Revision of the genus Philonome Chambers and its proposed reassignment to the family Tineidae (Lepidoptera, Tineoidea)

Abstract The New World genus Philonome Chambers, 1874 is revised. This genus comprises twelve species, seven of which are described as new: two species, Philonome nigrescens sp. n. and Philonome wielgusi sp. n., from the United States; four species, Philonome albivittata sp. n., Philonome curvilineata sp. n., Philonome kawakitai sp. n., and Philonome lambdagrapha sp. n., from French Guiana; and one species, Philonome penerivifera sp. n., from Brazil. Lectotypes are designated for Philonome clemensella Chambers, 1874 and Philonome rivifera Meyrick, 1915. Partially on evidence of their head morphology and particularly from molecular evidence, the genus Philonome, previously associated with Bucculatricidae or Lyonetiidae, is reassigned to Tineidae. A possible systematic position of Philonome within Tineidae is discussed. Eurynome Chambers, 1875, is synonymized with Argyresthia Hübner, 1825 (Argyresthiidae). Photographs of adults and illustrations of genitalia, when available, are provided for all described species of Philonome and two species previously misplaced in Philonome, Argyresthia luteella (Chambers, 1875) and Elachista albella (Chambers, 1877). In addition, DNA barcodes were used for the delimitation of most species.


Introduction
The monobasic genus Philonome was proposed by Chambers in 1874 for Philonome clemensella Chambers. Chambers (1875) later proposed a supposedly allied genus, Eurynome, also based on a single species, Eurynome luteella Chambers, and in 1877, added another congener, Eurynome albella Chambers. Eurynome, however, was recognized as a homonym and later replaced by Busckia Dyar, 1903. Chambers (1875, 1877, 1880 assigned Philonome to the Tineina, a conventional group name to accommodate primitive Microlepidoptera, and he further suggested that the genus is allied to Bucculatrix Zeller, 1839. The putative association between Philonome and Bucculatrix has been repeatedly expressed by subsequent researchers such as Meyrick (1915Meyrick ( , 1920 and Forbes (1923). Barnes and McDunnough (1917) included Philonome under Lyonetiidae, together with Bucculatrix, followed by Forbes (1923), but they treated Busckia (= Eurynome Chambers) as a genus of Elachistidae. McDunnough (1939) transferred Busckia to Lyonetiidae and synonymized it with Philonome. Sohn et al. (2013) conducted a molecular phylogeny of Yponomeutoidea (to which Lyonetiidae belongs), including Philonome clemensella, and found that the species is nested within the Tineidae (Fig. 1). However, the tineid association of Philonome has been so far supported only by molecular data, not yet by morphological evidence.
Philonome currently includes six species, which occur exclusively in the New World: two from the Nearctic Region and four from the Neotropical Region. Eurynome albella Chambers (Figs 17,70), known only from the unique holotype collected at Edgerton (38°57'24"N, 104°50'6"W; at ~ 6500 feet elevation), El Paso Co., Colorado, was once treated as Philonome (McDunnough 1939), but it was later assigned to Elachista of Elachistidae (Kaila 1999). Kaila (1999) found that the name Elachista albella (Chambers) had been preoccupied and hence he proposed a replacement name, Elachista dasycara. Chambers (1874Chambers ( , 1877 characterized Philonome and Eurynome on superficial appearance and wing venation. The adults resemble some species of Bucculatrix in wing pattern, notably B. adelpha Braun, 1963, or B. angustata Frey & Boll, 1876. However, Philonome differs from Bucculatrix in having an elongate, telescopic ovipositor and lacking an androconial scale pocket on the male abdomen (Braun 1963;Kobayashi et al. 2010). This suggests that their resemblance is due to convergence. The biology of Philonome is essentially unknown. Forbes (1923) stated that P. clemensella have been collected from hickory and linden trees. His statement, however, was based on the ambiguous label data of specimens from the United States National Museum of Natural History. No additional observation of the larvae of P. clemensella has been reported from these trees.
The goals of this paper are to redefine the generic characteristics of Philonome, to describe seven new species from the Nearctic and Neotropical Regions, to transfer a misplaced species, "Philonome" luteella to its correct genus, Argyresthia, and to provide morphological evidence of the tineid relationship of Philonome, which has been suggested from a recent molecular phylogenetic study (Sohn et al. 2013).  Sohn et al. (2013), based on 27 nuclear genes. Branches in bold indicate the > 70% bootstrapping support from at least one analysis attempted by Sohn et al. (2013). The 'A' in closed circle represents a well-supported subclade of Tineidae in which Philonome clemensella is included.
