Research Article |
Corresponding author: Donald R. Davis ( davisd@si.edu ) Academic editor: Erik J. van Nieukerken
© 2015 Jae-Cheon Sohn, Donald R. Davis, Carlos Lopez-Vaamonde.
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:
Sohn J-C, Davis DR, Lopez-Vaamonde C (2015) Revision of the genus Philonome Chambers and its proposed reassignment to the family Tineidae (Lepidoptera, Tineoidea). ZooKeys 494: 69-106. https://doi.org/10.3897/zookeys.494.8748
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The New World genus Philonome Chambers, 1874 is revised. This genus comprises twelve species, seven of which are described as new: two species, P. nigrescens sp. n. and P. wielgusi sp. n., from the United States; four species, P. albivittata sp. n., P. curvilineata sp. n., P. kawakitai sp. n., and P. lambdagrapha sp. n., from French Guiana; and one species, P. penerivifera sp. n., from Brazil. Lectotypes are designated for Philonome clemensella Chambers, 1874 and P. 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.
Argyresthia, Argyresthiidae, Bucculatricidae, COI, DNA barcoding, Eurynome, Lyonetiidae, new species, New World
The monobasic genus Philonome was proposed by Chambers in
Maximum likelihood phylogeny of Tineidae s. l. extracted from
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
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 (
Pinned specimens from five institutional collections were examined. The abbreviations of these depositories are as follows:
BMNH Natural History Museum (formerly British Museum of Natural History), London, UK;
MCZ Museum of Comparative Zoology, Harvard University, Cambridge, USA;
MSU Mississippi Entomological Museum, Mississippi State University, Starkville, Mississippi, USA;
USNM National Museum of Natural History (formerly United States Museum of Natural History), Washington DC, USA;
VOB Vitor O. Becker, Instituto Uiraçu, Camacan, Brazil.
Other abbreviations include:
ex. example, the specimens whose sex cannot be determined;
Co. county;
GSN genitalia slide number;
WSN wing slide number.
Selected specimens were dissected for genitalia and abdominal structures, following
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
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/).
Species | Process ID | Sample ID | Country | Accession number (NCBI, GENBANK) |
---|---|---|---|---|
Philonome clemensella | MEC875-04 | jflandry0875 | Canada | GU096008 |
Philonome clemensella | MNAL543-10 | CNCLEP00035968 | Canada | KP696787 |
Philonome clemensella | MNAI712-09 | CNCLEP00038457 | Canada | GU692620 |
Philonome clemensella | MNAI218-09 | CNCLEP00042501 | USA | GU693088 |
Philonome clemensella | MNAI241-09 | CNCLEP00042524 | USA | GU693064 |
Philonome curvilineata | LNOUA586-10 | CLV68110 | French Guiana | HQ571412 |
Philonome euryarga | LNOUA669-10 | CLV76410 | French Guiana | HQ571490 |
Philonome albivittata | LNOUA849-10 | CLV94410 | French Guiana | HQ571657 |
Philonome albivittata | LNOUA946-10 | CLV104110 | French Guiana | HQ571747 |
Philonome lambdagrapha | LNOUA928-10 | CLV102310 | French Guiana | HQ571730 |
Philonome kawakitai | GRANO044-11 | AK0044 | French Guiana | HQ571758 |
Philonome sp. | LNOUA958-10 | CLV105310 | French Guiana | KM224529 |
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
Philonome Chambers, 1874: 96;
Type species: Philonome clemensella Chambers, 1874, by monotypy.
Phillonome [sic]:
Phyllonome [sic]:
Head (Fig.
Thorax: Foreleg epiphysis slender. Midfemur with apical tuft of elongate scales. Hind-tibia hairy dorsally. Forewing pattern elements (Fig.
Adults. 2 Philonome cuprescens, ♂ (3.1 mm), abbreviations: AS = apical spot; CF = costal fascia; DB = dorsal bar; LF = longitudinal fascia; SS = subapical spot; TP = tornal patch 3 P. wielgusi, ♂ holotype (2.8 mm) 4 P. nigrescens, ♂ holotype (2.8 mm) 5 P. clemensella, ♂ (4.0 mm) 6 P. lambdagrapha, ♂ holotype (3.0 mm) 7 P. curvilineata, ♂ holotype (2.8 mm) 8 P. euryarga, ♂ (2.7 mm) 9 P. albivittata, ♂ holotype (2.8 mm). (Forewing lengths in parentheses).
