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ZOOTAXAISSN 1175-5326 (print edition)

ISSN 1175-5334 (online edition)Copyright © 2010 · Magnolia Press

Zootaxa 2693: 21–34 (2010) www.mapress.com/zootaxa/ Article

Key to the genera of Oxyptilini (Lepidoptera: Pterophoridae: Pterophorinae), with descriptions of two new genera

HELEN ALIPANAH1,2, ALIREZA SARI1, ALIMORAD SARAFRAZI2, CEES GIELIS3 & SHAHAB MANZARI2

1School of Biology, College of Science, University of Tehran, Tehran, Iran. E-mail:alipanah@khayam.ut.ac.ir; halipanah@gmail.com2Insect Taxonomy Research Department, Iranian Research Institute of Plant Protection, P. O. Box 1454, Tehran 19395, Iran3Mr. Haafkensstraat 36, 4128 CJ Lexmond, Netherlands

Abstract

Apoxyptilus, gen. nov., and Pseudoxyptilus, gen. nov., are described and illustrated, with Oxyptilus anthites Meyrick, 1936, and Oxyptilus secutor Meyrick, 1911, as their type-species, respectively. Apoxyptilus anthites (Meyrick), comb. nov., and Pseudoxyptilus secutor (Meyrick), comb. nov., are redescribed. An illustrated key to the genera of the tribe Oxyptilini is provided.

Key words: Lepidoptera, Pterophoridae, Pterophorinae, Oxyptilini, new genera, new combinations, identification key

Introduction

Pterophoridae are a well-defined family of microlepidoptera characterized by deeply cleft wings in most species, a T-shaped resting posture of adults, and venous scales on the underside of the hindwing (Gielis 1993). The family is worldwide in distribution and includes approximately 1,189 described species assigned to 90 genera and four subfamilies (Gielis 2003, 2006, 2008, 2009; Arenberger 2006; Alipanah & Ustjuzhanin 2005, 2006; Altermatt 2008, Gielis & Karsholt 2009). The largest subfamily, Pterophorinae, presently is divided into six tribes, i.e., Tetraschalini, Platyptiliini, Exelastini, Oxyptilini, Oidaematophorini, and Pterophorini (Gielis 2003).

As previously defined, Oxyptilini is represented by 15 genera and 113 described species (Zagulajev 2002; Gielis 2003, 2006, 2008, 2009; Arenberger 2006). Alipanah et al. (in press) recently transferred Antarchesfrom Exelastini to Oxyptilini and confirmed the assignment of Sphenarches to Oxyptilini as proposed by Arenberger (2002). They also revealed that only five species of the many species currently assigned to Oxyptilus, i.e., O. pilosellae (Zeller), O. parvidactyla (Haworth), O. chrysodactyla ([Denis & Schiffermüller]), O. ericetorum (Stainton), and O. delawaricus Zeller, are similar to each other (and the type species of the genus) in both external and genitalia characteristics. These species, which were found to represent a sister group to Crombrugghia spp., were assumed to be the only species correctly assigned to Oxyptilus, and they are the only ones considered as such in the key presented below. Alipanah et al. (in press) concluded that the total number of described species in Oxyptilini is 118. The monophyly of the tribe is supported by the presence of symmetrical valvae in the male genitalia, the presence of two longitudinal rows of venous scales on the underside of the hindwing, an extension of the venulae of sternite II to the end of the sternite in the male abdomen, and the presence of an inverse, nearly V-shaped sclerotized structure at the distal half of sternite II in the male (Alipanah et al. in press).

There have been few published generic keys to Pterophoridae. In the first, provided by Meyrick (1886), subfamilies were not recognized. Meyrick (1886) classified the family into 16 genera that were mostly assigned to subfamilies during the second half of the 19th century. Meyrick’s classification was based mainly on the fore- and hindwing characteristics including venation. A similar key, presented by Fletcher (1909),

Accepted by J. Brown: 22 Oct. 2010; published: 1 Dec. 2010 21

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included 13 genera. All of the subsequently published keys have focused primarily on regional faunas (i.e., Yano 1963a, b; Bigot 1966; Nel 1986; Landry & Gielis 1992; Gielis 1991, 1996, 2006; Arenberger 2002). This approach is problematic because it can result in an unstable system in which new species are constantly being described and assigned to genera that traditionally have been recognized regionally. A further consequence of the regional classification has been a tendency towards describing new genera on an ad hocbasis, where new species are identified that do not fit the existing system (for the region). Many generic concepts that have originated in this way require review and/or revision. The lack of an inclusive generic key in each of the six known tribes of the subfamily Pterophorinae is also noticeable.

