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The genus Zizyphia Chrétien, 1908, with notes on its systematic position and the first record of Z. cleodorella Chrétien, 1908 from Europe (Lepidoptera, Depressariidae, Cacochroinae)
expand article infoLauri Kaila‡, Ole Karsholt§, Txema Revilla|
‡ University of Helsinki, Helsinki, Finland
§ Zoological Museum, Natural History Museum of Denmark, Copenhagen, Denmark
| Unaffiliated, Berango, Spain

Corresponding author: Ole Karsholt ( okarsholt@snm.ku.dk )

Academic editor: David C. Lees
© 2024 Lauri Kaila, Ole Karsholt, Txema Revilla.
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: Kaila L, Karsholt O, Revilla T (2024) The genus Zizyphia Chrétien, 1908, with notes on its systematic position and the first record of Z. cleodorella Chrétien, 1908 from Europe (Lepidoptera, Depressariidae, Cacochroinae). Nota Lepidopterologica 47: 19-28. https://doi.org/10.3897/nl.47.115542
ZooBank: urn:lsid:zoobank.org:pub:E8D51484-4007-4B67-BCDD-1550B34CCC6E
Open Access

Abstract

Zizyphia cleodorella Chrétien, 1908 is reported as new to Spain and Europe. The genus Zizyphia and the species Z. cleodorella is diagnosed and its separation from the closely related Z. zizyphella Amsel, 1935 is elaborated. The adult habitus and genital structure, and the species’ habitat are illustrated. The systematic position of the genus is discussed. Because Orophiinae Lvovsky, 1974 is a secondary homonym of Orophiini Thomson, 1863 (Coleoptera: Ciidae), and Cryptolechiinae is considered here as being unrelated to it, the name Cacochroini Leraut, 1993 is reinstated as a valid family-group name as Cacochroinae within Depressariidae (Gelechioidea), with Eutornini Lvovsky, 2019, syn. nov. as a subjective junior synonym of Cacochroini.

Introduction

The genus Zizyphia Chrétien, 1908 was established based on its type species Z. cleodorella Chrétien, 1908 from Algeria. So far only one additional species Z. zizyphella Amsel, 1935 has been added to the genus. Since their description both species have only been mentioned a few times in the lepidopterological literature (Gerghiou 1977; Lvovsky 2009; Lvovsky et al. 2016; Christophides 2017).

During field work in October 2022 in southern Spain (province of Almeria) Txema Revilla and Peder Skou independently collected specimens of a rather small, brownish gelechiid-looking moth, which turned out to be Z. cleodorella. The specimens represent the first record of this species, and its genus, from both Spain and Europe. It is a little-known species, with a rather confused history regarding its systematic position. We therefore find it justified to discuss and illustrate it, in order to make it better known.

Materials and methods

Specimens

The studied material of Z. cleodorella was collected in the El Toyo/Retamar area and on the slopes of the Sierra Alhamilla in the province of Almería, Andalucía, Spain. It was attracted to ultraviolet light (8W/12V tubes and 125W mercury vapour bulbs), installed in portable light traps (Olsen et al. 1984) or attached to a vertical frame covered in a white sheet. We also studied dry, pinned museum specimens deposited in the institutional collections listed below. Genitalia were dissected using standard methods and mounted on permanent slides, with Euparal as the mounting medium.