Selected specimens were dissected for genitalia and abdominal structures, following Clarke (1941), except that Chlorazol black was used for staining. Dissected genitalia were mounted on microscope slides in Euparal resin (BioQuip Products Inc.) or Canada balsam. Pinned specimens were examined under a Leica MZ APO stereoscope. Slidemounted specimens were examined under a Leica LEITZ-DMRX microscope. All illustrations were drawn from dissections temporarily stored in glycerin, which were later permanently embedded in mounting medium. Terms for genitalia and wing venation follow Klots (1970) and Wootton (1979), respectively. The 8 th abdominal segment is abbreviated as A8 in the descriptions. Verbatim label data are given for primary types. Additional data by the present authors are given in brackets.
DNA was extracted from hind legs of dried specimens. DNA barcodes (658 bp of the COI mitochondrial gene) were generated at the Canadian Centre for DNA Barcoding (CCDB, Guelph). A total of seven specimens were sequenced (Table 1), all collected in French Guiana by the third author (CLV). These newly generated barcodes were compared to five DNA-barcodes of Philonome clemensella (Table 1), one (jflandry0875) available at the Barcode of Life Data Systems (BOLD; www.boldsystems.org; also see Ratnasingham and Hebert 2007) and the other four unpublished. Barcode data were analysed using the analytical tools of BOLD such as Neighbour Joining and pairwise genetic distance matrix.
Details on the date and site of collection for each specimen, as well as a photograph are available through the DOI (http://dx.doi.org/10.5883/DS-PHILONO). The same DOI provides access to the sequence records and GenBank accession numbers (Table 1). Table 1. Specimens used for the DNA barcoding analysis. Both the Process ID and sample ID codes are unique identifiers linking the record in the BOLD database and the voucher specimen from which the sequence is derived. Additional collecting and specimen data are accessible in BOLD's data set (http:// dx.doi.org/10.5883/DS-PHILONO) as well as GenBank (http://www.ncbi.nlm.nih.gov/genbank/). Adult. Head (Fig. 18): Vestiture of vertex rough with piliform scales; frons smooth with broad, flat, appressed scales; a band of broad, spatulate scales between the bases of the antennae, along the transfrontal suture, bounded both above and below by piliform scales. Antenna filiform in both sexes; antennal pecten absent; scape elongate, ~2.2-2.4× length of adjacent pedicel. Labial palpus without bristle-like setae; 2 nd segment 2× longer than 1 st , as long as 3 rd . Maxillary palpus 5-segmented, longer than labial palpus. Proboscis naked, shorter than maxillary palpus.
Biology. Chambers (1878) mentioned that he repeatedly collected Philonome clemensella from the type locality where Gleditschia triacanthos L., Ulmus americana L., Prunus serotina Ehrh., and Celtis occidentalis L. grow in the immediate vicinity. He then assumed that the larvae may feed on some weeds or shrubs growing nearby. Forbes (1923) noted that the larvae of P. clemensella feed on hickory and linden. These records were, however, based on ambiguous label data which state only plant names without details and thus require verification. Nothing is known about the biology for other congeners of P. clemensella.
Female unknown. Types. Holotype: male, "ARIZONA: Cochise Co.: Sierra Vista 5131 Bannock 2 IX 1988", "Attracted to (E, Z) -3 13 ODDOH @ 1615-1730 hrs.", "R. S. Diagnosis. This species is easily distinguished from all other congeners in possessing a black ground-color of the forewing and an elongate process on the transtila of the male genitalia.
Types   (Forbes 1923). These are from the label data in the USNM collection. The collection also includes a specimen whose label data states that it came from oak (Fagaceae: Quercus). The label data give no details other than plant common names. Therefore, it is not clear if these records refer to larval host plants or where the adults were collected. Diagnosis. This species is similar to Philonome curvilineata in external appearance but differs from the latter in having the longitudinal and costal fasciae separate (continuous in P. curvilineata and larger apical protrusion on the tegumen in the male genitalia.