So far, no inclusive identification key for the genera of Oxyptilini including all the genera previously assigned to Trichoptilini and Oxyptilini (Bigot et al. 1998; Arenberger 2002) has been published. To classify the extant, described genera, we here present a new generic classification without including most of the characters that were described in detail in the original generic descriptions. Careful examination of the available material resulted in the discovery of several highly informative characters that, in part, are used here in the construction of our new classification; some of them demonstrate the minimal value of homoplasy (i.e., position of the lateral concavity of abdominal tergite II of male and cone-shaped folded area(s) at the posterior of corpus bursae). We believe that the characters used in the key provided below easily separate the genera from each other.

The examination of the available specimens of Oxyptilini in the National Centre of Biodiversity, Naturalis, Leiden, also revealed that two species previously assigned to Oxyptilus, i.e., O. anthites Meyrick and O. secutor Meyrick (recently proposed by Alipanah et al. (in press) to be senior synonym of O. variegatus(Meyrick)), are different from other members of the genus, both in external features and genitalia characteristics. Further examination of adults showed that these species require placement in two new monotypic genera that are herewith proposed and described. Support for these new genera is provided in the phylogenetic analysis of the tribe presented by Alipanah et al. (in press).

Material and methods

This study is based on the examination of specimens of pinned adult moths representing the genera previously assigned to Oxyptilini. Most of the material is from the collection of Cees Gielis, which is housed in the National Centre of Biodiversity, Naturalis, Leiden. Other depositories are the ZMUC (Zoological Museum of the University of Copenhagen, Denmark), ZMA (Zoölogisch Museum, Amsterdam, The Netherlands), HMIM (Hayk Mirzayans Insect Museum, Iran), and the personal collections of Ernst Arenberger (Austria), Jaroslaw Buszko (Poland), Leif Aarvik (Norway), and Peter Ustjuzhanin (Russia).

Dissection and slide mounting methods for genitalia and wings were based on those of Clarke (1941) and Robinson (1976), except preparations were stained with chlorazol black and eosin, respectively, and mounted in Euparal. Characters of the genitalia, wings, and the whole body were examined using a stereomicroscope (maximum magnification 128×). Wing venation terminology follows Comstock and Needham (1918) and other morphological and genitalia terms follow Nielsen and Common (1991), Scoble (1992), Klots (1970), Gielis (1993, 1996) and Kristensen (2003).

Results

Key to genera of Oxyptilini

The key assumes some knowledge of wing shape, pattern, and venation, as well as both male and female genital characters. Many of the features are indicated with arrows in the figures.

1. First lobe of forewing narrow, gradually attenuate towards the apex and pointed apically (Fig. 1A) ......................... 2- First lobe of forewing nearly uniform (or slightly widened) medially (lanceolate), with or without termen (Figs 1B,

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C)................................................................................................................................................................................... 72. Individual longitudinal white scales present at base of dorsum of third lobe of hindwing (Figs 2A, B, E–H)............ 3- Individual white scales absent at base of dorsum of third lobe of hindwing (Fig. 2D)................................. Buckleria3. Individual longitudinal white scales at base of dorsum of third lobe of hindwing nearly as long as (or only slightly

shorter than) the longest dark scale tooth/teeth of dorsum (Figs 2A, E–H).................................................................. 4- Most individual longitudinal white scales at base of dorsum of third lobe of hindwing significantly longer than the

longest dark scale tooth/teeth of dorsum (Fig. 2B)....................................................................................................... 64. Vein Cu1 present in forewing ........................................................................................................................................ 5- Vein Cu1 absent in forewing............................................................................................................................. Stangeia5. In the forewing, vein R5 present, veins R2 and Cu2 absent; ventral scale brush of second segment of labial palpus

extending along third segment to three-fourths of its length (Fig. 3A) ........................................................... Dejongia- In the forewing, vein R5 absent, veins R2 and Cu2 present; ventral scale brush of second segment of labial palpus

extending along third segment to half of its length (Fig. 3B) ......................................................................Stenodacma6. Forewing with two radial veins ...................................................................................................................Trichoptilus- Forewing with more than two radial veins ............................................................................................ .Megalorhipida7. Anal angle of second lobe of forewing without pronounced wide group of elongate dark linear fringe scales (Figs

