Artículos

A review of the genus Holcophora Staudinger, 1871, with description of three new species and new data on the taxonomy of the genus (Lepidoptera: Gelechiidae)

Revisión del género Holcophora Staudinger, 1871, con descripción de tres especies nuevas y nuevos datos sobre la taxonomía del género (Lepidoptera: Gelechiidae)

Introduction

From the time of its description almost to the present the genus Holcophora Staudinger, 1871, with the type species H. statices Staudinger, 1871, was considered as monotypic. Recently, Aponoea Walsingham, 1905 (type species Aponoea obtusipalpis Walsingham, 1905) was synonymized with Holcophora, and Blastobasis inderskella Caradja, 1920, was transferred to that genus (ADAMSKI & SATTLER, 2019). These updates encouraged us to have a closer look at rich material of related taxa from different regions that was provisionally associated with Holcophora/Aponoea. Our study resulted in establishing two new generic synonyms: the monotypic Spermanthrax Meyrick, 1936 (type species S. pycnostoma Meyrick, 1936) syn. n. and the genus Epimesophleps Rebel, 1907 (type species E. symmocella Rebel, 1907), syn. n. as congeneric with Holcophora Staudinger, 1871. Based on the diagnosis of A. obtusipalpis provided by ADAMSKI & SATTLER (2019) we established that the northern African species Aponaea (sic!) pruinosella Chrétien, 1915, syn. n., and S. pycnostoma Meyrick, 1936, syn. n., are new synonyms of A. obtusipalpis whilst specimens from mainland Spain, although closely related to H. obtusipalpis, represent a separate species - Holcophorahispanica Gastón & Vives, sp. n. Our study of the holotype and other specimens of Gelechia molitor Walsingham, 1906, indicated the assignment of this species to Holcophora: Holcophora molitor (Walsingham, 1906) comb. n. We show that the re-description of H. inderskella by ADAMSKI & SATTLER (2019) refers to H.rostrella Bidzilya & Sattler, sp. n. Additionally, a new species, Holcophora centralasiae Bidzilya & Karsholt, sp. n., is described from Afghanistan and Pakistan. The diagnosis of Holcophora is improved by consideration of additional characters, and the systematic position of that genus within the subfamily Gelechiinae is briefly discussed. All eight species currently associated with Holcophora are diagnosed, and the data on their distribution and biology are summarised. Additional we briefly deal with Epimesophleps aphridias Meyrick, 1925, and formally transfer it to Holcophora. However, due to lack of material we are unable to discuss it in detail. We provide keys for species identification based on external and genitalia characters.

Materials and methods

Moths examined included type specimens as well as non-type specimens from the institutional and private collections listed below. Data from holotypes are cited exactly as on the labels of the specimens, whereas other material is organised in a standardized format rather than verbatim, viz., alphabetic after country and province, region etc.; material from the same province is listed chronologically. The photographs of the male genitalia are provided both in traditional ventral view and “unrolling” except for H. symmocella.

OB photographed pinned specimens and their genitalia as described by BIDZILYA et al. (2017).

Our terms for the male and female genitalia follow HODGES (1986) and PONOMARENKO (2008, 2009).

Our arrangement of the species within the genus Holcophora is based principally on similarities of the male genitalia.

Abbreviations of collections

MfN Museum für Naturkunde, Berlin, Germany

MNCN Museo Nacional de Ciencias Naturales, Madrid, Spain

MNHN Muséum National d’Histoire Naturelle, Paris, France

NHMUK Natural History Museum, London, United Kingdom

NHMB Hungarian Natural History Museum, Budapest, Hungary

NHMV Naturhistorisches Museum, Vienna, Austria

NMPC National Museum, Prague, Czech Republic

RCFG Research collection Friedmar Graf, Bautzen, Germany

RCJG Research Collection Javier Gastón, Getxo, Vizcaya, Spain

RCMD Research Collection Marek Dvorˇák, Smrèná, Czech Republic

RMHN Naturalis Biodiversity Center, Leiden, The Netherlands

SMNK State Museum of Natural History Karlsruhe, Karlsruhe, Germany

ZIN Zoological Institute, Russian Academy of Sciences, Sankt-Petersburg, Russia

ZMKU Zoological Museum, Kyiv Taras Shevchenko National University, Kyiv, Ukraine

ZMUC Zoological Museum, Natural History Museum of Denmark, Copenhagen, Denmark

Other abbreviations

AV Antonio Vives

comb. n. new combination

gen. slide genitalia slide

JG Javier Gastón

KS Klaus Sattler

OB Oleksiy Bidzilya

OK Ole Karsholt

syn. n. new synonym

TL Type locality

TS Type species

Abbreviations on figures

cc - cucullus; gn - gnathos; sc - sacculus; cul - culcitula; tg pr - process of tegumen; un - uncus

