some Aspilanta species produce these projections at least sometimes (Fig.
Pupa. The pupa of Heliozela sericiella was described by
Phylogeny. We use the recent molecular phylogeny based on transcriptome data, including up to 1049 nuclear genes (
Possible apomorphies for the Heliozela, Antispila, and Holocacista groups are:
(1) Size strongly reduced. The wingspan of all the species of these genera is small, always less than 9 mm, and usually smaller.
(2) Compound eyes in a posteroventral position. The anterior lateral margin of the compound eyes does not reach the anterior surface of the head capsule, but is present behind the laterally extended genae.
(3) Proboscis very long, two to three times as long as height of head capsule. However, as noted above, it is shorter again in the small moths of Holocacista, Antispilina, and Coptodisca.
Nielsen also listed absence of mandibles as an apomorphy, but we think that reduced mandibles are in fact present.
A possible apomorphy for the Heliozela group (including Tyriozela) is:
(4) Female with spear-shaped ovipositor tip. The flattened, deeply dented ovipositor observed in the Antispila and Holocacista groups is regarded as the relatively plesiomorphic state compared to the spear-shaped tip of the Heliozela group. The first mentioned type is present in most Incurvariidae and the most generalised genera of Prodoxidae.
Two other characters given by Nielsen are in our view incorrect; the retinaculum is not as long as Sc as he states, and although some species have petiole-mining larvae, this is not the only larval feeding mode; many Australian and Oriental species are leafminers, and others are gall formers.
Possible apomorphies for Antispila and the Holocacista group are:
(5) Head with transfrontal suture absent and scale-sockets on the entire frons and vertex. The area in all other Adeloidea is strengthened and devoid of scale-sockets.
(6) Foretibia without fully developed epiphysis.
(7) Pseudofrenular bristles absent in male.
(8) Maxillary palpi three-segmented or shorter.
(9) Microtrichiation reduced on forewings. Only the proximal third is provided with microtrichia; in Hoplophanes the entire wing surface is covered with microtrichia, what is also regarded as a ground-plan condition within Adeloidea.
(10)*. Larvae without legs and prolegs.
(11)*. Forewing with “Antispila pattern” (see above), secondarily derived in Coptodisca and a few species of Holocacista.
Possible apomorphies for Antispila s. str. are:
(12)* Cocoon folded lengthwise, resulting in a distinct keel.
(13)* The presence of warts on the 8th larval segment may be an additional apomorphy, but more likely only for a subset of the genus.
Nielsen gave the presence of an interapodemal process in female as important apomorphy for the Antispila group, but we see a similar structure, albeit somewhat reduced, in Holocacista (van
Possible apomorphies for the Holocacista group are:
(14). Venation strongly reduced: cell absent in both fore- and hindwings.
(15). Male retinaculum comprising a series of hook-shaped setae.
(16). Sublateral process of the transtilla in male genitalia long and slender.
(17). Larval case attached freely to a branch of the food-plant or other material.
(18)*. Male genitalia: vinculum extremely extended.
(19)*. Number of antennal segments reduced.
(20)*. Reduction of maxillary palps to one segment.
Possible apomorphies for Antispilina are:
(21). Larvae mining the leaves of herbaceous Polygonaceae.
(22)*. Labial palpi reduced to 2 segments.
A possible apomorphy for Holocacista is:
(23)*. Phallus with long, often recurved appendix.
Possible apomorphies for Aspilanta are:
(24)*. Forewing with apical spot.
Possible apomorphies for Coptodisca are:
(25). Forewing pattern specialised.
DNA barcoding. We provide COI DNA barcode sequences as an aid for identification for all known species of Aspilanta. A Neighbor-Joining tree of these (Fig.
1 | Forewing with an apical silver spot in addition to a fascia at ¼ and two opposite spots at 2/3; antenna dark throughout or with the last 1–3 segments white | 2 |
– | Forewing without apical spot; antenna with distinct white tip of 2 or 3 segments | A. viticordifoliella |
2 | Antennae with last 3 segments clearly white. Larvae mine Hydrangea | A. hydrangaeella |
– | Antennae dark throughout, or at most last segment white. Larvae mine Vitaceae or Myricaceae | 3 |
3 | Head with silvery white scales, forewing with distinct fringe line, fringe almost white. Larvae on Vitaceae |
A. oinophylla, ampelopsifoliella or “Vitis1” See van |
– | Head with darker bronze brown or brassy scales, forewing with rather indistinct fringe line, fringe grey brown. Larvae on Vitaceae or Myricaceae | 4 |
4 | Eastern species (ON & NL to NC), larvae on Myricaceae | A. argentifera |
– | Southwestern species (AZ, UT, TX), larvae on Vitis | A. voraginella |
Antispila oinophylla
van Nieukerken & Wagner, 2012: 38. Holotype ♂, USA: Georgia, Murray Co., Chattahoochee Nat. Forest, E of Chatsworth, GA rd 52, 523 m, 34.74066N, 84.71852W, hardwood forest along highway, leafmines on Vitis aestivalis var. aestivalis, 14.x.2010, EvN2010266, emerged 14.iv–4.v.2011, E.J. van Nieukerken & C. Doorenweerd, Genitalia slide EJvN 4204,
Antispila oinophylla;
[Antispila ampelopsifoliella;
[Antispila ampelopsiella;
Wingspan ca. 4.8–6.2 mm, forewing length 2.3–2.8 mm. Externally inseparable from A. ampelopsifoliella and A. “Vitis1_USA”. The silvery white head separates it from A. voraginella and A. argentifera, and A. hydrangaeella has distinct white antennal tips. Male genitalia with characteristic long curved process and a comb of 10–12 large teeth at phallotrema; tegumen truncate; valvae with 10–12 pecten sensilla. Female oviscapt with more cusps (4–5 at either side) than A. ampelopsifoliella (3). Leafmines differ from other Vitaceae miners by compact size, rather short linear portion close to a vein and frass in concentric lines, especially in thin leaves.