4A, B)............................................................................................................................................................................ 8- Anal angle of second lobe of forewing with pronounced wide group of elongate dark linear fringe scales (Figs 2A,

C, E–H) ....................................................................................................................................................................... 108. Surface of forewing lobes with longitudinal white lines (Fig. 4A) ......................................................... .Intercapperia- Surface of forewing lobes without longitudinal white lines ......................................................................................... 99. Pronounced dark scale teeth absent in both costa and dorsum of third hindwing lobe (Fig. 4B) .............. .Eucapperia- Pronounced dark scale teeth present only at dorsum of third hindwing lobe (Fig. 7A) ............................. .Apoxyptilus10. Tibia of legs without rough scales at base of spurs, or with very short rough scales .................................... .Tomotilus- Tibia of legs with pronounced long rough scales at base of spurs............................................................................. .1111. Forewing with vein R1 present; lateral concavity of abdominal tergite II of male situated almost at 0.45–0.55 length

of tergite II (Fig. 5A)................................................................................................................................................... 12- Forewing without vein R1; lateral concavity of abdominal tergite II of male at 0.65–0.75 length of tergite II (Fig. 5B)

.................................................................................................................................................................................... 1612. Second segment of labial palpus without ventral scale brush .................................................................................... 13- Second segment of labial palpus with dense ventral scale brush extending along third segment to three-fourths (or

more) of its length (Fig. 3C) ....................................................................................................................................... 1513. Uncus fused with tegumen............................................................................................................................. Antarches- Uncus articulated with tegumen ................................................................................................................................ .1414. Pronounced dark scale teeth present at both costa and dorsum of third hindwing lobe; in the female genitalia, poste-

rior margin of lamella antevaginalis without sclerotized area (Fig. 6II, A); with cone-shaped folded area(s) at the posterior end of corpus bursae (Fig. 6II, C)................................................................................................Sphenarches

- Pronounced dark scale teeth present only at dorsum of third hindwing lobe; in the female genitalia, posterior margin of lamella antevaginalis with a pair of large symmetrical sclerotized triangular structures (Figs 6II, B, 8I, J); no cone-shaped folded area(s) at the posterior end of corpus bursae (Fig. 8J).......................................... Pseudoxyptilus

15. Pronounced dark scale teeth present at both costa and dorsum of third hindwing lobe; distance between apical and subapical (subterminal) scale teeth at dorsum conspicuously less than length of subapical group (Fig. 2E) .........................................................................................................................................................................................Oxyptilus

- Pronounced dark scale teeth present only at dorsum of third lobe of hindwing; distance between apical and subapical scale teeth at dorsum slightly more than length of subapical group (Fig. 2F) ........................................ Crombrugghia

16. Pronounced individual white scales at base of dorsum of third hindwing lobe absent; tegula brown, without creamy-white scales caudo-dorsally; position of uncus when articulated with tegumen in front of tegumen, but hanging and not in the same direction with it (Fig. 6I, A); ductus bursae emerging from medio-lateral to basal part of the cone-shaped folded area at the posterior end of corpus bursae (Fig. 6II, D) ................................................................. Geina

- Pronounced individual white scales at base of dorsum of third hindwing lobe present; tegula pale brown, with creamy-white scales caudo-dorsally; position of uncus when articulated with tegumen in front of tegumen, in the same direction with it (Fig. 6I, B); ductus bursae emerging from near distal part of the cone-shaped folded area at the posterior end of corpus bursae (Fig. 6II, E) ................................................................................................................ 17

17. Single pronounced dark scale tooth of costal margin of third hindwing lobe apical or very close to apex (Fig. 2A) ............................................................................................................................................................................Capperia

- Single pronounced dark scale tooth of costal margin of third hindwing lobe subapical, located from basad of apex by slightly less than its own length (Fig. 2H) .................................................................................................................. 18

18. In the male genitalia, valva shaped as a regular paddle (Fig. 6I, C); maximum length of specialized eighth sternite/

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maximum length of the valva ≤ 0.45 ..........................................................................................................Procapperia- In the male genitalia, valva not shaped as a regular paddle (Fig. 6I, D); maximum length of specialized eighth stern-

ite/maximum length of the valva 0.55–0.70 .............................................................................................Paracapperia

FIGURE 1. Forewings. A, Megalorhipida leucodactylus. B, Geina didactyla. C, Eucapperia bullifera. Arrows indicate key characters.