Results

Check list of the species of Holcophora

Holcophora hispanica Gastón & Vives, sp. n.

Holcophora obtusipalpis (Walsingham, 1905)

= Mesophleps cinerellus Turati, 1930

= Aponaea (sic!) pruinosella Chrétien, 1915, syn. n.

= Spermanthrax pycnostoma Meyrick, 1936, syn. n.

Holcophora statices Staudinger, 1871

Holcophora inderskella (Caradja, 1920)

Holcophora centralasiae Bidzilya & Karsholt, sp. n.

Holcophora rostrella Bidzilya & Sattler, sp. n.

Holcophora molitor (Walsingham, 1906), comb. n.

Holcophora symmocella (Rebel, 1907), comb. n.

Holcophora aphridias (Meyrick, 1925), comb. n.

Generic descriptions

Holcophora Staudinger, 1871. Berl. ent. Z., 14(3/4): 313

TS: Holcophora statices Staudinger, 1871. Berl. ent. Z., 14(3/4): 313-314, by monotypy

= Aponoea Walsingham, 1904. Entomologist’s mon. Mag., 40: 216, nomen nudum

= Aponoea Walsingham, 1905. Entomologist’s mon. Mag., 41: 125

TS: Aponoea obtusipalpis Walsingham, 1905. Entomologist’s mon. Mag., 41: 125, by original designation and monotypy. (Synonymized by ADAMSKI & SATTLER, 2019: 18)

= Epimesophleps Rebel, 1907. Lepid. Südarabien u. Insel Sokótra: 95. Syn. n.

TS: Epimesophleps symmocella Rebel, 1907. Lepid. Südarabien u. Insel Sokótra:, 95, fig. 40, by monotypy

= Aponaea; Chrétien, 1915. Ann. Soc. ent. Fr., 84: 330, lapsus calami

= Spermanthrax Meyrick, 1936. Exotic Microlepid., 4(20): 624. Syn. n.

TS: Spermanthrax pycnostoma Meyrick, 1936. Exotic Microlepid., 4(20): 625, by monotypy

Generic diagnosis: The species of Holcophora are distinguished externally by a long labial palpus (up-curved or more or less porrect) with segment 3 usually shortened (1/5-1/3 length of segment 2), and segment 2 with brush of raised scales on its dorsal surface (Figs 1-12). The predominantly plain grey or pale forewing with indistinct markings and groups of raised scales (Figs 21-23) in conjunction with a weakly excavated hindwing termen are also diagnostic. The male genitalia of Holcophora usually are strongly modified. A tendency towards asymmetry in the dorsal sclerites of the male genitalia (uncus displaced laterally and turned anti-clockwise, tegumen bearing lateral process of irregular shape) is characteristic for most species. Additional diagnostic characters of the male genitalia are the presence of membranous lobes covered with long setae on both sides of the uncus base (culcitula), the usually shortened tegumen, the strongly reduced gnathos, anteromedially strongly broadened sacculus, weakly sclerotized distal part of the phallus with distinct lateral rod ending in short sclerite or long, strongly recurved apical process, and very long bulbus ejaculatorius. The female genitalia are characterized by the weakly modified, evenly sclerotized (except anterior margin) segment VIII with sternum usually deeply emarginated both anteriorly and posteriorly, the short, tubular colliculum, short apophyses anteriores and posteriores and the Gelechia-type signum.

Remarks: The systematic position of Holcophora within the Gelechiidae is uncertain and somewhat controversial. That is due to the strongly modified male genitalia and in parts still unresolved system of the family in general.