Adults. 10 P. penerivifera, ♀ paratype (3.6 mm) 11 P. kawakitai, ♀ holotype (3.8 mm) 12 P. rivifera, ♂ lectotype (3.4 mm) 13 P. rivifera, ♂ paralectotype (2.8 mm) 14 P. sp., CLV105310 (4.1 mm) 15 P. spectata, ♀ holotype (2.3 mm) 16 Argyresthia luteella, ♀ holotype (3.4 mm) 17 Elachista dasycara (= Eurynome albella),♀ holotype (4.0 mm). (Forewing lengths in parentheses).
Abdomen: Coremata on male sternum VIII present posterolaterally, short and stiff (P. albivittata sp. n., P. clemensella, P. euryarga, and P. wielgusi sp. n.), long and hair-like (P. lambdagrapha sp. n.) or absent (P. curvilineata sp. n. and P. rivifera).
Male genitalia: Paired processes (uncus, Fig.
Philonome, male genitalia. 21–25 P. cuprescens. 21 Ventral view of genital capsule 22 Lateral view 23 Lateral view of valva 24 phallus, lateral view 25 phallus, ventral view 26–30 P. wielgusi, male genitalia. 26 Ventral view of genital capsule 27 Lateral view 28 Ventral view of anellus and uncus 29 Lateral view of valva 30 Phallus. (Scale lengths in parentheses).
Female genitalia: Ovipositor telescopic with two primary segments; papillae anales semi-elliptical, setose; lamella antevaginalis conical or cylindrical; additional protrusion behind ostium bursae present (P. nigrescens and P. rivifera) or absent; ductus bursae slender; corpus bursae obovate or elliptical; signum absent.
Included species (arranged by the similarities in the forewing pattern and the male genitalia):
Philonome cuprescens Walsingham, 1914
Philonome wielgusi sp. n.
Philonome nigrescens sp. n.
Philonome clemensella Chambers, 1874
Philonome lambdagrapha sp. n.
Philonome curvilineata sp. n.
Philonome euryarga Meyrick, 1915
Philonome albivittata sp. n.
Philonome penerivifera sp. n.
Philonome kawakitai sp. n.
Philonome rivifera Meyrick, 1915
Philonome spectata Meyrick, 1920
Philonome sp.
1 | Forewing with longitudinal fascia extending near apex | 2 |
– | Forewing with longitudinal fascia not extending beyond discal cell | 5 |
2 | Forewing ground color brown or reddish brown | 3 |
– | Forewing ground color black | nigrescens sp. n. |
3 | Forewing with costal fascia curved | 4 |
– | Forewing with costal fascia straight | wielgusi sp. n. |
4 | Yellow lining along costal fascia of forewing narrow | clemensella Chambers |
– | Yellow lining along costal fascia of forewing broad | cuprescens Walsingham |
5 | Forewing with dorsal bar | 6 |
– | Forewing without dorsal bar | spectata Meyrick |
6 | Forewing with dorsal bar connected with longitudinal fascia | 7 |
– | Forewing with dorsal bar separated from longitudinal fascia | 8 |
7 | Fore- and hindwing fringes pale grayish orange | euryarga Meyrick |
– | Fore- and hindwing fringes brownish gray | albivittata sp. n. |
8 | Forewing with costal fascia | lambdagrapha sp. n. |
– | Forewing without costal fascia | rivifera Meyrick and allied species* |
* Four species, curvilineata sp. n., rivifera Meyrick, penerivifera sp. n., and kawakitai sp. n., are indistinguishable from one another based on external appearance; see Table
Comparison of three similar species of Philonome, P. curvilineata, P. penerivifera, and P. rivifera in the male genitalia.
Characters | curvilineata | penerivifera | rivifera |
---|---|---|---|
Apical area of valva | entire | emarginated | bifid |
Short spiniform setae on cucullus | present | absent | absent |
Saccular portion of valva | not separate | separate | separate |
Lateral area of vinculum | subtruncate | strongly protruding | slightly protruding |
Saccus | 1/4 as long as valva | 3/5 as long as valva | 1/2 as long as valva |
Philonome cuprescens Walsingham, 1914: 346;
(Fig.