FIGURE 2. Fore- and hindwings. A, Capperia fusca. B, Megalorhipida leucodactylus. C, Geina didactyla. D, Buckleria paludum. E, Oxyptilus pilosellae. F, Crombrugghia distans. G, Sphenarches anisodactyla. H, Paracapperia anatolicus. Arrows indicate specific characters.

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FIGURE 3. Head in lateral view. A, Dejongia lobidactylus. B, Stenodacma wahlbergi. C, Oxyptilus pilosellae. Arrows indicate specific characters.

Apoxyptilus, gen. nov.

Type species: Oxyptilus anthites Meyrick, 1936.

Diagnosis. Apoxyptilus is characterized by the absence of a pronounced ventral scale brush on the second segment of the labial palpus; a strongly oblique termen of the second lobe of the forewing; the distance between the subapical and apical scale teeth exceeding three times the length of the subapical tooth; male genitalia with the uncus reduced and the valva without a valvular lobe; and female genitalia with a cylindrical antrum and lacking a signum.

FIGURE 4. Fore- and hindwings. A, Intercapperia scindia. B, Eucapperia bullifera. Arrows indicate specific characters.

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FIGURE 5. First and second abdominal segments of males. A, Crombrugghia laetus. B, Capperia raptor. Arrows indicate key characters.

Description. Head. Frons and vertex smooth scaled. Labial palpus straight; second segment without pronounced ventral scale brush extending along third segment. Length of antenna ca. one-half length of forewing; dorsal surface ringed with black and white scales. Chaetosema absent. Thorax. Smooth scaled. Legs with pronounced rough scales at base of spurs. Forewing (Fig. 7A) with cleft positioned centrally; first lobe nearly lanceolate, pointed apically; second lobe decreasing in width approximately from mid-dorsum towards apex. Fringe yellowish white, at dorsum of first lobe with a narrow preapical group of rather long dark fringe bordered by whitish fringe. Hindwing (Fig. 7A) with first lobe tapering towards apex, nearly pointed apically; width of second lobe decreasing approximately from mid-dorsum or slightly beyond towards apex, with strongly oblique and slightly excavated termen. Fringe yellowish white with two dark scale teeth on dorsum of third lobe: subapical and apical; distance between these two scale teeth more than three times length of subapical one. Venous scales on underside of hindwing ferruginous brown, in two longitudinal rows. Forewing venation (Fig. 7B) with R1 absent; R2 and R5 free; R3 and R4 stalked; M1, M2, Cu1, and Cu2 present;

Cu1 branched from one-fourth length of M3 length; Cu2 branched from slightly beyond lower angle of discal

cell. Hindwing venation with Sc+R1 extending to middle of wing; M3, Cu1, and Cu2 present; Cu1 branched

from slightly beyond first cleft; third lobe of hindwing with one anal vein (An1). Abdomen. Lateral concavity

of tergite II of male at 0.65–0.75 length of tergite II; specialized eighth sternite of male unilobed. Male genitalia (Fig. 7C) with valvae symmetrical; nearly spoon-shaped; without costal processes and valvular lobes; cucullus undifferentiated. Uncus reduced. Gnathos arms absent. Anellus without arms. Phallus nearly straight, without conspicuous process. Female genitalia (Fig. 7D) with papillae anales setaceous on posterior margins. Apophysis posterioris slender throughout. Apophysis anterioris absent. Ostium and antrum exposed and positioned centrally. Antrum cylindrical, of a single plate. Ductus bursae narrow, parallel-sided. Corpus bursae lacking signum. Ductus seminalis arising from posterior end of corpus bursae.

Etymology. The generic name indicates that this genus not closely related to Oxyptilus.Remarks. As with Oxyptilus, Apoxyptilus is characterized by the posterior margin of the eighth

abdominal tergite covered by scales in the form of a V-shaped structure; gnathos arms absent; the phallus nearly straight; the ostium bursae exposed; and the ductus seminalis originating from the posterior end of the corpus bursae. Apoxyptilus differs from Oxyptilus in the shape of the second forewing lobe; forewing

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venation; position of the dark scale teeth on dorsum of the third lobe of the hindwing; absence of a ventral scale brush on the second segment of the labial palpus; absence of a transverse white band at the upper portion of the fronto-clypeus; pattern of the scales covering surface of tegula; lateral concavity of abdominal tergite II of male, which is at 0.65–0.75 length of tergite II; unsclerotized eighth tergite; unilobed specialized eighth sternite; reduced uncus; shape of the valva, anellus, and saccus; absence of a signum in the corpus bursae; and a cylindrical antrum. In addition, preliminary results of a phylogenetic analysis based on morphological data (see Alipanah et al. in press) separate it convincingly from Oxyptilus species. Apoxyptilus shares the absence of vein R1 in the forewing and stalked R3 and R4 with Capperia, Procapperia, Paracapperia, Intercapperia,