Thorax: Patagium dark brown; tegula lustrous pale yellow, intermixed with orange scales basally; mesonotum silvery white with dark brown transverse band along anterior and posterior margins and at anterior 1/3, with orange transverse band at middle. Foreleg with coxa lustrous pale yellow; femur lustrous pale yellow, intermixed with pale gray- ish brown laterally; tibia dark brown dorsally, pale grayish yellow ventrally; tarsomeres dark brown dorsally, pale orange ventrally. Midleg with coxa and femur lustrous pale yellow; tibia and tarsomeres grayish brown dorsally, pale yellow ventrally. Hindleg consumed for DNA extraction. Forewing length 3.0 mm (n = 1), reddish brown; costa black in distal 1/3; longitudinal fascia extending to apical streak, straight, white on basal 1/2, juxtaposed with a slender, intermittent black line along lower border, sinuous, black on distal 1/2; costal fasciae slender, extending to apex; subapical streak white, juxtaposed with slender black line along lower border; apical streak white, connected with longitudinal fascia; dorsal bar white, juxtaposed with black along outer border; dorsal margin sparsely irrorated with black scales on basal 1/6 and at middle; tornal patch elongate, white, juxtaposed with black along upper border, irrorated with dark brown scales along outer border; marginal streak dark brown; fringe brown on distal 1/3 of costa, pale yellowish gray along termen. Hindwing brownish gray; fringe pale grayish brown.
Male genitalia (Figs 37-39): Tegumen nearly as long as valva, semi-elliptical on basal 3/4, rectangular on distal 1/4, with small lateral protrusion dorsoposteriorly. Valva elongate, lobate, sparsely setose on outer surface. Juxta liguiform, 1/2 as long as valva. Vinculum broad, gradually broadened anteriorly, with medial and lateral protrusions along anterior margin; Phallus slightly curved at basal 2/5, broadened posteriorly. Distribution. French Guiana. Etymology.The species name is derived from the Greek letter 'lambda' and a suffix derived from the Greek 'graphein' meaning "to write", and refers to the white fascia of the forewing resembling a lambda (λ). Diagnosis. This species is indistinguishable from Philonome rivifera Meyrick in external appearance but differs from the latter in having the apex of the valva in the male genitalia entire (vs. bifid in P. rivifera).
Thorax: Scales of patagium orange with dark brown tips; tegula pale orange, intermixed with orange scales basally; mesonotum lustrous orange white, with pale orange transverse band along anterior and posterior margins and at middle. Foreleg with coxa and femur lustrous pale orange; tibia orange, intermixed with dark brown scales dorsally, orange white ventrally; tarsomeres orange dorsally, pale orange ventrally; first tarsomere sparsely intermixed with dark brown scales dorsally. Midleg with coxa and femur lustrous pale orange; tibia brownish orange dorsally, lustrous orange white ventrally; tarsomeres pale orange dorsally, pale yellow ventrally. Hindleg consumed for DNA extraction. Forewing length 2.8 mm (n = 1), reddish brown, slightly paler along dorsal area; costal area yellowish brown on basal 1/2, brownish white above the curvature of longitudinal fascia, pale orange on distal 1/4, intermixed with black scales on middle and distal 1/4 of costa; longitudinal fascia continuous to near apex; convex at distal 1/3, white, juxtaposed with slender black line along lower border; dorsal bar straight, white, juxtaposed with slender, intermittent, black line along outer border; black irroration at middle of dorsal margin and on tornal area; fringe orange on distal costa and apex; scales of fringe along termen pale yellowish gray on basal 2/3, black on distal 1/3. Hindwing brownish gray; fringe pale grayish brown.
Thorax: Patagium lustrous pale yellow; tegula white, intermixed with pale orange scales basally; mesonotum white on anterior half, lustrous reddish brown on posterior half, with a dark brown transverse band medially. Foreleg with coxa lustrous pale yellow; femur lustrous dark grayish brown laterally, lustrous pale yellow mesally; tibia and tarsus dark brown dorsally, pale grayish yellow ventrally. Midleg with coxa lustrous pale yellow; femur lustrous pale orange dorsally, lustrous pale yellow laterally and ventrally; tibia pale orange, intermixed with dark brown scales dorsally; tarsomeres orange dorsally, pale orange ventrally. Hindleg with coxa and femur lustrous pale orange; tibia pale brownish orange dorsally, pale orange ventrally, with stiff piliform scales; tarsomeres pale orange. Forewing length 2.8-3.1 mm (n = 3); reddish brown; costa brown; longitudinal fascia white, spanning entire costal area except costa; lower margin sinuous, accompanied with narrow, dark brown line; dorsal bar white, at basal 1/3 of dorsum, dentiform, accompanied with dark brown bar along upper margin; marginal area dark brown; fringe brownish gray. Hindwing and fringe brownish gray.