Aspilanta species, mounted adult moths. 1 A. oinophylla, male, USA, VT,
Vitaceae: Parthenocissus quinquefolia, P. vitacea, Vitis aestivalis, V. labrusca, V. riparia, V. vinifera, V. vulpina.
(Figs
Yellowish green, with green gut contents; head and prothorax brown.
Larvae in Canada and NE United States in July, August and September, more south until October; in Italy in June–July and August–October. Adults from June to August; apparently bivoltine.
Canada: Ontario, Quebec; USA: Connecticut, Georgia, Kentucky, Massachusetts*, Minnesota*, New York, North Carolina*, Oklahoma*, Tennessee, Vermont, Wisconsin*; Europe: Italy (introduced).
BIN: BOLD:AAI4367 average distance 0.42%, max. distance 1.28% (n = 41), distance to nearest neighbour 9.79% (A. voraginella).
Eulophidae: Cirrospilus pictus (Nees, 1834) (Italy) (BOLD:ACZ7659), Cirrospilus sensu lato, including Burkseus (see
In the original description it was noted that the species had not been found on Parthenocissus in North America. Since then some specimens of this species were barcoded and reared from Parthenocissus by J.-F. Landry (material in
Adults examined. CANADA – Ontario • 2; Normandale; 42.71N, 80.31W; Freeman & Lewis leg.; Vitis; emerged 22–25 Mar. 1957; EventId: 56–168; CNCLEP00122325–00122326. • 2; Normandale; 42.71N, 80.31W; Freeman & Lewis leg.; Vitis; emerged 19–25 Feb. 1960; EventId: 59–198; CNCLEP00122327–00122328. • 1 ♂; Ottawa; 45.41N, 75.69W; G.G.Lewis leg.; Vitis; emerged 25 Feb. 1971; EventId: 70–53; Genitalia slide: MIC1871; CNCLEP00100104. • 1 ♀ 1; Ottawa; 45.41N, 75.69W; G.G.Lewis leg.; Parthenocissus; emerged 15, 29 Mar. 1971; EventId: 70–48; Genitalia slide: MIC1877; CNCLEP00100105–00100106. • 1 ♂1 ♀ 3; Ottawa; 45.41N, 75.69W; Freeman, Lewis leg.; Vitis; emerged 21 Mar. – 08 Apr. 1958; EventId: 57–112; Genitalia slide: MIC1872, MIC1873; CNCLEP00100467–00100468, 00122312–00122314. • 1 ♂ 1; Ottawa; 45.41N, 75.69W; T.N. Freeman leg.; Vitis; emerged 16–18 Apr. 1957; EventId: 56–183; Genitalia slide: MIC1875; CNCLEP00122315, 00122316. • 1 ♂ 6; Overbrook; 45.42N, 75.65W; G.G. Lewis leg.; Vitis; emerged 15–24 Aug. 1955, 04–06 Jul. 1956; EventId: 55–53; Genitalia slide: MIC1874, MIC1878; CNCLEP00122317 – 00122324. – Québec • 1; Gatineau, Aylmer, 48 rue du Couvent; 45.3967N, 75.849W; alt. 80 m; 14 Jul. 2010; J.-F. Landry leg.; Vitis riparia; CNCLEP00097660. • 3; same data as preceding; emerged 31 Aug. – 02 Sep. 2011; CNCLEP00091691–00091693. • 4; same data as preceding; Parthenocissus quinquefolia; emerged 01–02 Sep. 2011; CNCLEP00091694– 00091697. • 10; same data as preceding; alt. 80 m; J.-F. Landry leg.; Vitis riparia; emerged 31 Aug. 2011; CNCLEP00097700–00097709. • 1; Hull; 45.435N, 75.708W; T.N. Freeman leg.; Vitis; emerged 07 Jul. 1956; EventId: 55–228; CNCLEP00122305. • 3; Québec, Hull; 45.435N, 75.708W; T.N. Freeman leg.; Vitis; emerged 01–07 Jul. 1956; EventId: 55–228; CNCLEP00122306–00122308. • 2; Hull; 45.435N, 75.708W; G.G. Lewis leg.; Vitis; emerged 12–15 Aug. 1955; EventId: 55–60; CNCLEP00122309–00122310. • 1; Québec, Hull; 45.435N, 75.708W; G.G. Lewis leg.; Vitis; emerged 30 Jun. 1956; EventId: 55–60; CNCLEP00122311.
USA – Oklahoma • 1; Payne Co., Mehan; 36.014339N, 96.996744W; 10 Jul. 2016; Michael W. Palmer leg.; Vitis; emerged 10 Aug. 2016; EventId: CSE2971; CSEC. – Vermont • 1 ♂, 9; Addison Co, Button Bay SP, Lake Champlain borders; 44.18154N, 73.36892W; alt. 40–50 m; 16 Sep. 2011; E.J. van Nieukerken leg.; Vitis riparia; emerged 21 May–05 Jun. 2012; EventId: EvN no 2011253–1K; Genitalia slide: EvN4378;
CANADA – Ontario • leafmines and larvae, rearing failed; Chatham–Kent Div., Rondeau Prov. Park, Campground; 42.3223N, 81.8438W; alt. 177 m; 24–25 Jul. 2015; E.J. van Nieukerken leg.; Vitis riparia; EventId: EvN no 2015091–1K;
USA – Massachusetts • Franklin Co., Northfield; 42.646762N, 72.42527W; 13 Sep. 2016; Charley Eiseman leg.; Vitis aestivalis; CSEC.