Tomotilus and Geina; the position of the lateral concavity of tergite II of the male with Capperia, Procapperia, Paracapperia, Intercapperia and Geina; the spoon-shaped valva with Paracapperia esuriensand all Capperia species except C. raptor and C. insomnis; the position of dark scale teeth on the dorsum of the third hindwing lobe with Trichoptilus and two Megalorhipida species, viz., M. angusta and M. pseudodefectalis; the unilobed, specialized eighth sternite with Stangeia, and the reduced uncus with Buckleria and Trichoptilus pygmaeus. It also differs from all other known species incorrectly assigned to Oxyptilus species in wing shape and genitalia characteristics.

FIGURE 6. Stylized drawings of some parts of the male (I) and female genitalia (II). I, A–B, uncus-tegumen complex; light grey areas refer to the uncus. I, C–D, valva. II, A–B, posterior end of the female genitalia including the seventh abdominal sternite, ostium bursae, antrum, antrum plate, ductus bursae, and sclerotized parts at the posterior margin of the lamella antevaginalis (dark grey). II, C–E, anterior end of the female genitalia including the corpus bursae, anterior side of ductus bursae and ductus seminalis.

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FIGURE 7. Apoxyptilus anthites (Meyrick) comb. nov. A, adult male. B, fore- and hindwing venation. C, male genitalia (ventral view) and phallus. D, female genitalia.

Apoxyptilus anthites (Meyrick), comb nov.

Material examined. 1 male, Kenya, Rift valley, Turi, 2000 m, 7.i.1999, leg. D. Agassiz; 1 male, Tanzania, Arumeru Distr., Usa river, 1170 m, 31.vii.1991, leg. L. Aarvik (coll. CG 21953).

Diagnosis. This is the single species currently assigned to this new genus. It is characterized by four diagnostic characters: creamy-white tegula; bilobate anellus; circular and flat tip of the spoon-shaped valva and arch-shaped saccus.

Redescription. The following information is added to the original description. Head. Frons smooth. Vertex with large scales, almost twice size of those of frons. Labial palpus smooth-scaled; length approximately twice vertical diameter of eye; terminal (third) segment nearly as long as the second. Thorax. Smooth scaled, with metallic sheen. Tegula nearly bean-shaped, covered with creamy-white scales throughout. Wingspan 9–11 mm; forewing (Fig. 7A) yellowish white; each lobe of forewing with two transverse white bands on upperside: a complete preapical line and a wide line separate from apex; a very narrow transverse blackish line at base of cleft. Hindwing (Fig. 7A) whitish; subapical dark scale tooth of third hindwing lobe narrow (≤ 8 scales); pronounced individual dark scales on dorsum of third hindwing lobe present. Length of costal margin nearly 1.5× length of dorsum margin. Other characteristics of the wings as

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described for the genus. Abdomen (Fig. 7A). Whitish; dorsally with ferruginous-brown, small scattered patches and some darker ones towards the posterior end. Maximum length of unilobed specialized eighth sternite to maximum length of valva less than 0.3. Semicircular eighth abdominal tergite of male not sclerotized, with a depression at posterior margin in the middle. Male genitalia (Fig. 7C) as described for the genus. Additional diagnostic characters include circular and flat tip of spoon-shaped valva, bilobate anellus, almost arch-shaped saccus and absence of cornuti. Female genitalia (Fig. 7D) as described for the genus. Ring-shaped and weakly sclerotized antrum plate and the very slightly sclerotized posterior end of lamella antevaginalis are additional diagnostic characters for this species.

Ecology. Larvae feed on buds of Dombeya emarginata (Malvaceae) (Meyrick 1936).Distribution. Apoxyptilus anthites is known from Uganda, Tanzania, Kenya, and South Africa (Meyrick

1936; Gielis 2003).

Pseudoxyptilus, gen. nov.

Type species: Oxyptilus secutor Meyrick, 1911.