Thorax: Scales of patagium pale orange, with dark brown tips; tegula reddish brown basally, paler to apex, pale orange apically; mesonotum pale orange, transversely intermixed with dark brown scales at middle. Fore-and midlegs with coxa lustrous yellowish white; femur, tibia, and tarsomeres dark brown dorsally, lustrous yellowish white laterally and ventrally. Hindleg pale brownish gray dorsally, lustrous yellowish white laterally and ventrally. Forewing length 3.2-3.6 mm (n = 2), coloration and patterns similar to P. rivifera. Hindwing dark brownish gray; fringe brownish gray on costal and apical area, yellowish gray along posterior margin.
Abdomen: Terga pale grayish orange or pale grayish brown; sterna lustrous, white or pale orange.
Types. Holotype: male, "Holo-Type" [circular label with red border], "Para Brazil Distribution. Brazil (Amazonas, Federal District, Pará). Etymology. The species name is derived from the Latin prefix 'pene (= paene)', meaning "almost", and the preexisting species name, rivifera, and refers to the overall similarity of this species to Philonome rivifera.
Remarks. The holotype and three paratypes of Philonome penerivifera in the BMNH collection were misidentified as P. rivifera by Edward Meyrick. Diagnosis. The female genitalia of P. kawakitai is similar to those of P. penerivifera but differ from the latter in having the lamella antevaginalis of the seventh sternite more rounded (obliquely truncate posteriorly in P. penerivifera) and in the absence of microscopic spicules in the corpus bursae. Philonome kawakitai is distinguished from P. curvilineata, P. penerivifera, and P. rivifera in having the dorsal bar not reaching the dorsal margin and the complete subterminal line on the forewing. Adult (Fig. 11). Head: Vertex pale brown, sparsely intermixed with dark brown scales posterolaterally, pale orange on anterior 1/3; frons lustrous pale grayish yellow. Antenna 8/9 as long as forewing; scape pale orange dorsally, lustrous yellowish white ventrally; first flagellomere dark brown dorsally, pale yellow ventrally; second to 9 th flagellomeres pale orange dorsally, pale yellow ventrally; remaining flagellomeres pale grayish yellow. Labial palpus 1/2 as long as maxillary palpus, dark yellowish brown. Maxillary palpus dark yellowish brown.

Philonome rivifera
Thorax: Patagium pale orange, tinged with dark brown distally; tegula lustrous pale yellow, intermixed with brown-tipped, orange scales basally; mesonotum pale orange, anterior 1/3 and posterior 1/3 lustrous pale yellow, with a narrow transverse band of dark brown-tipped scales. Fore-and midlegs with coxa lustrous pale yellow; femur pale orange dorsally, lustrous pale yellow laterally and ventrally; tibia and tarsomeres pale brown dorsally, pale orange ventrally. Hindleg with coxa and femur lustrous pale orange; tibia lustrous pale orange, with long piliform scales ventrally; tarsomeres orange dorsally, pale orange ventrally. Forewing length 2.8-4.6 mm (n = 3), reddish brown in medial area, orange in terminal 1/3 of costal area, pale orange in basal 2/3 of costal area and in basal 1/2 of dorsal area; longitudinal fascia white, continuing to subapical area, accompanied with a slender, dark brown line along lower margin, curved to costa at terminal 1/3; dorsal bar white, narrow, connected to white spreading on dorsum; distal area of costa, termen, and apical area densely irrorated with dark brown; elongate scales of fringe pale grayish brown, with dark brown tip; piliform scales of fringe pale orange. Hindwing dark grayish brown; fringe purplish gray.

Philonome spectata
Thorax: Patagium and mesonotum reddish brown; tegula white. Legs pale orange. Forewing length 2.3 mm (n = 1), reddish brown; longitudinal fascia white, covering most costal area, lower margin sinuous, accompanied with very narrow dark brown line; costa suffused with pale orange subbasally and in terminal 1/3; elongate scales of fringe around apex reddish brown with dark brown tips; piliform scales of fringe on  Distribution. Brazil (Pará).

Remarks.
Only the holotype of Philonome spectata is known to exist. It was not possible to examine this specimen and to illustrate the genitalia. This species can be distinguished from other congeners in lacking the dorsal bar on the forewing. Philonome sp. Fig. 14 Note. Forewing length 4.1 mm (n = 1). This species is indistinguishable from P. rivifera in superficial appearance. Our DNA-barcoding data show that it is distinct from other congeners from French Guiana and P. clemensella, and may be genetically closest to P. kawakitai (Fig. 71). The only specimen of this species has its abdomen missing. Its description is pending until additional specimens are found. Distribution. French Guiana.
Male unknown.