Abbreviations of collections

MZH Finnish Museum of Natural History, Helsinki, Finland

SMNK Staatliches Museum für Naturkunde, Karlsruhe, Germany

TxR Research collection of Txema Revilla, Berango (Vizcaya), Spain

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

Results

Diagnosis of Cacochroinae

We consider the genus Zizyphia to belong to Cacochroinae (Depressariidae) (see Discussion on systematics and nomenclature below). Members of Cacochroinae share the usual characteristics of Gelechioidea, i.e. the basally-scaled haustellum and long, upcurved labial palpi, the second segment of which is particularly long. They are relatively small and slender winged compared to most Depressariidae. The forewing pattern usually contains longitudinal and/or oblique stripes or bands, sometimes with raised scales or scale tufts. The scape of the antenna lacks pecten. The tergal segments of the abdomen lack areas covered by spines. The male genitalia either lack the uncus or it is vestigial, but they have a well-developed tuba analis. They usually have a gnathos that is mesially divided into a paired spinose knob, with Cacochroa and Rosetea Corley & Ferreira, 2019 being exceptions that lack the gnathos. The shape of the valva varies, but it often contains a curved lobe at the apex of the costa and/or the sacculus is distally separated. The lateral lobes of the juxta are prominent and sometimes very long. The phallus is pistol-shaped with a curved caecum, and is attached to the anellus, that can be membranous or sclerotized. The female genitalia of Cacochroinae have a telescopic ovipositor, but otherwise they vary considerably between genera, and seem not to have any particular shared features amongst all genera. The larvae of only few species are known, but at least some are leaf miners to begin with and then tie or roll leaves.

The male genitalia of Cacochroa permixtella (Herrich-Schäffer, 1854) are anomalous, compared to virtually all other species of Gelechioidea, in having an enormous-sized vinculum (cf. Corley and Ferreira 2019). The female genitalia are rather similar to that of Zizyphia, but the signum is more similar to that of Rosetea. Along with Rosetea, it also differs from others in not having a gnathos. It should be noted that the structure called the uncus by Corley and Ferreira (2019) for C. permixtella is actually the anellus that surrounds the phallus. The true uncus is vestigial and weakly sclerotized, situated dorsad of the more distinctive tuba analis. Even though C. permixtella has such distinctive features, it also shares characters with Rosetea, including the vestigial uncus and well-developed tuba analis, and divided valva, in addition to a markedly similar outer appearance. It also has broadly separated valvae that are typical of other genera. Therefore, we consider C. permixtella a close relative of other genera treated here. However, discussion of the status or species composition of the established cacochroine genera is beyond the scope of this publication.

Zizyphia Chrétien, 1908

Zizyphia Chrétien, 1908: 166. Type species: Zizyphia cleodorella Chrétien, 1908, by monotypy. Type Locality: Algeria; Meyrick, 1925: 36.

Diagnosis

Zizyphia adults externally most closely resemble those of some species of Orophia and the predominantly East-Asian – Australasian genus Eutorna in having a similar wing shape, with the forewings being relatively narrow. The wing pattern is also similar to that of several Eutorna species and Orophia zernyi (Szent-Ivány, 1942). The male genitalia of Zizyphia share with other cacochroines the vestigial or entirely lacking uncus, the well-developed tuba analis, divided mesial knob of the gnathos, the bilobed valva and the broad vinculum. Note that in the genera Cacochroa and Rosetea the gnathos is absent. The knobs of the gnathos are significantly larger in Zizyphia than in other genera. This seems to be the only male genitalia character that separates Zizyphia from Orophia. Externally, the species in these genera are also fairly similar, with the apex of the forewing being somewhat more acute in Orophia than in Zizyphia. In their male genitalia, unlike in Rosetea, the costa of the valva is weakly sclerotized, and the sacculus is extended into a prolonged hook. The vinculum is broad, but not as long as in Cacochroa. In the Eutorna species examined (E. leonidi Lvovsky, one unidentified species from Australia, and another from New Zealand) the valvae are undivided. The females of Zizyphia differ from those of Orophia as follows: in Orophia the ductus bursae and the colliculum are narrow and elongate, being longer than the corpus bursae. The ductus bursae is otherwise not sclerotized. In Zizyphia the ductus bursae is broad and there is a separate, elongate sclerotization on its ventral side.

Remarks

Chrétien (1908) did not compare Zizyphia with other genera, but placed it next to Holcophora Staudinger, 1871, Sophronia Hübner, 1825) (both Gelechiidae) and Holcopogon Staudinger, 1879 (now Autostichidae: Holcopogoninae). Meyrick (1925) stated that he had not seen material of Zizyphia, but based upon its original description, he doubted that it was related to Sophronia.