Diagnosis. Pseudoxyptilus is characterized by the absence of a ventral scale brush on the second segment of the labial palpus; hindwing with double rows of dark scaled teeth on dorsum of third lobe: distance between subapical and apical tooth more than three times length of the latter; presence of a wide group of short, dark scales on costal margin of third hindwing lobe almost above the subapical dark scale tooth of dorsum; male genitalia with half-pyramid-shaped, apically bifurcate uncus and flat valva without valvular lobe; and female genitalia with a pair of sclerotized structures at the posterior end of lamella antevaginalis, a wide ductus bursae, and the absence of a signum.

Description. Head. Frons and vertex smooth scaled. Labial palpus upturned, smooth scaled, second segment without pronounced ventral scale brush along third segment. Length of antenna approximately one-half length of forewing; dorsal surface ringed with black and white scales. Chaetosema absent. Thorax. Smooth scaled. Legs with pronounced rough scales at base of spurs. Forewing (Fig. 8A) with cleft in middle or slightly beyond; first lobe nearly lanceolate, pointed apically; second lobe slightly widened distally, with an oblique excavated termen. Fringe greyish white; at dorsum of first lobe, a narrow preapical group of rather long dark fringe bordered by whitish fringe. Hindwing (Fig. 8A) with first lobe tapering to nearly pointed apex; width of second lobe decreasing from about mid-dorsum towards apex. Fringe greyish white with two dark scale teeth on dorsum of third lobe: subapical and apical; distance between these two groups slightly more than length of subterminal one; venous scales on underside of the wing ferruginous brown, in two longitudinal rows. Forewing venation (Fig. 8B) with R1 and R2 stalked proximad of cleft base; R3 and R4

stalked; R5 free; M1, M2, Cu1, and Cu2 present; Cu1 branched from 0.3 length of M3; Cu2 branched from beyond lower angle of discal cell. Hindwing with Sc+R1 extended to middle of wing; M3, Cu1, Cu2, and An1 present. Abdomen. Lateral concavity nearly at middle of tergite II in male; anterior half of sternite II with an inverse V-shaped sclerotized structure at posterior margin; eighth abdominal tergite almost rectangular (Fig. 8H), not sclerotized; specialized eighth sternite bilobed, with lobes connected to each other internally-laterally or slightly separated near tips; apex of each lobe relatively wide and rounded (Figs 8C, G); anterior margin of specialized eighth sternite lacking internal flap; ventral surface without conspicuous setal tufts. Male genitalia (Figs 8C–G) with valvae symmetrical, flat, narrowing towards base, without costal processes; valvular lobe absent; cucullus undifferentiated; basal sclerotized processes on outer surface of valvae almost semi-circular to nearly oval. Uncus above tegumen, half-pyramid-shaped with bifurcate tip. Tegumen unilobed. Gnathos arms absent. Anellus without arms, sclerotized throughout; an elongated rectangle in anterior view and in antero-lateral view as a nearly asymmetric cone that is basally extending entire length of rectangle. Saccus somewhat T-shaped. Phallus straight, bulbus ejaculatorius positioned antero-dorsally to the phallobase, with a fish-hook-shaped sclerite inside. Female genitalia (Figs 8I, J) with papillae anales setaceous on posterior margins. Apophysis posterioris slender throughout, nearly as long as length of sternite VIII. Apophysis anterioris absent. Ostium bursae exposed; ostium and antrum positioned centrally. Antrum wide, somewhat funnel-shaped. Antrum plate present, single. Posterior end of lamella antevaginalis with pair of symmetrical sclerotized structures. Ductus bursae wide. Corpus bursae without signum. Ductus seminalis origniating from posterior end of corpus bursae.

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FIGURE 8. Pseudoxyptilus secutor (Meyrick) comb. nov. A, adult male. B, forewing venation. C, male genitalia (ventral view). D, phallus. E, male genitalia (lateral view). F, saccus. G, specialized eighth sternite (ventral view). H, eighth abdominal tergite of male. I, posterior margin of the lamella antevaginalis, including antrum and antrum plate. J, female genitalia.