Thorax: Patagium orange; tegula white, tinged with brown basally, yellowish brown subbasally; mesonotum brown with white transverse band at anterior 1/5, lined with yellowish brown anteriorly. Foreleg with coxa, femur and tibia dark brown on exterior surface, lustrous pale reddish brown on interior surface; tarsomeres pale brown dorsally, pale orange ventrally. Midleg with coxa lustrous pale orange; femur lustrous pale orange, tinged with dark grayish brown apically; tibia dark brown dorsally, pale orange ventrally; tarsomeres pale reddish brown dorsally, pale orange ventrally. Hindleg with coxa pale orange; femur pale grayish brown, tinged with pale orange ventrobasally; tibia brown dorsally, pale orange ventrally, with mixture of pale orange and pale brown piliform scales of tuft ventrally; tarsomeres pale reddish brown dorsally, pale orange ventrally. Forewing length 2.8–3.9 mm (n = 2), brown, intermixed with dark brown scales in postmedian area; longitudinal fascia white, closer to costa than to dorsum, accompanied with yellowish brown fascia anteriorly; costal fascia yellowish brown, curved to apex at the middle, accompanied with narrow, white line along lower margin in costal 1/2; dorsal bar white, curved in terminal 1/4, accompanied with yellowish brown spreading in dorsal area, almost connected with longitudinal fascia; subapical spot white, narrow, curved; apical spot white, suffused with reddish brown costally; tornal patch very small; fringe elongate scales dark brown, hairy scales dark grayish brown. Hindwing and fringe dark grayish brown.
Abdomen: Terga lustrous, dark reddish brown; sterna lustrous, pale yellow.
Male genitalia (Figs
Female unknown.
Holotype: male, “Type” [circular label with red borders], “Amula, 6000ft. GUERRERO MEXICO VIII 18 (H.H.Smith) (Gdm. Slvn) 66776”, “Walsingham Collection, 1910-427”, “Philonome ♂ cuprescens Wlsm. Biol. C. Am. Lep. Het. 4. p346, 1914 TYPE ♂ descr” [label with black marginal lines], BMNH. Paratypes: Same data as holotype: 1♂, 1 ex. [hindwing & abdomen missing], Type no. 66778 & 66779, [GSN] USNM 34210 (♂), USNM.
Mexico: Same locality as holotype: 1♂, 2 ex., 18 September [no year] (HH Smith), BMNH.
Mexico (Guerrero).
This species is similar to P. clemensella in external appearance but can be distinguished from the latter in having shorter longitudinal fascia and straight costal fascia on the forewing. In the male genitalia, the lateral processes on the uncus are larger in P. wielgusi than in P. clemensella and the valvae are not divided at all in P. wielgusi.
(Fig.
Thorax: Patagium yellowish brown. Tegula dark yellowish brown on basal 1/4, pale grayish brown on distal 3/4, paler distad. Mesonotum lustrous white on anterior 1/3, dark grayish brown on posterior 2/3. Foreleg with coxa dark grayish brown laterally, silvery gray mesally; femur and tibia dark grayish brown, paler ventrad; tarsomere I to II pale orange, with narrow dark brownish gray patch dorsally, the remaining tarsomeres entirely dark grayish brown. Midleg with coxa and tibia dark grayish brown laterally, lustrous pale orange mesally; tibia dark brownish gray dorsally, lustrous pale orange ventrally; tarsi dark brownish gray. Hindleg with coxa and femur lustrous, pale orange; tibia lustrous, yellowish gray, sparsely hairy dorsally, with dense spiniform setae ventrally; tarsi dark brownish gray. Forewing length 2.3–3.0 mm (n = 8), brown, dorsum dark brownish gray basally; longitudinal fascia, white, extending from base to basal 1/3 of forewing, costal fascia white, at distal 2/5 of costa, oblique, adjacent to a slender dark brown line on anterior side; subapical and apical spots white; tornal patch white, triangular; dorsal bar white, oblique; fringe dark orange, each scale with dark brown tip. Hindwing and fringe gray.
Abdomen: Terga lustrous, gray; sterna lustrous, pale orange. Tergum VIII of male rectangular; sternum VIII subrectangular, broadly emarginated posteriorly, with oblique furrow laterally and short, stiff coremata posterolaterally.
Male genitalia (Figs
Female unknown.