Zizyphia cleodorella Chrétien, 1908

Zizyphia cleodorella Chrétien, 1908: 167; Meyrick 1925: 36; Mück 1985: 20, 163; Báez and García 2005: 87.

Examined material

Spain, prov. Almeria, Sierra Alhamilla, 400 m, 1♀ 3/4.x.2022, leg. Tx. Revilla (TxR); prov. Almeria, Retamar, 25 m, 1♀ 3/4.x.2022, leg. Tx. Revilla (TxR); same locality but 15 m, 3♂, 4/5.x.2022, 4♂, 2♀, 16.x.2022, P. Skou leg. (MZH, ZMUC).

Description (Fig. 1)

Forewing brown, costal side broadly grey. Along basal 1/3 of wing length at centre, grey spot varying from small, dark grey and distinctive to indistinctly delineated grey area blended with the costal grey area. Along fold white, longitudinal stripes, irregular in number and strength. Near apex from costa to apex, oblique, white stripe; before apex black spot. In the male genitalia (Fig. 3) sacculus of valva separated from the rest, and terminating in a curved, sclerotized hook. Phallus short and broad, pistol-shaped with broad and bent caecum, with one long and another shorter cornutus, and near base comb-like dense group of spines. Female genitalia (Fig. 4) characterized by broad ductus bursae, with short and broad, sclerotized antrum, ductus bursae otherwise membranous; on ventral side of ductus bursae a conspicuous sclerotization, three times as long as broad. Ductus seminalis basally bulbous and situated in posterior end of corpus bursae; corpus bursae with one small, star-shaped signum.

Figures 1, 2. 

Zizyphia adults. 1. Z. cleodorella Chrétien; 2. Z. zizyphella Amsel, holotype.

Biology

The larva and pupa were described in detail in the original description (Chrétien, 1908), but the life history was only briefly summarized. The larva feeds during May on Ziziphus (probably Z. mauritiana Lam. or Z. lotus Lam.) (Rhamnaceae), afterwards descending to the ground to pupate. The adults emerge in June (Chrétien 1908) and October. Mück (1985: 20, 163) lists Z. cleodorella from Ziziphus mauritiana in the Cape Verde Islands, without a detailed locality, but it is unclear if this was based on personal observation.

The biotope where this species occurs in Spain is a sandy area with narrow, mobile dunes (Fig. 7), within the semi-arid thermo-Mediterranean climatic zone with high atmospheric humidity, where extensive formations of Ziziphus lotus (“azufaifo” in Spanish) occur, together with patches of annual grassland. Tamarix sp., Ephedra sp., Ononis sp., Thymelaea sp. etc. complete the markedly African botanical character of this landscape, distinctive of the Retamar area where Z. cleodorella occurs.

Figures 3–6. 

Zizyphia genitalia. 3. Z. cleodorella Chrétien, ♂; 4. Z. cleodorella Chrétien, ♀; 5. Z. zizyphella Amsel ♂; 6. ♀. In the male genitalia images the anellus has been severed from the rest of the genitalia and is surrounding the phallus.

Distribution (Fig. 8)

Algeria, Cape Verde Islands and Spain.

Remarks

Zizyphia cleodorella was described from an unstated number of specimens bred from larvae found in Biskra, Algeria.

Zizyphia zizyphella Amsel, 1935

Zizyphia zizyphella Amsel, 1933: 127 (nomen nudum); Amsel 1935a: 298, pl. 11, fig. 101; 1935b: 210; 1935c: 266; Gerghiou 1977: 198; Lvovsky 2009: 149–152, figs 1–2; Gozmány 2012: 348–349; Lvovsky et al. 2016: 108.

Examined material

Image of the holotype ♀ with labels: Jericho (Pal), 30.4.1930 H. Amsel (Lichtfang); Typus leg. H. Amsel; ex coll. H.G. Amsel (SMNK).