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Etymology. The generic name reflects the superficial similarity of this genus to Oxyptilus. Remarks. This genus differs from Oxyptilus in having a greater distance between the two dark scale teeth

on the dorsum of the third hindwing lobe; a rectangular eighth abdominal tergite of male; the shape of the specialized eighth sternite; the half-pyramid-shaped uncus; the unilobed tegumen; the position of the bulbus ejaculatorius in relation to the phallobase and the presence of a sclerotized structure within it; the shape of the anellus, saccus, and basal sclerotized process on outer surface of the valve; the absence of a signum in the corpus bursae; and the shape of the antrum, the antrum plate, and the sclerotized areas at the posterior end of lamella antevaginalis. Furthermore, the results of a phylogenetic analysis of Oxyptilini (Alipanah et al. in press) based on morphological data place it convincingly outside of Oxyptilus. This genus is closer to the Oxyptilus generic group than Trichoptilus. It shares a similar distance between the two dark scale teeth on the dorsum of the third hindwing lobe with Crombrugghia; the common stalks of R1 and R3 with R2 and R4 with Sphenarches (except for the position of common stalk of R1 and R2); the bilobed specialized eighth sternite whose lobes are separated near the tip with Capperia, Geina, and Sphenarches ontario; the T-shaped saccus, position of the bulbus ejaculatorius, and sclerotized structure within it with Capperia, Procapperia, Intercapperia, Eucapperia, Geina, and Sphenarches; the rectangular anellus with Capperia, Procapperia, Paracapperia, Intercapperia; the position of the uncus above the tegumen with most Trichoptilus and Megalorhipida species; semicircular to oval shape of the basal sclerotized process on outer surface of the valva with Dejongia, Trichoptilus, Megalorhipida; and the presence of a pair of sclerotized structures at posterior end of the lamella antevaginalis with Eucapperia.

Pseudoxyptilus secutor (Meyrick), comb nov.

Material examined. Holotype (male), Republic South Africa, Pretoria, 5.xii.1907, A.J.T Janse (gen. prep. 14976); 1 male, Republic South Africa, Natal, Spioenkop, NR 12.ii.1995, leg. H.W. V.D. Wolf (coll. CG 11042).

Diagnosis. Pseudoxyptilus is currently defined as monotypic, with the single species secutor. It is characterized by four diagnostic characters: flat valva with two distal angles; specialized eighth sternite that is slightly longer than tergite VIII; trapezoidal antrum plate that is sinuate posteriorly, and a pair of symmetrical sclerotized triangular structures at the posterior end of lamella antevaginalis.

Redescription. The following information is added to the original description. Head. Frons and vertex smooth scaled. Labial palpus twice length of vertical diameter of eye. Thorax. Smooth scaled. Tegula light brown with creamy-white scales caudo-dorsally. Wingspan 17–19 mm. Forewing (Fig. 8A) brownish white; each lobe of forewing on upperside with two complete white bands, a transverse preapical line and a wide line separate from apex; the bands in both lobes oriented in the same direction; base of first lobe near the cleft with a small nearly circular white patch followed distally by a small blackish spot; basal one-third with a distinct blackish spot. Fringe of forewing with three groups of dark scales at three-fourths distad of dorsum: the first one behind middle, the second in the middle, the third near apex of wing. Hindwing (Fig. 8A) light brown; subapical dark scale tooth of third hindwing lobe relatively wide (more than 10 scales); pronounced individual dark and white scales present on dorsum of third lobe. Abdomen. Light brown dorsally with whitish longitudinal lines. Length of specialized eighth sternite 1.2–1.4 times length of tergite VIII. Male genitalia (Figs 8C–G) as described for the genus. The flat valva with two latero-distal angles is an additional diagnostic character for this species. Female genitalia (Figs 8I, J) as described for the genus. Additional diagnostic characters for this species include the triangular shape of paired symmetrical sclerotized structures at posterior end of lamella antevaginalis, width of antrum which is nearly as wide as papillae anales, width of ductus bursae (0.1–0.2× width of ductus seminalis) which sometimes slightly decreasing towards corpus bursae, and occasional presence of a rather elongated sclerotized plate on one side of it.

Ecology. Unknown.Distribution. Specimens have been collected in Transvaal (Louis Trichardt), Natal, and Pretoria in South

Africa (Meyrick 1911, 1920; Gielis 2003).

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Discussion

Oxyptilus (sensu lato) constitutes one of the most difficult and challenging genera of the tribe Oxyptilini. The genus was proposed by Zeller (1841) with five species, and most of its other included species were described by Meyrick during the second half of the eighteenth century. Alipanah et al. (in press) transferred O. regulusto the genus Nippoptilia, synonymised Oxyptilus variegatus with Oxyptilus secutor, and we herein transfer the latter species to a newly described genus, leaving 20 species in Oxyptilus. However, as mentioned previously, among the 20, only five species are considered to belong to Oxyptilus. The remaining 15 do not convincingly key to Oxyptilus. Some of them can be easily excluded from the genus. For example, Alipanah et al. (in press) revealed that O. vibrans Meyrick probably belongs to Deuterocopinae, close to the genus Deuterocopus. Although Oxyptilus wallecei Fletcher was described in Oxyptilus, Fletcher (1911) mentioned that "it will be removed from this genus in the near future and will probably form the type species of a new genus somewhat intermediate between Oxyptilus and Sochchora." A close affinity bewteen Tetraschalis and O. erythrodactylusFletcher was also demonstrated by Fletcher at that time.