Holotype: male, “ARIZONA: Cochise Co.: Sierra Vista 5131 Bannock 2 IX 1988”, “Attracted to (E, Z) – 3 13 ODDOH @ 1615-1730 hrs.”, “R. S. Wielgus Collector”, USNM. Paratypes (78♂): USA: Arizona: Cochise Co.: Chiricahua Mountains., Sunny Flat Campground: 1♂, 28 July 1989, B & JF Landry, CNC. Sierra Vista: 5131 Bannock, 10♂, 31 August 1988 (RS Wielgus), on pheromone trap; 18♂, 1 September 1988, [GSN] USNM 31056; 12♂, 2 September 1988, [GSN] USNM 29950; 2♂, 5 September 1988; 1♂, 6 September 1988; 7♂, 9 September 1988; 1♂, 10 September 1988; 1♂, 14 September 1988; 3♂, 15 September 1988; 8♂, 16 September 1988; 4♂, 17 September 1988; 7♂, 18 September 1988, USNM. Graham Co.: Pinaleno Mountains: Wet Canyon: 3♂, 22 July 1989, B & JF Landry, CNC.
Southwestern United States (Arizona).
The species name is a patronym in honor of Mr. Ronald S. Wielgus, who collected nearly the entire type series.
As reported by the collector, Ronald Wielgus, and indicated on specimen labels, nearly all moths were collected in the late afternoon, between 16:15 and 17:30 hours. All 157 adults collected thus far are males.
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.
(Fig.
Thorax: Patagium white on distal half, dark brown on basal half; tegula white, tinged with dark brown basally; mesonotum dark brown with coppery luster. Foreleg with coxa to tarsomeres lustrous orange-white, narrowly tinged with gray dorsally. Midleg with coxa to tibia lustrous orange-white; femur with broad pale reddish brown patch dorsally; tibia broadly dark gray dorsally; tarsomeres dark gray, paler ventrally. Hindleg with coxa lustrous pale orange, tinged with brown basally; femur lustrous pale orange; tibia and tarsomeres dark gray dorsally, lustrous pale orange ventrally; tibia spinose dorsally, with hair tufts ventrally. Forewing length 2.1–3.2 mm (n = 7), dark brown with coppery luster; longitudinal fascia white, extending from base to basal 1/3 of forewing; costal fascia white, straight, broadened at costa; dorsal bar white; subapical, apical and tornal spots white; fringe gray. Hindwing gray, paler to base; fringe gray.
Abdomen: Terga lustrous, dark grayish brown; sterna lustrous, yellowish gray ventrally.
Male genitalia (Figs
Female genitalia (Figs
Holotype: male, “ARIZONA: Cochise Co. Sierra Vista 5131 Bannock 18 IX 1988”, “R Wielgus Collector”, “Attracted to 1988 Farchan (Z, Z)-3, 13 ODDA@1620hrs. in pheromone trap” [hand-written], USNM. Paratypes (8♂, 1♀): USA: Arizona: Cochise Co.: Same locality as holotype: 1♂, 13 May 1988, attracted to pheromone trap; 3♂, 28 August 1988 (R S Wielgus), attracted to pheromone trap; 1♂, 17 September 1988, all USNM. Pima Co.: Station Catalina: 1♂, [“iss”] 13 June 1913 (Hopk.), “from trunk of evergreen Oak”, [GSN] USNM 16406, USNM. Santa Cruz Co.: Peña Blanca Campground: 2♂, 22 August 1988, attracted to pheromone trap, [WSN] USNM wing 29949, USNM. New Mexico: Grant Co.: Silver City: 1♀, 5 June 1974 (PM Jump), [GSN] USNM 34355, USNM.
The species name is derived from the Latin verb ‘nigrescere’ meaning “verging on black” and refers to the black ground color of the forewing of this species.
Southwestern United States (Arizona, New Mexico).
Philonome clemensella:
Philonome staintonella:
(Fig.
Thorax: Patagium reddish brown; tegula pale orange-white, suffused with reddish brown basally; mesonotum pale orange-white. Foreleg with coxa lustrous orange-white, tinged with gray dorsobasally; femur and tibia dark brownish gray dorsally, lustrous orange-white ventrally; tarsi orange-white, lustrous ventrally. Midleg with coxa lustrous orange-white; femur to tarsi pale orange dorsally, lustrous orange-white. Hindleg with coxa to tarsi lustrous orange-white; femur narrowly tinged with pale orange dorsally; tibia spinose dorsally, with hair-tufts ventrally. Forewing length 2.8–4.4 mm (n = 70), reddish brown; longitudinal fascia, white from base to the middle of forewing, often connected with white dorsal bar; costal fascia at distal 1/3 of costa, white, terminal 1/3 curved to apex, accompanying a row of dark brown scales caudad; subapical spot orange-white; tornal patch white, semicircular, borders blurred; elongate scales on apex, terminal 1/3 of costa, termen, with dark brown tips; fringe orange-white. Hindwing and fringe lustrous pale gray.