Distribution. Western United States (Colorado). Chambers (1875) stated "Spanish Bar", now Fall River in Larimer County, Colorado, as the collecting locality. On the label of the holotype of P. luteella, "Kentucky" was given as collecting locality with strikethrough mark indicating that the locality is not correct.
Remarks. The forewing pattern and the female genital morphology of Eurynome luteella suggest that it is not congeneric with Philonome. Its forewing pattern is similar to some species of Argyresthia, especially A. cupressella Walsingham, 1891, andA. freyella Walsingham, 1891. The female genitalia of E. luteella include a denticulate signum of which the shape is typical for Argyresthia. This species, consequently, has been reassigned to Argyresthia.
Despite the strong support from molecular data, the tineid association of Philonome has never been addressed with morphological studies. Among the morphological characters associating Philonome with Tineidae are the reduced, naked haustellum with unassociated galeae, 5-segmented maxillary palpi, and vein Rs 4 terminating on costa before the forewing apex. It now appears that Philonome is most allied to the tineid subfamily Hieroxestinae also on the basis of morphological similarities. These include the wedge-shaped head (lateral view), vestiture of head partially consisting of appressed, laminate scales, and elongate scape without pecten. Previous association of this genus with Bucculatrix and Lyonetiidae was most likely decided largely by the presence of the broadly scaled antennal scape which forms an eyecap, a feature absent or poorly developed in Tineidae but typical for the latter two families. Eleven genera and 289 species are now recognized globally within Hieroxestinae, with 180 species assigned to Opogona (Robinson 2009). Within this subfamily, Philonome appears most similar morphologically to Oinophila Stephens, 1848, a holarctic genus currently restricted to two species. In particular, the head vestiture of both genera share unusual specializations not observed in other Hieroxestinae. The adult heads of Hieroxestinae typically possess a smooth, broad scaled frons and occiput, and a rough vertex consisting of a tuft of erect, piliform scales. The heads of Philonome and Oinophila are unusual in having the piliform scales of the vertex divided by a narrow, transverse band of broad, flat scales extending between the bases of the antennae (Davis 1978;Robinson and Tuck 1997). Philonome and Oinophila also possess similar wing venation, with the R vein lacking in the forewing and Rs with 4 branches. The heads of both genera possess a relatively raised vertex, and the rudimentary mandibles are better developed than in other genera of the subfamily. The antennal scape of Oinophila differs from that of Philonome in being more slender, smoothly scaled, and not formed into an eyecap. The female genitalia of Philonome differ from other known Hieroxestinae by lacking a signum in the corpus bursae.
Despite some possible synapomorphies between Philonome and the Hieroxestinae, we find them insufficient for a final taxonomic placement, and therefore leave the genus unplaced in Tineidae. Figure 71 shows a neighbor joining tree based on the DNA barcode sequences for 12 individuals of Philonome available at BOLD systems (www.barcodinglife.org). The resulting tree and the distance matrix (Table 3) indicate the presence of seven unique taxonomic units which can be assigned to the separate Barcode Index Numbers (BINs: Ratnasingham and Hebert 2013). These include two previously known species of Philonome; P. clemensella (five individuals) and P. euryarga (one individual); four species described in this paper, P. albivittata, P. curvilineata, P. kawakitai, and P. lambdagrapha (all except P. albivittata based on singleton); and one species (Fig. 14: CLV105310) from French Guiana which cannot be named due to the loss of the abdomen.

DNA barcoding
DNA barcodes of the seven species analysed are very distinctive (Fig. 71, Table 1). Indeed, DNA barcodes show high levels of interspecific genetic distance (Table 3). All species analysed show distinct DNA barcodes with a minimum interspecific pairwise genetic distance of 6.9% among all species. The maximum intraspecific genetic variation ranged from 0.9 to 0.3, much lower than interspecific distances, suggesting the existence of a barcode gap although current intraspecific sampling is too limited. Jean-François Landry of the Department of Entomology, Agriculture and Agri-Food Canada/Agriculture, Ottawa, kindly provided specimen data and allowed us to use four unpublished barcodes of P. clemensella. We would like to thank the editor Erik van Nieukerken and two anonymous reviewers for their valuable comments on our manuscript. Field work in French Guiana was funded by the CNRS program 'Amazonie', Nouragues research grants 2009 and 2010 to C.L.V. Funding for DNA barcoding was provided by the government of Canada through Genome Canada and the Ontario Genomics Institute in support of the International Barcode of Life project, and by NSERC. The first author especially appreciates the financial support from the Peter Buck Postdoctoral Fellowship (2013-2015), Smithsonian Institution.