Staudinger placed the genus Holcophora originally between Anarsia Zeller, 1839, and Hypatima Hübner, [1825] (“Chelaria Haworth, 1828”); it was subsequently included by MEYRICK (1925) in the Dichomeris group (now Dichomeridinae). In some regional catalogues and faunistic lists (KARSHOLT et al., 1996; ELSNER et al., 1999) it was tentatively placed in the tribe Chelariini, subfamily Gelechiinae, but no characters were given in support of that position. PONOMARENKO (2009) treated Holcophora as a member of the tribe Gelechiini within the subfamily Gelechiinae based on her study of the functional morphology of the male genitalia, and that concept was accepted by ADAMSKI & SATTLER (2019).

The subfamily Gelechiinae is considered as monophyletic based on the presence of mediobasal processes of the valva and on glands of the genital segment (PONOMARENKO, 2005: 73, 74, 90). KARSHOLT et al. (2013) following HUEMER & SATTLER (1995) recognized Gelechiinae based on the horizontal division of the male pregenital abdominal segment VIII into free dorsal and ventral flaps that permit wider opening of that segment for the extrusion of the genitalia during copulation. Although different authors proposed a different suite of autapomorphies, the monophyly of Gelechiinae (with tribes Gelechiini, Gnorimoschemini and Litini) is well supported by a molecular analysis (KARSHOLT et al., 2013: 344), and the subfamily in this sense may be considered satisfactorily defined both morphologically and molecularly. Within Gelechiinae, the tribes Litini and Gnorimoschemini are considered as monophyletic by all authors. The third tribe, Gelechiini, is clustered with Gnorimoschemini based on the laterally dilated vinculum (PONOMARENKO, 2005). According to a molecular analysis Gelechiini are paraphyletic with respect to Litini and Gnorimoschemini (KARSHOLT et al., 2013: 345). Thus, the monophyly of Gelechiini still remains unresolved.

Amongst characters that justify the placement of Holcophora as a member of Gelechiini, PONOMARENKO (2009: 140) listed the ridge-shaped tegumen without a ventral wall, the basally broadening valva and the well-developed muscle m 22 in the male genitalia. In our opinion the first two characters are rather vague: the degree of “broadening” of the valva is subjective and difficult to estimate. The “ridge-shaped tegumen” seems also present in some genera of other tribes and can hardly be considered as phylogenetically significant. The reduced muscle m 22 is considered as one of the diagnostic character for Gelechiini (PONOMARENKO, 2005: 74), and its presence in Holcophora would contradict the placement of the genus in that tribe. However, Ponomarenko seems to contradict herself with regard to muscle m 22 the reduction of which she considers as a Gelechiini character in 2005 whereas in 2009 it is the well-developed m 22. Nevertheless, we agree with Ponomarenko that Holcophora cannot be placed in Dichomeridinae (including Chelariini) as proposed earlier but should be assigned to Gelechiini, a position that is supported by the characteristic Gelechia-type signum bursae. In view of the strongly modified male genitalia we are still unable to assign Holcophora to a convincing place within that tribe.

In Gelechiidae asymmetry in the male genitalia is known in a number of genera, e.g. Caulastrocecis Chrétien, 1931, Horridovalva Sattler, 1967 (Anomologinae), Anarsia Zeller, 1839 (Chelariini), Coleotechnites Chambers, 1880 (Litini), Chionodes Hübner, [1825], Photodotis Meyrick, 1911 (Gelechiinae), Thiotricha Meyrick, 1886 (Thiotrichinae) etc. In all these genera the asymmetry affects mainly the valvar complex, but the asymmetry in the uncus and tegumen of Holcophora appears to be unique in the family Gelechiidae. A reduced gnathos, characteristic of Holcophora, is found in many unrelated genera but most commonly in Anomologinae, Apatetrinae, and Litini of the Gelechiinae; it has not yet been found amongst Gelechiini. Another unique character of Holcophora is the rounded, densely setose lobes of the posterior margin of the tegumen on both sides of the uncus (Figs 65, 67, 76). Somewhat similar structures are known in some species of Anarsia, e.g. A. spartiella (Schrank, 1802) (Chelariini), and Platyedra subcinerea (Haworth, 1828) (Pexicopiini), however, their homology is uncertain. Ponomarenko suggested that these in Holcophora can be interpreted as the modified culcitula, a structure commonly present in some related genera of Gelechiini.