Abdomen: Terga lustrous, yellowish gray; sterna lustrous orange-white. Male tergum VIII trapezoidal; male sternum VIII subrectangular, broadly emarginated posteriorly, with oblique furrow laterally and short, stiff coremata posterolaterally.
Male genitalia (Figs
Female genitalia (Figs
Lectotype (designated here): male, “48” [hand-written], “Type No. 522 U.S.N.M.” [red label], “Philonome clemmensella [sic] K[entuck]y. 5961Lis1 Cham[bers]” [hand-written], USNM. Paralectotypes: USA: Kentucky: 4♀, 3 ex., June 14 [no year] (Chambers), Type no. 1311, MCZ.
Canada: Ontario: Ottawa-Careton, Dunrobin: 1♀, 9 July 2007, CNCLEP00035968; 28 July, 2007, CNCLEP00038457, (L Scott). Quebec: 2♂, 21 July 2004, CNCLEP00006545, (JF Landry), CNC. USA: Alabama: Monroe Co.: Haines Island Park (31°43'23"N, 87°28'10"W): 2♂, 2♀, 26–27 May 1995 (R Brown, J MacGown & D Pollock), MSU. District of Columbia: Unspecified locality: 1♂, no date (Fernald); 1♂, 28 June 1885 (Fernald), “on oak”; 1♀, 21 June 1886 (Fernald); 1♂, 11 July 1896, “Hickory”; 3♀, 2–4 June 1897, “from Linden”; 1♀, September 1953, USNM. Florida: Pinellas Co.: Dunedin: Hammock Park: 1♂, 22 April 1987 (LC Dow), [GSN] USNM 96414, USNM. Kentucky: No specified locality: 1♂, no date & collector, USNM. Illinois: Macon Co.: Decatur: 1♂, 8–15 June [no year], USNM. Putnam Co.: 1♀, 30 June 1976 (MO Glenn), USNM. Maryland: Montgomery Co.: Takoma Park: 1♂, 1♀, 8 July 1986 (WE Steiner); 1♂, 7 July 1986, USNM. Wicomico Co.: 1km SW Sharptown at Plum Creek: 1♀, 12 July 1986 (JM Hill et al.), USNM. Massachusetts: Dukes Co.: Martha’s Vineyard: 1♂, 13 July [no year] (FM Jones); 1 ex., 29 July [no year], all USNM. New Jersey: Burlington Co.: Medford: Lake Pine: 1♂, 13 July 1974 (DC Rentz), USNM. Essex Co.: Caldwell: 3♂, 2♀, 8 July 1900 (WD Kearfott), USNM. Essex County Park: 1♀, 20 May [no year] (WD Kearfott); 1♂, 7 July [no year], GSN: USNM 29977; 1♂, 12 July 1901; 1♂, 15 July [no year], USNM. Montclair: 1♀, 10 July [no year] (WD Kearfott), USNM; 1♂, 18 July [no year] (WD Kearfott), USNM. New York: Tompkins Co.: Ithaca: Six Mile Creek: 1♀, 23 July 1960 (RW Hodges), USNM. Unspecified locality: 1♀, “4971/ WLSM. 1906” (Beutenmueller), USNM. North Carolina: Craven Co.: Cherrypoint: 1♀, 3 July 1961 (SS Nicolay); 1♀, 12 July 1961; 1♀, 21 July 1961, all USNM. Harnett Co.: Spout Springs: 1♂, 25 August 1984 (WE Steiner et al.), USNM. Ohio: Hamilton Co.: Cincinnati, 1 ex., 23 June 1906 (A Braun); 1♂, 1♀, 27–28 June 1906, [GSN] USNM 16405 (♂); 1♀, 24 July 1907; 1♂, 3 August 1907; 1♂, 16 June 1908, all USNM. PENNSYLVANIA: Adams Co.: Arendtsville: 5♂, 6 July 1921 (SW Frost), USNM, GSN: USNM 29575. Allegheny Co.: Oak Station: 1♂, 6 July 1907 (F Marloff); 1♂, 11 July 1907, all USNM. Beaver Co.: New Brighton: 2♂, 11 July 1907 (Merrick Museum); 1♂, 23 July 1907; 1♂, 2♀, 26 July 1907, [GSN] USNM 34213 (♀), USNM. South Carolina: Charleston Co.: McClellanville: Wedge Plantation: 1♀, 11 May 1981 (RW Hodges), USNM, GSN: USNM 34212. Tennessee: Cocke Co.: Great Smoky Mt. National Park: Foothills Parkway (35°48'59"N, 83°13'11"W): 3♂, 1♀, 9 June 2002 (RL Brown & SM Lee), MSU. Texas: Harris Co.: 1♂, 20 May 1984 (Bellaire), [GSN] USNM 96415; 1♂, 2 April 1986, all USNM. Virginia: Fairfax Co.: 1km E Fairfax City: 1♂, 9 July 2005 (J Brown), USNM. Unspecified locality: 1♀, 27 June 1886, USNM. West Virginia: Morgan Co.: Sleepy Creek Forest: 2♂, 1 July 2010 (J Glaser); 1♂, 16 July 2011; 1♂, 19 July 2011; 1♂, 21 July 2011, USNM.