Examined paratypes: 1♂ with same collecting data as in holotype, but the collecting date 28.iv.1930; with labels Zizyphia zizyphella Amsel teste A.L. Lvovsky; L. Kaila prep. 6358 (SMNK); 1 ♀: Jericho Z.312 Zyzyphus spina-christi 4.5.; Palästina expedition 18.2.–14.6.1930 H.G. Amsel, L. Kaila prep 6359 (SMNK).

Diagnosis (Fig. 2)

Similar to Z. cleodorella, but forewings paler ochreous brown with only a little grey along costa; near apex usually two inconspicuous dark spots (forewing in Z. zizyphella dark brown, costal side broadly grey, near apex one more prominent spot). Male genitalia (Fig. 5) similar to those of Z. cleodorella. The sclerotization in the female ostium+antrum is shorter and broader, and likewise, the ductus bursae is with shorter and broader sclerotization in Z. cleodorella (Fig. 6) than in Z. zizyphella. In Z. cleodorella the sclerotization is three times as long as broad, whereas in Z. zizyphella it is four times as long as broad.

Biology

According to the original description the type series was collected as larvae in late April and early May on Ziziphus spina-christi (L.) Willd., but according to a label on the holotype it was apparently collected at light. In the original description Amsel (1935a: 298) gave no details on the life history, but stated that the larvae were common. In another paper from the same year (Amsel, 1935c) he described the larva in some detail, stating that it lives in a boat-shaped leaf, which it skeletonizes. When disturbed it becomes very lively. Gerghiou (1977: 198) reported larvae being a leaf tier on leaves on Paliurus sp. (Rhamnaceae) in January and July-August, mentioning that these specimens had been identified by H. G. Amsel, the author of the species. According to Christophides (2017: 282), records of Paliurus from Cyprus refer to Zizyphus lotus.

Distribution (Fig. 8)

Known only from the type locality Jericho (Palestine/Israel) and the southern shore of the Dead Sea (Amsel, 1935b: 210), and from Cyprus, but without an exact locality (Gerghiou 1977). Z. zizyphella was not included in Fauna Europaea (Karsholt and Nieukerken 2004–2023) although this website treats Cyprus as a part of Europe.

Remarks

The name Zizyphia zizyphella was first listed by Amsel (1933: 127) in a checklist of Palestinian Lepidoptera, but without a description. It was subsequently described by Amsel two years later (Amsel 1935a). In that publication the species was illustrated from a black/white photograph of low quality, making it look much darker that the holotype, which may in turn be bleached.

Although Z. zizyphella is similar to the other species of the genus, Z. cleodorella, Amsel (1935a: 298) wrote: “Mit keiner Art näher verwant” (Not related to any species).

Zizyphia zizyphella was described within the Gelechiidae, and stayed there until Lvovsky (2009) transferred it into the Depressariidae (tribe Orophiini). During a recent survey of the Gelechiidae of Israel (Bidzilya et al. 2019) no additional specimens were examined.

Discussion

The name of the genus Zizyphia refers to the host plant, from which the larvae of the type species, Z. cleodorella were found. Ziziphus is a genus of evergreen shrubs or medium-sized trees. Some species, e.g. Z. mauritiana and Z. jujuba (English: Jujube or Chinese Date), originating from South Asia, have been planted in other parts of the world, because they are edible (especially the fruits) and can survive in dry areas. Considering that they are grown for horticultural purpose, it is surprising that we have only been able to find little published information on the life histories and distribution of these two species. Therefore, Z. cleodorella may have expanded its distribution northward due to climate change rather than human activity.

Figures 7, 8. 

7. Habitat of Zizyphia cleodorella Chrétien, Spain, province Almeria, Retamar; 8. Distribution map of Zizyphia cleodorella Chrétien and Z. zizyphella Amsel.