CANADA – Ontario • 3; Grand Bend, Pinery Provincial Park, Site 2; 43.2699N, 81.8271W; alt. 178 m; 25 Jun.–09 Jul. 2014; CBG Collections Staff leg.; EventId: GMP#03351; BIOUG33534–B01, BIOUG33534–C01, BIOUG33534–D01. • 1; Point Pelee National Park, Cactus Field; 41.939N, 82.516W; alt. 168 m; 27 Jun.–04 Jul. 2012; Tyler Peters leg.; EventId: GMP#00175; BIOUG03514–B07. • 1; Wellington County, Puslinch Township, Concession 11/Hume Rd; 43.537N, 80.134W; alt. 320 m; 10–17 Jul. 2010; Paul Hebert leg.; EventId: L#PHPUS–016; BIOUG02834–E04. • 1; Lambton Co., Port Franks; 43.22N, 81.91W; 02 Jul. 2017; K. H. Stead leg.; EventId: KSLEP1081–17; KSLEP1081–17; Research Collection of Ken Stead. • 1; Lambton Co., Port Franks; 43.2257N, 81.916W; alt. 188 m; 30 Jul. 2012; K.H.Stead leg.; BIOUG20646–E04. • 1; same locality data as preceding; 02 Aug. 2012; K.H.Stead leg.; BIOUG16764–D05. – Québec • 1; Montreal, Montreal Botanical Garden, Trap 2; 45.5594N, 73.5668W; alt. 52 m; 11–18 Jun. 2014; Maxim Larrivee leg.; EventId: GMP#04699; BIOUG25652–B07.
ITALY – South Tyrol • 1; Bolzano, Kaltern/ Altenburger Wald, Umg. Ziegelstadel; 46.379N, 11.229E; alt. 705 m; 10 Aug. 2015; Huemer P. leg.;
ITALY (all from http://www.lepiforum.de/lepiwiki.pl?Antispila_Oinophylla) – South Tyrol • 1 adult; Bolzano, Bozen–Rentsch; 46.502N, 11.366E; alt. 265 m; 08 Aug. 2014; Werner Pichler leg.. • many adults; South Tyrol, Bolzano, Bozen–St. Magdalena; 46.503N, 11.372E; alt. 250–600 m; 01 Jul.–30 Aug. 2015; Werner Pichler leg.; Vitis vinifera. • Bolzano, Eisacktal, Klausen; 46.64N, 11.56E; alt. 650 m; 09–11 Jul. 2019; Dieter Robrecht leg.; Parthenocissus vitacea; Robrecht, Dieter, personal collection. • 15 adults; South Tyrol, Bolzano, Eisacktal, Klausen [Chiusa]; 46.64N, 11.56E; alt. 650 m; 09–11 Jul. 2019; Dieter Robrecht leg.; Vitis vinifera; emerged 01–06 Aug. 2019; Robrecht, Dieter, personal collection.
USA – Minnesota • Wabasha Co., Weaver Dunes Preserve; 44.258N, 91.932W; 17 Jul. 2015; Charley Eiseman & Julia Blyth leg.; Vitis; https://www.inaturalist.org/observations/44819628. – North Carolina • Durham Co., 17–acre wood preserve; 36.024N, 78.925W; 29 Sep. 2017; Tracy Feldman leg.; Vitis; EventId: CSE4561; https://bugguide.net/node/view/1447964. – Wisconsin • Dane Co., Cross Plains; 43.12N, 89.66W; 19 Sep. 2011; Ilona Loser leg.; Vitis riparia; https://bugguide.net/node/view/578758/bgimage.
Antispila hydrangaeella Chambers, 1874a: 170. Syntypes leafmines and larvae: [USA: Kentucky, Covington] on Hydrangea nivea [probably lost].
Antispila hydrangiaeella Chambers, 1878: 113. Subsequent incorrect spelling.
Antispila hydrangaeella;
Antispila hydrangiaeella;
Wingspan ca. 5.0–5.8 mm, forewing length 2.2–2.8 mm. Externally easily separable from other Aspilanta species with terminal spot by the white terminal antennal segments (3 flagellomeres with 6 scale rings). Male genitalia characterised by the two long curved terminal spines at phallotrema, the bearded setae on the juxta, the larger number of sensilla (ca 17–21) of the valval pecten and the bilobed tegumen. Female genitalia not examined.
Aspilanta oinophylla, adult (male) morphology,
Hydrangeaceae: Hydrangea arborescens.
(Figs
Colourless or whitish except for green gut contents; head and prothorax dark brown, some darker dots on most segments visible (Fig.
Poorly known; larvae found in June, August, September and October; adults emerged the following spring from larvae collected in October, but dates are too early due to forced emergence. Chambers provides no information whatsoever;
USA: Georgia, Illinois, Kentucky, Maryland*, North Carolina, Ohio*, Tennessee*.
Georgia and Ohio population: BIN: BOLD:AAV5059 (n = 2, average distance = 2.15%), Tennessee population: BOLD:AAV5060 (n = 1), distance between these nearest neighbours 5.25%.
Eulophidae: Pediobius albipes (Provancher, 1887) (GA) (BOLD:ACZ8030), P. ocellatus Peck, 1985 (GA, NC) (BOLD:ACZ8031); Braconidae: Microgastrinae sp. (GA) (BOLD:ADA0313).