H. molitor and H. symmocella share a phallus with an unusually long, narrow, strongly recurved apex and uniquely modified, possibly androconial, scales on the apex of the sacculus (Figs 74-77). Similarly specialised scales were otherwise observed in Gelechiidae only in some species of Anarsia Zeller, 1839.

The long, strongly recurved process at the apex of the phallus in H. molitor and H. symmocella is a unique specialisation that indicates a closer relationship of these two species vis-à-vis the rest of the genus. The short ring-like male abdominal segment VIII in Holcophora (Figs 51-52) with laterally fused sternum and tergum is not characteristic for Gelechiinae, but was also observed in Athrips Billberg, 1820, the assignment of which to that subfamily is commonly accepted (HUEMER & KARSHOLT, 1999; PONOMARENKO, 2004; BIDZILYA, 2005). The raised scales on the forewing are found in all tribes of Gelechiinae, but occur also in some Pexicopiini (e.g. Sitotroga exquisita Bidzilya & Mey, 2011) and Dichomeridinae (Dichomeris alacella (Zeller, 1839)).

It appears that an appropriate position of Holcophora within the Gelechiidae cannot be satisfactorily established based alone on traditional morphological characters or on the functional morphology of the male genitalia. However, that genus shares more diagnostic characters (valva divided into cucullus and basally broadening sacculus, free phallus, Gelechia-type signum, raised scales on forewing) with Gelechiinae (Gelechiini) than with any other currently recognised Gelechiidae subfamily. The location of Holcophora within Gelechiini remains uncertain at this stage.

Description: Adult. Head smoothly scaled except dishevelled lateral tufts on neck and on base of antennae, scales plain grey or grey with brown tip; ocelli present; labial palpus (Figs 1-12) long, more or less porrect or moderately up-curved, segment 2 2 to 5 times as broad as segment 3, with distinct tuft of raised scales on dorsal surface (absent in H. centralasiae sp. n.), more or less uniformly coloured without distinct rings on outer surface, segment 3 usually short, 1/4-1/5 length of segment 2 (1/3-1/2 in H. centralasiae sp. n.), slender, acute; proboscis long; in H. rostrella sp. n. frons strongly modified, bearing crater-like structure with long, pointed central process (ADAMSKI & SATTLER, 2019: figs 10-13); antennal scape without pecten, 1.5-2 times as broad as adjacent part of flagellum, flagellomeres dark, grey to brown or black with white rings, denuded (H. centralasiae sp. n.), with short (H. statices) or longer (H. obtusipalpis, H. hispanica sp. n.) cilia in some species (Figs 13-17); thorax and tegulae concolorous with head; forewing usually plainly coloured, pale, light grey to brown, scales with black tips, pattern, if present, represented by oblique basal fascia and diffuse markings in cell, in H. statices with more distinct black spots and streaks, raised scales usually gathered in small tufts distal to basal fascia and in middle of cell (Figs 21-23); fringe concolorous with ground colour of forewing; hindwing usually light grey to almost white, but dark, blackish-brown in H. hispanica sp. n. frenulum simple in male, composed of three acanthi in female (Fig. 20); anal zone of hindwing and dorsolateral surface of metathorax with brushes of androconial scales (Figs 18-20).

Abdomen: Male segment VIII as long as or slightly longer than segments III-VII (Figs 51-52), with tergum and sternum fused laterally, forming ring with posterior part of vinculum and anterior margin of tegumen; in female segment VII 1.5 times to twice as broad as segments II-VI; tergum I broader than long, stronger sclerotized in distal part; sternum II longer than broad, with distinct venulae and long apodemes (Figs 53-55); all segments bearing short hairs in addition to scales (Fig. 56). Male genitalia. Uncus short, triangular, bearing long setae (H. hispanica sp. n., H. obtusipalpis) or displaced laterally, narrow, hook-shaped, or long, narrow or moderately broad process, usually bent over posterior margin of tegumen; gnathos membranous lobe with straight posterior margin and reduced medial sclerite (H. hispanica sp. n., H. obtusipalpis, H. statices) or completely reduced; tegumen usually broader than long (except in H. rostrella sp. n.), with paired rounded lobes on distal margin at both sides of uncus densely covered with long setae (culcitula) in most of species, anteromedial emargination large, broadly rounded, lateral margins usually bearing processes of irregular shape; cucullus slender, straight or weakly curved, broadening apically in H. rostrella sp. n., strongly broadening in basal half (except in H. rostrella sp. n.) and fused basally with sacculus; sacculus broader than cucullus (except in H. rostrella sp. n.), its basal part distinctly broadening to form medially fused dorsal processes (transtilla), and weakly inflated ventral sclerites fused with vinculum anteriorly and with base of cucullus laterally, apex of sacculus simple or bifurcated, with specialized (?androconial) scales in H. symmocella and H. molitor; vinculum narrow, band-like; saccus stout, broad, long, triangular to sub-rectangular; phallus with distinctly swollen caecum, distal part membranous except for strongly sclerotized lateral rod(s) ending in short apical sclerite or long, strongly recurved process in H. symmocella and H. molitor, bulbus ejaculatorius long, anterior part of ductus ejaculatorius in some species with small ring-like sclerotization.