With ongoing climate change and the consequent increase in temperatures, the distributions of “exotic species” are expanding to include regions that until recently did not provide suitable climates for them. The Spanish Mediterranean coast, affected by recent climate change, offers clear examples of such range shifts in the Microlepidoptera fauna, with recently recorded appearances of other species of Gelechioidea, e.g. Batrachedra amydraula Meyrick, 1916 (Batrachedridae) in Mojacar (Almería) in 2019 (Revilla and Gastón 2019), Labdia semicoccinea (Stainton, 1859) (Cosmopterigidae) in Motril (Granada) in 2022 (Olvera et al. 2022), and perhaps also Xenotorodor stygioxanthus Sterling, Lees & Grundy, 2023 (Lecithoceridae) near Tarifa (Cádiz).

Notes on systematics and nomenclature on Cacochroinae

Heikkilä et al. (2014) and Wang and Li (2020) preliminarily placed Cacochroinae (as Cryptolechiinae) in the family Depressariidae in their summary of the family classification of Gelechioidea. However, Heikkilä et al. (2014) discussed the possible distinctiveness of Cryptolechiinae and Cacochroinae, suggesting that Cryptolechia might belong to Lypusidae instead (p. 581). The name Orophiini Lvovsky 1974, also used as the subfamily name Orophiinae, based on Orophia Hübner, [1825], has occasionally been understood to include some or all of the genera Eutorna, Orophia, Cacochroa and Zizyphia (e.g. Tokár et al. 2005; Lvovsky 2009, 2012). Recently Corley and Ferreira (2019) added a new genus, Rosetea, to this group of genera. Orophiinae was synonymized with Cryptolechiinae by Minet (1990), but because it is a junior homonym of Orophiini Thomson, 1863 (Coleoptera: Ciidae) (Bouchard et al. 2011), it is unavailable. Lvovsky (2019) proposed Eutornini as a replacement name for Orophiini. However, because these names have different type genera, Eutornini is not an objective synonym. The name Cryptolechiinae, (originally Cryptolechiidae Meyrick, 1883, with the type species Cryptolechia straminella Zeller, 1852), has been widely applied to these genera (Minet 1990; Kaila 2004; Kaila et al. 2011; Heikkilä et al. 2014; Kim et al. 2016; Wang and Li 2020). The application of this name has, however, been inconstant. In addition to the name-bearing genus Cryptolechia, and the group here treated, it has also been used for Periacmini Lvovsky 2005, whose author originally considered it as a tribe within Amphisbatinae. Heikkilä and Kaila (2010) narrowed the concept Amphisbatidae, this name though proven to be a junior synonym of Lypusidae after inclusion of Lypusa (van Nieukerken et al. (2011), and excluded Periacmini. Periacminae is now a subfamily in Autostichidae (Wang and Li 2020). According to Heikkilä et al. (2014) C. straminella is likely related to Lypusidae, and not to Periacminae or the Cacochroa group of genera. We support the view that C. straminella (for images of the holotype, see http://www2.nrm.se/en/lep_nrm/s/cryptolechia_straminella.html) is not closely related to the genus group treated here, because of the different wing shape, the presence of a well-developed uncus, unilobed gnathos, and absence of lobes attached either laterad of juxta or the base of the valva. As the name Orophiini is unavailable, and Cryptolechiinae is here considered as unrelated, the valid name is Cacochroinae. As we consider Eutorna and Cacochroa to belong to the same tribe, Eutornini Lvovsky, 2019, syn. nov. becomes a subjective junior synonym of Cacochroini Leraut, 1993.

Acknowledgements

We would like to express our gratitude to Peder Skou, Denmark for providing specimens for this work, to Michael Falkenberg, SMNK for providing photographs and lending out material, to Alicia Morente, Spain for commenting on an earlier version of the manuscript, to Francisco Morente, Spain for his advice and contribution on the vegetation of the area, to Ursula Grimm and Peter Huemer, Tiroler Landesmuseum, Innsbruck, Austria and Eddie John, U.K., who helped with literature searches, and to “la Consejería de Agricultura, Ganadería, Pesca y Desarrollo Sostenible de la Junta de Andalucía”, Spain for the granting of the corresponding permits, which have allowed us to carry out our studies. We are grateful to three reviewers and Conrad P.D.T. Gillett, MZH, for commenting on the manuscript and improving the English text.

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