In our barcode analysis we found two clusters with a distance of 5.25%, even the distance of two specimens of the Georgia population is rather large with 2.15%. By analysing the characters of adults of the two barcoded populations, we did not find any supporting argument for the earlier suggestion that two species might be involved (van
USA – Tennessee • 1; Blount Co., NP Great Smoky Mts, Lead Cove Trail; 35.59976N, 83.73998W; alt. 690 m; 03 Oct. 2010; E.J. van Nieukerken & C. Doorenweerd leg.; Hydrangea arborescens; emerged 04 Apr. 2011; EventId: EvN no 2010141–K;
USA – Ohio • 2 larvae, Hocking Co., South Bloomingville, Deep Woods Farm; 39.406165N, 82.574946W; alt. 229 m; 13 Sep. 2012; C. Eiseman leg.; Hydrangea arborescens; EventId: CSE-OH;
USA – Georgia • Gilmer Co., Ellijay; 34.651N, 84.608W; 23 Sep. 2019; Lisa Kimmerling leg.; Hydrangea arborescens; https://www.inaturalist.org/observations/33268524. – Kentucky • Bullitt Co., Bernheim Arboretum and Research Forest; 37.913N, 85.648W; 30 June 2019; Mike Plagens leg.; Hydrangea arborescens; https://www.inaturalist.org/observations/29522629. – Maryland • Harford Co., Susquehanna State Park; 39.605N, 76.152W; 11 Aug. 2018; Josh Emm leg.; Hydrangea arborescens; https://bugguide.net/node/view/1573751/bgimage. – North Carolina • Madison Co., Along Appalachian Trail southbound from Devil’s Fork Gap; 36.0082N, 82.6098W; 05 Oct. 2019; Jim Petranka leg.; Hydrangea arborescens; https://bugguide.net/node/view/1736188/bgimage. – Ohio • Hocking Co., Rockbridge, Crane Hollow Preserve; 39.48N, 82.584W; 05 Aug. 2016; Charley Eiseman & Julia Blyth leg.; Hydrangea arborescens; https://www.inaturalist.org/observations/44820974.
Antispila ampelopsifoliella
Chambers, 1874a: 168. Syntypes: leafmines [USA: Kentucky, Covington] on Ampelopsis quinquefolia [= Parthenocissus quinquefolia], “pseudotypes”, Kentucky, Covington (
Antispila ampelopsisella Chambers, 1874b: 197. Subsequent incorrect spelling.
Antispila ampelopsiella Chambers, 1874b: 198. Subsequent incorrect spelling.
Antispila ampelopsilofoliella: internet error.
Antispila ampelopsifoliella;
Antispila ampelopsiella;
Wingspan ca. 5.0–5.3 mm, forewing length 2.4–2.6 mm. Externally inseparable from A. oinophylla (see above) and A. “Vitis1_USA”. Male genitalia characterised by the non-bilobed tegumen, pecten with 11–14 sensilla, and phallus with rather small curved spine at phallotrema. Female oviscapt with 3 cusps at either side. Leafmines recognised by long linear portion that is often sinuous.
Vitaceae: Parthenocissus quinquefolia, P. vitacea.
(Figs
Aspilanta species, male genitalia. 21 A. oinophylla, phallus separate,
Colourless or whitish except for green gut contents, in contrast with yellow-green larva of A. viticordifoliella on the same host plant (compare Figs
Poorly known. Larvae found from June (Oklahoma), August and September, field caught adults only from early July. The adult from the June larva emerged the following spring, like those from August and September larvae, suggesting that this species is univoltine.
Canada: Ontario; USA: Connecticut, Kentucky, Massachusetts*, New York, Oklahoma*, Rhode Island*, Vermont, Wisconsin*.
BIN: BOLD:ABA3237, average distance 0.3%, max. distance 1.12% (n = 10), distance to nearest neighbour 8.79% (BOLD:ACO9420, an unidentified Heliozelidae from Mexico, most likely an Aspilanta species).
Eulophidae: Chrysocharis paradoxa Hansson, 1985 (MA), Closterocerus cinctipennis Ashmead, 1888 (MA); Braconidae: Microgastrinae sp. (MA), Pseudognaptodon sp. (VT).
The identity of this species follows from the discussion in van
CANADA – Ontario • 3 ♀; Normandale; 42.71N, 80.31W; Freeman, Lewis leg.; Parthenocissus; emerged 15–23 Mar. 1958; EventId: 57–157; Genitalia slide: MIC1865, MIC1869, MIC1867; CNCLEP00100465, 00122296, 00122301. • 1 ♂; Normandale; 42.71N, 80.31W; Freeman, Lewis leg.; Parthenocissus quinquefolia; emerged 13 Mar. 1958; EventId: 57–157; Genitalia slide: MIC1866; CNCLEP00100464. • 6; Normandale; 42.71N, 80.31W; Freeman & Lewis leg.; Parthenocissus (quinquefolia); emerged 14–19 Mar. 1958; EventId: 57–157; CNCLEP00122294, 00122295, 00122297–00122300. • 1 ♂; Overbrook; 45.42N, 75.65W; G.G. Lewis leg.; Parthenocissus vitacea; emerged 05 Jul. 1956; EventId: 55–57; CNCLEP00122409. • 1; Simcoe; 42.83N, 80.31W; T.N. Freeman leg.; emerged 16 Jan. 1960; EventId: 65–80; CNCLEP00100466.
USA – Connecticut • 1 ♂; Winham Co, Windham airport, Mansfield Hollow SP; 41.74783N, 72.16409W; alt. 80 m; 09 Sep. 2011; E.J. van Nieukerken leg.; Parthenocissus quinquefolia; emerged 24 Apr. 2012; EventId: EvN no 2011178–2K; Genitalia slide: EvN4377;
USA – Massachusetts • Franklin Co., Northfield; 42.646558N, 72.426541W; 16 Sep. 2016; Charley Eiseman leg.; Parthenocissus quinquefolia; CSEC. • Hampshire Co., Pelham; 42.363212N, 72.460107W; 30 Aug. 2013; Charley Eiseman leg.; Parthenocissus quinquefolia; EventId: CSE914; CSEC; • 1 larva; Nantucket Co., Nantucket, Squam Swamp; 41.319937N, 70.00244W; alt. 2 m; 07 Sep. 2011; C.S. Eiseman leg.; Parthenocissus quinquefolia; EventId: C4;
CANADA – Ontario • 1; St Williams, Turkey Point Provincial Park; 42.7052N, 80.3285W; alt. 222 m; 23 Jun. 2014–07 Jul. 2014; CBG Collections Staff leg.; EventId: GMP#03287; BIOUG35664–A01. • 1; Thorold, Short Hills Provincial Park; 43.1129N, 79.2738W; alt. 94 m; 23 Jun. 2014–07 Jul. 2014; CBG Collections Staff leg.; EventId: GMP#03255; BIOUG35571–A12.