Female genitalia: Papillae anales ovate or sub-triangular, covered densely with short setae; apophyses posteriores slender, slightly longer than papillae anales, with distinctly swollen apex in H. rostrella sp. n.; apophyses anteriores 1/4-1/2 length of sternum VIII, straight, with broadening base in H. statices; tergum VIII sub-rectangular, evenly sclerotized except for strongly edged anterior margin, with deep broadly anteromedial emargination; sternum VIII weakly sclerotized, medially membranous, pair of spiracular openings present, anterior margin strongly sclerotized, narrow band-shaped or moderately broadening, straight or projecting anteriorly, with more or less distinct sclerites on both sides of ostium in some species; ostium opening near anterior margin of sternum VIII; antrum short, tubular or funnel-shaped, posterolaterally sclerotized, with medial sclerite in H. obtusipalpis, with indistinct transition to ductus bursae or well separated (H. obtusipalpis, H. molitor, H. symmocella) from it; ductus bursae long, slender, weakly broadening towards corpus bursae, or moderately broad (H. molitor, H. symmocella), ductus seminalis slender, arising from border between antrum and ductus bursae; corpus bursae egg-shaped to rounded, varying in size; signum plate rounded to rhomboid or sub-hexagonal, with serrated margins, medial zone varying considerably in width from narrow slit (H. centralasiae sp. n.) to large rounded plate occupying almost entire surface of signum plate (H. rostrella sp. n.), in H. hispanica sp. n. signum long lanceolate plate with serrated borders, pointing towards fundus bursae, and fixed by cylindrical process to wall of corpus bursae.

Notes: Despite some superficial similarity, most species of Holcophora can be distinguished by their wing pattern as well as colour and shape of the labial palpus. The female genitalia of Holcophora species have a rather similar ground plan. The most constant characters for species separation are found in the shape of the antrum, presence or absence of subostial sclerites (colliculum) and the comparative length of the ductus bursae and corpus bursae. The signum is also diagnostic for most species, especially for H. hispanica sp. n. However, this structure varies considerably in some species and in such cases can only be used for species diagnosis in combination with other characters. Contrary to the rather uniform external appearance and female genitalia, the male genitalia of Holcophora show rather broad variation. Two species with symmetrical genitalia and a short triangular uncus (H. hispanica sp. n. and H. obtusipalpis) are in contrast to all other species which are characterized by the strong asymmetry of tegumen and uncus. H. molitor and H. symmocella can easily be recognized by the modified (? androconial) scales on the apex of the sacculus and by the phallus with a long, narrow, strongly recurved apex. All other species are easily distinguished by the shape of the uncus, processes of tegumen, phallus and other characters. Despite some considerable morphological variation the male genitalia of Holcophora species follow an identical ground plan that justifies their treatment as members of one genus. It is the reason for considering the monotypic Spermanthrax Meyrick, 1936 (type species S. pycnostoma Meyrick, 1936) and the genus Epimesophleps Rebel, 1907 (type species E. symmocella Rebel, 1907) as congeneric with Holcophora Staudinger, 1871.

Molecular data are available only for H. obtusipalpis and H. hispanica sp. n. (HUEMER et al., 2020: suppl. data 2).