Eastern Canada and the United States west to Texas.
Hickory (Juglandaceae: Carya) and linden (Tiliaceae: possibly Tilia americana L.) (
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.
(Fig.
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 grayish 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.
Abdomen: Male tergum VIII and sternum VIII subquadrate; coremata piliform, as long as tergum VIII.
Male genitalia (Figs
Philonome, male genitalia. 37–39 P. lambdagrapha 37 Ventral view of genital capsule and phallus 38 Lateral view 39 Phallus 40–41 P. curvilineata. 40 Ventral view of genital capsule and phallus 41 Lateral view 42–46 P. euryarga 42 Ventral view of genital capsule 43 Ventral view 44 abdominal tergum VIII (caudal end directed upward) 45 Lateral view of phallus 46 Anterior base of phallus, ventral view.
Female unknown.
Holotype: male, “ID#: CLV102310 [red letters] French Guiana: [Régina, Nouragues Research Station] [Lt:4.1 Ln:52/] Carlos Lopez Vaamonde 23-Jan-2010 DNA Barcode LNOUA928-10 [green letters in blue row]”, “Nou68”, “Genitalia slide DRD ♂ USNM 34621” [green label], USNM.
French Guiana.
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).
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).
(Fig.
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.
Abdomen: Male tergum VIII rectangular; male sternum VIII rectangular, broadly emarginated posteriorly.
Male genitalia (Figs
Female unknown.
Holotype: male, “ID#: CLV68110 [red letters] French Guiana: [Régina, Nouragues Research Station] [Lt:4.1 Ln:52/] Carlos Lopez Vaamonde 20-Jan-2010 DNA Barcode LNOUA586-10 [green letters in blue row]”, “Nou37”, “Genitalia slide DRD ♂ USNM 34620” [green label], USNM.
French Guiana.
The species name, an adjective, is derived from the Latin words ‘curvus’ and ‘lineatus’, together meaning “curved line” and refers to the curved longitudinal fascia on the forewing of this new species.
Philonome euryarga Meyrick, 1915: 250.
(Fig.
Thorax: Patagium and tegulae white; mesonotum white in anterior 1/2, reddish brown in posterior 1/2, with a dark brown transverse band medially; mesoscutellum brownish gray. Foreleg lustrous yellowish white, with narrow brownish gray area dorsally. Midleg reddish orange dorsally, lustrous yellowish white ventrally. Hindleg pale orange dorsally, lustrous yellowish gray ventrally. Forewing length 2.7 mm (n=1), 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; elongate scales of fringe pale reddish brown, with dark brown tips; hairy scales of fringe pale yellowish gray. Hindwing pale grayish orange; fringe yellowish gray.
Abdomen: Male tergum VIII sclerotized, subtrapezoidal, narrower caudally, emarginated posteriorly, with dense pores on posterior 1/3 and long process posterolaterally; male sternum VIII subrectangular, with short coremata posterolaterally.
Male genitalia (Figs
Female genitalia not examined.
Holotype: female, “Holo-type” [round label with red borders], “Bartica, Brit[ish] Guiana. Parish. 2.13”, “euryarga Meyr.” [hand-written], “Philonome euryarga 1/1 Meyr[ick] E. Meyrick det. in Meyrick Coll.”, BMNH.
French Guiana: Régina: Nouragues Research Station (Lt: 4.1, Ln: 52): 1♂, 19 January 2010 (C. Lopez-Vaamonde), DNA Barcode LNOUA669-10, ID#: CLV76410, [GSN] USNM 34622, USNM.
Guyana and French Guiana.
This species is similar to another congener, P. euryarga Meyrick in overall external appearance, but differs from the latter in having darker hindwings. Their male genitalia possess several distinct differences including the tegumen with lateral projections in P. albivittata; the saccus present only in P. euryarga; and in the form of the costal portion of valva (Figs
(Fig.