USA – Massachusetts • Nantucket Co., Nantucket, Stump Pond; 41.292454N, 70.002666W; 05 Sep. 2019; Charley Eiseman leg.; Parthenocissus quinquefolia; https://www.inaturalist.org/observations/44821470. • New York • Putnam Co., Putnam Valley, 392 Dennytown Rd.; 41.41309N, 73.86508W; 09 Aug. 2016; Even Dankowicz leg.; Parthenocissus; https://www.inaturalist.org/observations/17908231, https://www.inaturalist.org/observations/17908234. – Rhode Island • Washington Co., Block Island; 41.195N, 71.5652W; 08 Aug. 2019; Aaron Hunt leg.; Parthenocissus; https://bugguide.net/node/view/1741669/bgimage. – Vermont • Chittenden Co., Shelburne; 44.39445N, 73.23003W; 14 Sep. 2019; Spencer Hardy leg.; Parthenocissus quinquefolia; https://www.inaturalist.org/observations/32714140. – Wisconsin • Dane Co., Madison, UW Lakeshore Nature Preserve; 43.084N, 89.429W; 25 Aug. 2019; Tom Klein leg.; Parthenocissus quinquefolia; https://bugguide.net/node/view/1717010/bgimage.
Antispila voraginella
Braun, 1927b: 191. Holotype ♂: USA: [Utah: Washington County] “B1206/Zion Canyon/Utah i.iv.9 [1926]- Antispila / voraginella / Type Braun.”, Genitalia slide EJvN 3916 [reared from mines on Vitis arizonica] (
Antispila voraginella;
Wingspan ca. 4.7–5.4 mm, forewing length 2.3–2.6 mm. Externally separated from A. oinophylla and ampelopsifoliella by the darker brassy brown head scaling, the absence of a fringe line and the overall less shining metallic pattern. Relatively similar to A. argentifera, but unlikely to be sympatric with that species. Male genitalia with distinctly bilobed tegumen, fewer pecten sensilla (8–10) and phallotrema with smaller spines only.
Aspilanta species, larval morphology, final feeding instar (4th) and non-feeding instar (5th, 36). 34 A. ampelopsifoliella, ventral aspect, ethanol preserved larva,
Phylogeny of the cosmopolitan leafmining clade of Heliozelidae, part of fig. 1 in
Vitaceae: Vitis arizonica, unidentified Vitis.
(Figs
Pale yellowish, head and prothorax hardly darker, mouthparts stand out as darker brown; a row of 5–7 brown spots on abdomen visible (Fig.
Larvae found between 4 June and 9 August (Arizona); in all cases the adults emerged the next spring (April to June), suggesting that the species is univoltine.
USA: Arizona, Texas (west), Utah.
BIN: BOLD:AAU3733, average distance 0.92%, max. distance 1.38% (n = 3), distance to nearest neighbour 8.83% (BOLD: AAV5059, A. hydrangaeella).
None known.
Adults: see van
USA – Arizona • Cochise Co., Portal; 31.90918N, 109.25228W; 03 Aug. 2019; Laura Gaudette leg.; Vitis; https://www.inaturalist.org/observations/30135988, https://www.inaturalist.org/observations/32296900. – Utah • Washington Co., Zion National Park; 37.23N, 112.963W; 22 Jul. 2008; Charley Eiseman & Noah Charney leg.; Vitis; https://www.inaturalist.org/observations/44822019.
Antispila argentifera
Braun, 1927a: Holotype ♀, Canada, Ontario, Sparrow Lake, 16.vii.1926, A. Braun, on young leaves of birch. (
Antispila argentifera;
Antispila
species on sweet fern;
Antispila
n. sp.,
Wingspan ca. 4.0–5.2 mm, forewing length 1.8–2.4 mm. Externally separable from A. oinophylla and ampelopsifoliella by the darker brassy head scaling, the grey fringe and the indistinct fringe line; A. argentifera is also smaller. Relatively similar to A. voraginella which is only known from the Southwestern states. Male genitalia with tegumen distinctly bilobed, valvae with 9–12 pecten sensilla, phallus without strong appendices (only one slide examined). Mines on Morella distinguished from those of an unnamed Coptodisca species, which occurs especially in the Southeast, by the larger cut-out, and the overall larger mine.
Aspilanta species, live adult moths. 45 A. oinophylla, male, Italy, Südtirol, Eisacktal, Klausen, 650 m, e.l. 1.viii.2019, photograph Dieter Robrecht 46 A. hydrangaeella, female, dorsal view, NC, photograph EJvN 47 A. ampelopsifoliella, female, Pelham MA, photograph CSE 48 A. ampelopsifoliella, male, dorsal view, VT, photograph EJvN 49 A. argentifera, female, Stockbridge MA, photograph CSE 50 A. viticordifoliella, female, Pelham MA, photograph CSE.
Aspilanta oinophylla, leafmines, larvae and shields. 51, 52 GA, Chattahoochee NF, 14.x.2010, type locality, Vitis aestivalis, EvN2010266 53, 56 MA, Northfield, 13.ix.2016, V. riparia, CSE 54, 58 TN, Great Smoky Mts NP, 2.x.2010, V. vulpina, EvN2010119 55, 57 VT, Button Bay SP, 16.ix.2011, V. riparia, EvN2011253.
Myricaceae: Comptonia peregrina, Morella caroliniensis (including M. pensylvanica, cf.
(Figs
Pale yellowish green, head and prothorax brown.
Bivoltine. Larvae found in July and September to November, adults in June, July and August; specimens reared from fall mines emerged (indoors) in April to May, from July mines in August. Mines of the first generation seem to be very scarce.