According to Alipanah et al. (in press), Oxyptilus causodes Meyrick differs from other Oxyptilus species in wing shape, wing pattern, wing venation, shape of the scales in the dark scale teeth at the dorsum of the third lobe of the hindwing, and in characteristics of the male and female genitalia. All the other remaining species, viz. O. catathectes Meyrick, O. celebratus Meyrick, O. epidectis Meyrick, O. erebites Meyrick, O. idonealis Walker, O. languidus Felder & Rogenhofer, O. mycites Meyrick, O. orichalcias Meyrick, O. praedator Meyrick, O. scutifer Meyrick and O. tessmanni Strand, are also different from the type species of Oxyptilus and its congeners, especially in the shape of the genitalia. Moreover, the position of dark scale teeth on the costa and dorsum of the third hindwing lobe, if present, is different from that of other Oxyptilus species. These putatively misplaced species also differ from species of the two new genera described herein by the shape of the genitalia and some external features. Accordingly, we here restrict the genus Oxyptilus to the five above-mentioned species, and it seems probable that several distinct genera are presently concealed within Oxyptilus (sensu lato). In other words, each of the remaining Oxyptilus species may be considered as a member of a new genus or even be synonymized with some previously described species in the future. Molecular studies may help reveal their exact positions.

The same situation is present in Megalorhipida and Trichoptilus. Although all the described species included in these two genera may key to the “correct” genus, further study of interspecific variation within each genus may result in the description of additional genera. For example, species of Megalorhipida can be divided into two distinct groups. According to Alipanah et al. (in press), Megalorhipida dulcis (Walsingham), M. deboeri Gielis, M. paraiso Gielis, M. madoris Gielis, and M. dubiosa Gielis (i.e., group 2) differ from the remaining Megalorhipida species in the shape of the genitalia and some of the hindwing characteristics easily distinguish them from the remaining Megalorhipida species (i.e., group 1). The groups may represent one or two new genera. All Megalorhipida species will key to that genus using the key presented above; however, members of these two groups can be distinguished by length of the antennae and costal scale tooth of the hindwing third lobe. In members of group 1 (Megalorhipida angusta Arenberger, M. fissa Arenberger, M. leptomeres (Meyrick), M. leucodactylus (Fabricius), M. prolai (Gibeaux) and M. pseudodefectalis Gielis), the length of the antenna is conspicuously less than half the length of the forewing, whereas in group 2 it is nearly half the length of the forewing. Moreover, in the latter group (except M. madoris), pronounced dark scale teeth are present at both the costa and dorsum of the third lobe of the hindwing, but in the former group, as well as in M. madoris, these are only at the dorsum of the third hindwing lobe. We have not separated the two groups in the key. We believe that the exact position of each group will be revealed after revision of known species that are included in the two genera Trichoptilus and Megalorhipida.

From the 15 known Trichoptilus species, only nine, viz. T. pygmaeus Walsingham, T. animosus Meyrick, T. cryphias Meyrick, T. festus , T. maceratus Meyrick, T. pelias Meyrick, T. varius Meyrick, T. viduus and T. vivax Meyrick, were examined here; they key convincingly to that genus. No specimens were available of the remaining species; however, because of the interspecific variation within this group, as stated by Adamczewski (1951), several distinct genera may also exist in this group as currently circumscribed.

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Acknowledgments

Our special gratitude is expressed to Jaroslaw Buszko (Copernicus University, Institute of Ecology and Environmental Protection, Toruń, Poland), Ernst Arenberger (Naturhistorisches Museum, Wien, Austria), Leif Aarvik (Natural History Museum, University of Oslo, Norway), Peter Ustjuzhanin (Siberian Division of the Russian Entomological Society, Russia), and Ole Karsholt (Zoological Museum of the University of Copenhagen (ZMUC), Denmark), who provided relevant material for this study. We gratefully acknowledge W. Gerald Tremewan (Truro, Cornwall, U.K.) for editing the English text.

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