Biology: Host-plants: Limoniastrum guyonianum Durieu ex Boiss., Limonium dichotomum (Cav.) O. Kuntze, Limonium meyeri (Boiss.) O. Kuntze, and Limonium pruinosum (L.) Chaz. (Plumbaginaceae). Known for three of the eight species of Holcophora. The larva lives quite exposed on its host-plant.

The currently known Holcophora species occur in open country - grasslands, mountain steppe slopes, semideserts and deserts up to 2300 m in mountains of Central Asia.

Distribution: Palaearctic and Afrotropical regions.

Key to the species of Holcophora based on external characters

1. Head with prominent frontal modification (Fig. 46) .......................................... H. rostrella sp. n.

- Head unmodified ........................................................................................................................... 2

2. Segment 2 of labial palpus dorsally with large triangular tuft of raised scales (Figs 1-4) ........... 3

- Segment 2 of labial palpus dorsally unmodified or with short brush of raised scales in distal 1/3 (Figs 5-8, 10-12) .............................................................................................................................. 4

3. Forewing grey with randomly spread black scales, hindwing light grey (Figs 30-35) .............. H. obtusipalpis

- Forewing dark grey with black scales concentrated mainly along veins, hindwing dark brown (Figs 24-29) .......................................................................................................... H. hispanica sp. n.

4. Segment 2 of labial palpus covered dorsally with smooth scales (Fig. 7) ... H. centralasiae sp. n.

- Segment 2 of labial palpus dorsally with short brush of raised scales in distal 1/3 ...................... 5

5. Forewing dark grey, contrasting, veins and fold mottled with light brown, black markings distinct, hindwing dark grey (Figs 36-38) ......................................................................... H. statices

- Forewing light grey to light brown, plain, without light brown irroration along veins, black markings indistinct (Figs 42-44, 47-50) ........................................................................................... 6

6. Segment 2 of labial palpus weakly curved, far protruded forwards, 3 times as broad as segment 3 (Fig. 8) ...................................................................................................................... H. inderskella

- Segment 2 of labial palpus distinctly turned upwards, segment 2 twice breadth of segment 3 (Figs 10-12) ...................................................................................................................................... 7

7. Apex of hindwing broadly rounded or just weakly pointed (Fig. 47) .................... H. symmocella

- Apex of hindwing more pointed (Figs 48-50) ................................................................ H. molitor

Key to the species of Holcophora based on male genitalia

1. Genitalia symmetrical (Figs 57-62) ............................................................................................. 2

- Genitalia asymmetrical (Figs 63-77) ............................................................................................. 3

2. Anteromedial emargination of tegumen extending to 1/3 length of tegumen, cucullus about same width of sacculus (Figs 57-59) .................................................................... H. hispanica sp. n.

- Anteromedial emargination of tegumen extending to 1/2-2/3 length of tegumen, cucullus distinctly narrower than sacculus (Figs 60-62) .......................................................... H. obtusipalpis

3. Uncus placed medially, tegumen longer than broad (Figs 71-73) ..................... H. rostrella sp. n.

- Uncus placed on left corner of tegumen, tegumen as long as broad or broader than long ............ 4

4. Sacculus covered apically with modified scales, apical sclerite of phallus long strongly recurved (Figs 74-77) ...................................................................................................................................... 5

- Sacculus apically without modified scales, apical sclerite of phallus short, straight .....................6

5. Uncus rounded apically, without dorsal process, sacculus 3 times as broad as cucullus, saccus parallel-sided to 3/4 length (Fig. 77) .......................................................................... H. symmocella

- Uncus pointed apically, with dorsal process, sacculus twice as broad as cucullus, saccus triangular (Figs 74-76) ....................................................................................................... H. molitor

6. Uncus short, twice as long as broad, process on left margin of tegumen long, apex of phallus with two sclerites facing in opposite direction (Figs 68-70) ....................................... H. inderskella

- Uncus long, considerably longer than broad, process on left margin of tegumen short or absent, apex of phallus with one sclerite (Figs 63-67) ................................................................................. 7

7. Uncus sharply pointed, tegumen with short process on left margin and longer process on right margin, sacculus 3 times as broad as cucullus (Figs 65-67) ............................................. H. statices

- Uncus not pointed, tegumen without process on left margin but short process on right margin, sacculus almost length of cucullus (Figs 63-64) .............................................. H. centralasiae sp. n.