Canada: Newfoundland*, Nova Scotia, Ontario, Prince Edward Island*, Quebec*; USA: Connecticut*, Massachusetts*, New York*, North Carolina*, Vermont.
BIN: BOLD:ACG8654, average distance 0.66%, max. distance 2.25% (n = 15), distance to nearest neighbour 11.67% (BOLD:ACF9350, an unidentified Heliozelidae from Honduras).
Eulophidae: Pediobius ? albipes (Provancher, 1887) (NY), Pnigalio sp. (MA).
Annette
Specimens from Prince Edward Island were previously regarded as misidentifications (van
CANADA – Nova Scotia • 1 ♂; Barrington; 43.56N, 65.57W; G.G. Lewis leg.; Comptonia peregrina; emerged 20 Aug. 1970; EventId: 70–28; CNCLEP00122407. – Ontario • 1 ♀ (Holotype, photograph examined, Fig.
USA – Massachusetts • 1 ♀; Berkshire Co, Stockbridge, Kampoosa Bog; 42.294364N, 73.304347W; 16 Jul. 2017; C. S. Eiseman leg.; Myrica gale; emerged 09 Aug. 2017; EventId: CSE4096;
USA – Connecticut • 1 larva, leafmines; New London Co, Connecticut College Arboretum; 41.37929N, 72.11121W; alt. 60 m; 10 Sep. 2011; E.J. van Nieukerken leg.; Morella caroliniensis; EventId: EvN no 2011194–2M;
CANADA – Newfoundland and Labrador • 1 ♀; Barachois Pond Provincial Park, Erin Mtn trail; 48.469N, 58.256W; alt. 240 m; 29 Jun. 2011; G.R.Pohl, D.W.Langor leg.; CCDB–23267–F01; NFC. • 1 ♂; J. T. Cheeseman Provincial Park; 47.633N, 59.255W; 27 Jun. 2011; G.R.Pohl, L.Lafosse leg.; CCDB–23267–E06; NFC. • 1 ♀; Hwy 1, 5 km N jct Rte 480; 48.561N, 58.264W; alt. 138 m; 25 Jun. 2011; G.R.Pohl, D.W.Langor leg.; CCDB–23267–F05; NFC. – Quebec • 3; Mingan Archipelago National Park Reserve, Quarry Island; 50.2135N, 63.7979W; 19–26 Jun. 2013; Park Staff leg.; EventId: GMP#01092; BIOUG12280–D02, BIOUG12281–E02, BIOUG12281–E06. • 3; same locality data as preceding; 02–09 Jul. 2013; Park Staff leg.; EventId: GMP#01096; BIOUG12352–A11, BIOUG12352–H01, BIOUG12398–F05. • 1; same locality data as preceding; 16–24 Jul. 2013; Park Staff leg.; EventId: GMP#01100; BIOUG12408–H03.
USA – Massachusetts • Plymouth Co., Marshfield, Hoyt Hall Preserve; 42.066N, 70.678W; 29 Oct. 2016; Steven Whitebread leg.; Morella caroliniensis; (photographs examined). – New York • Orange Co., Cornwall, Black Rock Forest; 41.39621N, 74.02522W; 30 Aug. 2019; Charley Eiseman & Julia Blyth leg.; Comptonia peregrina; https://www.inaturalist.org/observations/37506602. – North Carolina • Scotland Co., Laurinburg, St. Andrews University; 34.747N, 79.477W; 27 Sep. 2016; Tracy Feldman leg.; Morella cerifera; https://bugguide.net/node/view/1298044/bgimage same locality as preceding; 08–09 Nov. 2016; Tracy Feldman leg.; Morella cerifera; https://bugguide.net/node/view/1312510/bgimage, https://bugguide.net/node/view/1312770/bgimage • same locality as preceding; 16 Oct. 2017; Tracy Feldman leg.; Morella cerifera; https://bugguide.net/node/view/1456019/bgimage.
Antispila viticordifoliella
Clemens, 1860: 209. Syntype mines, larva [USA: Pennsylvania, Easton], larvae on “wild grapes” [Vitis vulpina], August–September, Brackenridge Clemens (
Antispila viticordifoliella;
Antispila cf. viticordifoliella
van
Wingspan ca. 5.5 mm, forewing length 2.2–2.5 mm. Externally different from all other Aspilanta species by the absence of the apical spot in the forewings. Differs from most Antispila species, which have a similar wing pattern, by the antennae with distinct white tips, the complete absence of androconial scales in the males and the smaller size. Genitalia not yet examined. Leafmines differ from other Aspilanta on Vitaceae by the absence of a linear portion and by the larva that is yellow-green (vs. whitish/colourless in A. ampelopsifoliella and oinophylla); from Antispila species by the black dispersed frass rather than brown and the markedly smaller cut-out. The record of an Antispila sp. from similar mines (see Remarks) complicates this, though.
Aspilanta hydrangaeella, leafmines, larvae and shields, on Hydrangea arborescens. 59, 60, 64 GA, Chattahoochee NF, 14.x.2010, EvN2010279 61, 63, 68 OH, South Bloomingville, 13.ix.2012, CSE, (61=
Aspilanta ampelopsifoliella, leafmines, larvae and shields, on Parthenocissus quinquefolia. 69, 74 VT, Williston, 28.viii.2016, CSE3764 (same mine) 70, 76–78 MA, Pelham, 30.viii, 4.ix.2013, CSE1102, 1108 71 MA, Nantucket, Squam Swamp, 7.ix.2011, CSE-C4 72 MA, Northfield, 16.ix.2016, CSE 73 MA, Nantucket, Stump Pond, 5.ix.2019, CSE 75 VT, Button Bay SP, 16.ix.2011, EvN2011254 79 MA, Bridgewater, 16.viii.2013, CSE983.
Vitaceae: Parthenocissus quinquefolia, P. vitacea, Vitis vulpina.