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.
Abdomen: Male tergum VIII rectangular; male sternum VIII subrectangular, with oblique furrow and short coremata laterally.
Male genitalia (Figs
Philonome, male genitalia. 47–48 P. albivittata 47 Ventral view of genital capsule and phallus 48 Lateral view 49–52 P. penerivifera 49 Ventral view of genital capsule and phallus 50 Lateral view, with ventral detail of uncus 51 Lateral view of valva 52 Phallus 53–56 P. rivifera. 53 Ventral view of genital capsule and phallus 54 Lateral view 55 Lateral view of valva 56 Phallus.
Female unknown.
Holotype: male, “ID#: CLV10410 [red letters] French Guiana: [Régina, Nouragues Research Station] [Lt:4.1 Ln:52/] Carlos Lopez Vaamonde 16-Jan-2010 DNA Barcode LNOUA009 [sic: 946] -10 [green letters in blue row]”, “Genitalia slide DRD ♂ USNM 34623” [green label], USNM. Paratype: French Guiana: Régina: Nouragues Research Station (Lt:4.1, Ln:52): 1♂, 20 January 2010 (C Lopez-Vaamonde), “ID#: CLV94410”, “DNA Barcode LNOUA849-10”, [GSN] 34625, USNM.
French Guiana.
The species name is derived from the Latin adjectives, ‘albus’ and ‘vittatus’, meaning “white” and “banded” respectively, and refers to the white longitudinal band on the forewing of this new species.
This species is indistinguishable from Philonome rivifera in external appearance. Both species can be clearly distinguished from each other by the male genitalia (Table
(Fig.
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.
Male genitalia (Figs
Female genitalia (Figs
Holotype: male, “Holo-Type” [circular label with red border], “Para Brazil Parish 6 -19.”, “Philonome rivifera 7/17 Meyr. E. Meyrick det. in Meyrick Coll.” [old label attached before this study], “Meyrick Coll. B.M. 1988-290.”, “B.M. ♂ Genitalia slide No. 32828”. Paratypes (1♂, 4♀, 1 ex.): Brazil: Amazonas: Munaos [= Manaus], 2♀, “11.19” (Parish), BMNH. Tefé, 1 ex., “1.20” (Parish), BMNH. Federal District: Planaltina (15°35’S, 47°42’W, alt. 1000m): 1♀, 3 May 1984 (VO Becker), “BECKER 56394”, VOB; 1♀, 15 June 1985, “BECKER 57636”, [GSN] USNM 34615, USNM. Pará: Óbidos, 1♂, “9.19” (Parish), BMNH.
Brazil (Amazonas, Federal District, Pará).
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.
The holotype and three paratypes of Philonome penerivifera in the BMNH collection were misidentified as P. rivifera by Edward Meyrick.
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.
(Fig.
Thorax: Scales of patagium pale orange, with dark brown tips; tegula pale brown on basal 1/3, pale orange on distal 2/3; mesonotum pale orange, sparsely intermixed with dark brown-tipped scales. Fore- and midlegs with coxa lustrous pale orange; femur, tibia, and tarsomeres dark brown mesally, lustrous yellowish white laterally. Hindleg with coxa pale orange; femur and tibia pale yellowish grayish dorsally, pale yellow ventrally; tarsomeres pale grayish yellow. Forewing length 3.8 mm (n = 2), reddish brown, paler along dorsal area; costal area pale orange, intermixed with dark brown scales densely on basal 1/3 and sparsely on distal 1/3; longitudinal fascia continuous to near termen, convex and narrowed at distal 1/3, white, juxtaposed with slender black line along lower border; dorsal bar as triangular patch on anterior half, combined to longitudinal fascia and as slender, intermittent, black line on posterior half; subterminal line connecting distal 1/8 of costa and tornus, dark brown, intermittent; fringe pale brownish gray, with three dark brown, transverse lines. Hindwing dark brownish gray; fringe pale brownish gray.
Male unknown.
Female genitalia (Figs
Holotype: female, “FRENCH GUIANA: Nouragues Nature Reserve Nouragues Research Station Sep[tember]-07-2010 collected by light trapping”, “Genitalia slide DRD ♀ USNM 34652” [green label], USNM.
French Guiana.
The species name is a patronym in honor of Dr. Atsushi Kawakita who collected the holotype.
Philonome rivifera Meyrick, 1915: 251.
(Figs
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.