(Figs
Yellow-green except for green gut contents, in contrast with colourless or whitish larva of A. ampelopsifoliella on the same host plant (compare Figs
Larvae found from August to September through most of the range, but in Florida in April. Field caught adults found in July; larvae collected in fall resulted in emerging adults in April-May, the adult from the April larva emerged in May.
Canada: Ontario, Québec*; USA: Connecticut, Florida, Kentucky, ? Maine, Massachusetts*, New York, Pennsylvania, ? Texas, Vermont. The previous record from Vermont by
BIN: BOLD:ABA2832 (CT, FL), average distance 0.32%, max. distance 0.8% (n = 5), distance to NN 3.53% (next BIN), BIN: BOLD:AAZ5646 (NY, VT), average distance 0.25%, max. distance 0.46% (n = 4), distance to NN 2.89% (BOLD:ACA6487). The single specimen in a Malaise trap with that BIN: BOLD:ACA6487, most likely also belongs to A. viticordifoliella, to be confirmed by morphological examination.
Antispila viticordifoliella was described on the basis of larvae and leafmines on Vitis vulpina (as V. cordifolia), with insufficient detail to recognise the species. The first adults were described by
Earlier we excluded the specimens reared from Parthenocissus from this species (van
From very similar mines in Parthenocissus collected in North Carolina, CSE and Tracy Feldman have reared an unidentified species of Antispila; we have thus omitted possible records of Aspilanta viticordifoliella from Iowa, Kansas and Ohio that are based only on photographed mines. In photographs of the North Carolina mines taken in the field, the larvae appear to be paler than those of A. viticordifoliella, but this requires confirmation; the Antispila mines are otherwise only recognisable when the typical keeled case is formed. The reared adults in fact also resemble A. viticordifoliella because of the white-tipped antennae; only checking wing venation and/or the genitalia will separate the two.
CANADA – Ontario • 1 ♀; Ottawa; 45.41N, 75.69W; G.G. Lewis leg.; emerged 02 Apr. 1971; EventId: 70–48; Genitalia slide: MIC1876; CNCLEP00100475.
USA – Massachusetts • 1 ♀; Hampshire Co., Pelham, 88 Arnold Rd.; 42.3629N, 72.4598W; 30 Aug. 2013; C.S. Eiseman leg.; Parthenocissus quinquefolia; emerged 13 May. 2014; EventId: CSE1109; CSEC.
CANADA – Québec • 1 vacated mine; Gatineau, Aylmer E, near Ottawa river; 45.39023N, 75.79139W; alt. 56 m; 23 Aug. 2015; E.J. van Nieukerken leg.; Parthenocissus vitacea; EventId: EvN no 2015160–1H;
USA – Massachusetts • Hampshire, Northampton Co., Northampton Bikeway west of King St.; 42.329151N, 72.637529W; 13 Sep. 2013; Charley Eiseman leg.; Parthenocissus quinquefolia; CSEC. – Vermont • Chittenden Co., Williston, Mud Pond; 44.413625N, 73.075697W; 28 Aug. 2016; Charley Eiseman leg.; Parthenocissus quinquefolia; CSEC.
CANADA – Ontario • 1; Point Pelee National Park, Cactus Field; 41.939N, 82.516W; alt. 168 m; 27 Jun.–04 Jul. 2012; Tyler Peters leg.; EventId: GMP#00175; BIOUG03514–A11. • 1; Wellington County, Puslinch Township, Concession 11/Hume Rd; 43.537N, 80.134W; alt. 320 m; 18–25 Jul. 2010; Paul Hebert leg.; EventId: L#PHPUS–017; BIOUG01146–B07.
USA – Tennessee • 1 ♀; Great Smoky Mountains National Park, Twin Creeks Science and Education Center; 35.6859N, 83.4986W; alt. 559 m; 10–17 Jul. 2012; Becky Nichols leg.; EventId: GMP#00037; BIOUG02919–H02.
CANADA – Ontario • Renfrew, Killaloe, Hagarty and Richards; 45.60491N, 77.59134W; 05 Sep. 2019; Jason J. Dombroskie leg.; Parthenocissus; https://www.inaturalist.org/observations/32173691. • Renfrew, Wilno; 45.60508N, 77.59131W; 05 Sep. 2019; Jason J. Dombroskie leg.; Parthenocissus; https://www.inaturalist.org/observations/32173324.
USA – Massachusetts • Berkshire Co., Great Barrington; 42.197001N, 73.335001W; 16 Sep. 2017; Charley Eiseman & Julia Blyth leg.; Parthenocissus quinquefolia; https://www.inaturalist.org/observations/44823345.
Antispila
Vitis1; van
Externally not distinguishable from A. oinophylla or A. ampelopsifoliella. Genitalia and leafmines not yet examined.
Vitaceae: Vitis aestivalis, Vitis spec.
Not examined.
Larvae in June and July, single adult reared in July. Possibly bivoltine.
USA: Connecticut, Florida.
BIN: BOLD:ABA2832, average/max. distance 0.35% (n = 2), distance to NN 9.72% (BIN: BOLD:AEA4069, unknown, not a public record).
See van
Aspilanta from SW USA, leafmines, larvae and shields on Vitis arizonica. 80, 81 A. voraginella, AZ, Cochise Co., 3.viii.2019, photographs by Laura Gaudette 82, 83 A. voraginella, US, Zion National Park (type locality), 22.vii.2008, lighting from resp,. above and beneath mines, photographs by Noah Charney 84–87 Aspilanta “Vitis.arizonica_USA”, AZ, Cochise Co., 11.xi.2012, CSE–L141.
Aspilanta argentifera, leafmines, larvae and shields. 88 NC, Laurinburg, 27.ix.2016, Morella cerifera, CSE 89 MA, Northfield, 26.x.2015, Comptonia peregrina, CSE 90 VT, Chickering Bog, 6.ix.2015, Myrica gale, EvN2015222, showing wilted leaf tip 91, 96 MA, Erving, 27.x.2014, Comptonia peregrina, CSE1557 92, 93 MA, Nantucket, Radar Hill, 15.xi.2015, Morella caroliniensis, CSE2398, same mine at different stages 94 MA, Stockbridge, 16.vii.2017, Myrica gale CSE4096, showing very long linear portion directly along midrib 95 MA, Nantucket, UMass field station, 5.xi.2017, Morella caroliniensis, CSE4546.
Antispila
sp.;
Adult unknown. Larvae solitary, not forming communal mines as in A. voraginella; different timing.
Vitaceae: Vitis arizonica.
(Figs
Not examined.
Larvae collected in November.
USA: Arizona.
A barcode of 407 bp groups in our tree (Fig.
It is yet not certain whether this species belongs really to Aspilanta, as the (incomplete) barcode seems to group more with Holocacista species.
USA – Arizona • 1 larva (barcoded), vacated mines; Cochise Co., Coronado National Forest, near Chiricahua National Monument; 31.978922N, 109.357056W; alt. 1675 m; 11 Nov. 2012; C.S. Eiseman leg.; Vitis arizonica; EventId: CSE–CH1/ CSE–L141;
Aspilanta viticordifoliella, leafmines, larvae and shields on Parthenocissus quinquefolia and P. vitacea (104). 97, 99, 100, 105, 106 MA, Northampton, 13.ix.2013, CSE, 105 shows the left mine of 100, 106 showing flat shield to distinguish it from Antispila species 98 VT, Williston, 28.viii.2016, CSE 101 MA, Great Barrington, 16.ix.2017, CSE 102, 103 VT, Button Bay SP, 16.ix.2011, EvN2011254–2 104 QC, Aylmer, 23.viii.2015, EvN2015160.
The phylogenetic basis for the erection of the genus Aspilanta is the transcriptomic study, which provides a high support for its monophyly (
1) The inclusion of the whole Holocacista group in an enlarged genus Antispila. Although this would have solved the problem of polyphyly, it would create a large genus with huge differences in many characters, both encompassing species with reduced venation and the more complete venation, and make recognition of species belonging to this genus almost impossible.
2) Combining all genera of the Holocacista group into one genus. Such a genus would be monophyletic and be based on a whole suite of characters, such as the reduced venation. Such a genus should also encompass the undescribed group I (
The choice of generic borders is in principle not ruled by any other criterion than monophyly, and thus partly subjective. In all three options a number of changes in names are to be expected, but in the chosen option this is limited to just six species, only one of which is an economically important insect. In alternative 1) the number of new combinations would be highest, 26 in total (all Coptodisca, Holocacista and some others, including five or more economically important species), in 2) still 15 are needed (Holocacista, the current Aspilanta species, Antispilina, Ischnocanaba, including at least three economically important species). So in conclusion, the choice for a new genus Aspilanta was obvious in terms of stability and recognisability of the groups. Also biogeographically this choice has advantages, as we have now two old world genera (Holocacista, Antispilina) and two new world genera (Aspilanta, Coptodisca), whereas Antispila remains almost global (unknown from Australia, and so far from the Neotropics). Further study of the Neotropical fauna is important to see whether the monophyly of Aspilanta will stand, especially in the light of the position of the Patagonian group of species cited as “Neospila”. If they belong to Aspilanta, which seems likely morphologically, the paraphyly of the combination Genus14 + Antispila group II as noted by
We choose to describe this genus without a full revision of its species. For several of the species we lack sufficient material, and additional fieldwork is much needed, not only for this genus, but for all North American Heliozelidae. Additionally, in Antispila, Coptodisca, and Heliozela we already know of several unnamed species, partly only from Malaise trapped specimens for barcoding campaigns, but also online photographs suggest there are more species. Inclusion of the rich Mexican fauna in such revisions would also be desirable. Further it would be important to settle the status of some old names by dissecting old types while extracting DNA, and for some species to designate neotypes.
By removing six species from Antispila to Aspilanta, the remaining five named North American species feeding on Cornaceae, Nyssaceae, and Vitaceae belong in the monophyletic genus Antispila, as shown by a combination of the molecular studies and their morphology (
Antispila eugeniella
Busck, 1900: 236. Holotype adult, USA, Florida, Palm Beach, leafmine on Eugenia sp., bred 25 Feb. 1900, “819”, [leg. Dyar], Type No. 4945, USNMENT 01476200 (
Antispila eugeniella;
This species is rare; since the unique holotype was reared, we are only aware of a few specimens reared by David L. Wagner (pers. comm.); all subsequent literature records to
We illustrate the holotype here (Fig.
The colour pattern of Antispila and Aspilanta is almost always with a fascia at one third and two opposite spots at two-thirds. The pattern of H. eugeniella is different, with a postmedial fascia, and resembles more some species of the Old World genus Holocacista (van
For information, discussions, assistance with analyses, and loans of specimens we are grateful to the following persons: Richard L. Brown, Donald R. Davis, Camiel Doorenweerd, Mike Halsey, Doug Hilton, Axel Kallies, Jean-François Landry, Liz Milla, Greg R. Pohl, Jeremy deWaard, David L. Wagner, and Andy Young. Christer Hansson, Sandrine Ulenberg, and José Fernández-Triana are thanked for the identification of parasitoids and further information on these. We thank Noah Charney, Laura Gaudette, Mike Palmer, and Dieter Robrecht for allowing us to reproduce photographs they made. BOLD staff helped providing and correcting records from the BOLD database. Camiel Doorenweerd, Frank Stokvis, and Ruben Vijverberg performed DNA analyses and Sjaak Koster made larval and genitalia preparations. EvN thanks Jean-François Landry and Vazrick Nazari for hospitality and assistance during a three-month stay at