Abdomen: Terga grayish brown; sterna pale orange. Male tergum VIII rectangular; male sternum VIII subrectangular, broadly emarginated posteriorly; coremata absent.
Male genitalia (Figs
Female genitalia (Figs
Lectotype (designated here): male, “LECTO-TYPE” (round label with indigo boarders), “Bartica Brit[ish] Guiana Parish. 2.13”, “Meyrick Coll. B.M. 1938-290.”, “Philonome rivifera 10/17 Meyr[ick] E. Meyrick det. in Meyrick Coll.”, BMNH. Paralectotypes: Guyana: same data as lectotype: 1♂, 4♀, [GSN] BM 31892 (♂) & BM 32829 (♀), BMNH.
Guyana.
Philonome spectata Meyrick, 1920: 359;
(Fig.
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 terminal 1/4 of costa and on tornal area yellowish brown with dark brown tips. Hindwing lustrous, yellowish gray, paler to base; fringe pale yellowish gray.
Abdomen: Terga and sterna lustrous white.
Female genitalia not examined.
Holotype: female, “Holo-type” [round label with red borders], “Para Brazil Parish 7-19.”, “Meyrick Coll. B.M. 1938-290.”, “Philonome spectata 1/1 Meyr[ick] E. Meyrick det. in Meyrick Coll.”, BMNH.
Brazil (Pará).
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.
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.
French Guiana: Régina: Nouragues Research Station (Lt: 4.1, Ln: 52): 1♂, 24 January 2010 (C. Lopez-Vaamonde), DNA Barcode LNOUA958-10, ID#: CLV105310, USNM.
French Guiana.
Philonome luteella (Chambers) was originally the type species of Eurynome Chambers, 1875. The generic name was found to be preoccupied by Eurynome Leach, [1814] and was replaced with Busckia Dyar, 1903.
Phalaena goedartella Linnaeus, 1758, by subsequent designation by
Eurynome Chambers, 1875: 304. A junior homonym of Eurynome Leach, [1814] [Crustacea]. syn. n. Type species: Eurynome luteella Chambers, 1875, by monotypy.
Busckia Dyar, 1903: 563. An objective replacement name of Eurynome Chambers, 1875. syn. n.
Eurynome luteella Chambers, 1875: 304.
Busckia luteella (Chambers):
Philonome luteella (Chambers):
(Fig.
Thorax: Patagium pale saffron yellow (
Male unknown.
Female genitalia (Figs
Holotype: female, “Kentucky [sic] Chambers”, “Type 14964” [red label], “Eurynome luteella Chambers” [hand-written on folded paper], “Genitalia slide MCZ-L122 Prep. by JC Sohn” [label with black border lines].
Western United States (Colorado).
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, and A. 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.
Philonome has been associated frequently with Bucculatrix, since
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 Rs4 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 (
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
Kimura 2-parameter (K2P) distances (%) for barcode DNA sequences of the seven analyzed species in genus Philonome. Minimal pairwise distances between species are given for each species pair. Values in square brackets represent maximal intraspecific distances.
clemensella | curvilineata | euryarga | albivittata | lambdagrapha | sp. | kawakitai | |
---|---|---|---|---|---|---|---|
clemensella | [0.9] | ||||||
curvilineata | 6.9 | ||||||
euryarga | 11.5 | 13.3 | |||||
albivittata | 12 | 13.2 | 11.7 | [0.3] | |||
lambdagrapha | 10.7 | 12.4 | 11.7 | 11.2 | |||
sp. | 13 | 14.1 | 13.9 | 14.1 | 12.1 | ||
kawakitai | 12.2 | 12.4 | 11.4 | 12.8 | 12.1 | 7.9 |
DNA barcodes of the seven species analysed are very distinctive (Fig.
We would like to thank Kevin Tuck (retired) and Geoff Martin, both from the Natural History Museum, London, for allowing the first author to examine the museum collection under their responsibility. We are also grateful to David Adamski (Systematic Entomology Lab, US Department of Agriculture, Beltsville, Maryland) for checking some type specimens of Philonome and Atsushi Kawakita for donating one specimen for analysis. We are indebted to Young Sohn, Vichai Malikul, Donald Harvey, and Karolyn Darrow of the Department of Entomology, Smithsonian Institution, for their assistance preparing the illustrations, graphics, and plates used in this publication. We also thank colleagues at the Biodiversity Institute of Ontario, University of Guelph, Ontario, Canada for their assistance in the production of DNA barcodes. 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.