New distributional records for sixteen Mordellidae species from the Western Palearctic (Insecta, Coleoptera, Mordellidae)

Abstract A list of 22 new distributional records is presented for 16 Mordellidae species from the Western Palearctic: Variimorda caprai (Franciscolo, 1951) (Montenegro); V. mendax Méquignon, 1946 (Montenegro); Mordellistena falsoparvula Ermisch, 1956 (Bosnia and Herzegovina, Montenegro); M. olympica Ermisch, 1965 (Cyprus, Montenegro); M. kraatzi Emery, 1876 (Morocco); M. longicornis Mulsant, 1856 (Morocco); M. dives Emery, 1876 (Kazakhstan); M. krujanensis Ermisch, 1963 (Montenegro); M. tarsata Mulsant, 1856 (Cyprus, North Macedonia); M. michalki Ermisch, 1956 (Kyrgyzstan); M. thuringiaca Ermisch, 1963 (Bulgaria, Montenegro, Slovakia, Spain); M. koelleri Ermisch, 1956 (Italy, Montenegro); Mordellistenula longipalpis Ermisch, 1965 (Montenegro); Mordellochroa milleri (Emery, 1876) (Italy); Dellamora palposa Normand, 1916 (Italy). Information about the distributional range is summarised for each species, and notes on habitat and host plants are also provided.


Introduction
Mordellidae Latreille, 1802 is a rather diverse but poorly known and sporadically studied family of beetles. It comprises more than 2300 described species distributed nearly worldwide except for the polar and subpolar zones. Mordellid beetles inhabit various ecosystems including tropical rainforests, temperate deciduous forests, grasslands and ruderal ecosystems. The majority of species are pollinivorous in adult stages, feeding on a wide variety of plant species. Exceptions are the members of the genus Glipa Leconte, 1859, reported to feed on fern spores (Takakuwa 2000), or the South American Boatia albertae Franciscolo, 1985 which was found to have the foregut filled with fungal spores (Lawrence and Ślipiński 2010). Larvae of some mordellid beetles are wood borers (e.g., Mordella Linnaeus, 1758, some Mordellistena Costa, 1854), the others develop in sporocarps of Polyporaceae fungi (e.g., Curtimorda Méquignon, 1946, Mordella marginata Melsheimer, 1845, or in stems of various herbaceous plants (e.g., Mordellistena Costa, 1854, Mordellistenula Shchegoleva- Barovskaya, 1930). In some cases, mordellid larvae seem to have predaceous habits against other insect larvae (Tooker and Hanks 2004) or can be inquilines of termite colonies (Hill 1922).
The majority of distribution records available on Palearctic Mordellidae are usually included in publications focused primarily on the taxonomy (e.g., Ermisch 1963bErmisch , 1965Ermisch , 1977Horák 1985;Plaza 1985). Several authors have summarised the distributional information on regional level (e.g., Ermisch 1956Ermisch , 1963aErmisch , b, c, 1969bErmisch , 1977Köstlin and Vogt 1971;Batten 1976b;Franciscolo 1995;Odnosum 2003Odnosum , 2010Zemoglyadchuk 2007;Ruzzier 2013;Ruzzier et al. 2017). Comprehensive catalogues were published, for instance, by Heyden et al. (1906), Csiki (1915), and Horák (2008). Despite the great effort of these authors, the knowledge of the distribution of many Palearctic species can be still considered poor, and several species remain to be known only from the localities stated in the original description.
Herein we provide new distributional and bionomical data obtained during the recent collecting activities of both authors (2010)(2011)(2012)(2013)(2014)(2015)(2016)(2017)(2018) and by the re-examination of the material deposited in museums or private collections listed below. We also summarise the previously published distributional data and provide the lists of countries from which the species have been previously reported.

Materials and methods
The present study is based on the material sampled by the authors or accessed from the museums and private collections listed below. Specimens were collected individually from flowers or by sweeping the vegetation. Larvae of Mordellistena species were reared from stems of herbaceous plants collected in April, cut to the smaller pieces (ca. 20 cm), transferred to the laboratory, and stored in plastic containers with air access. Adult specimens were killed in ethyl acetate. Dry specimens were relaxed in water with a small amount of acetic acid, then dissected and glued on cards. Dissected genitals were glued with the respective specimen using dimethyl hydantoin formaldehyde (DMHF) or stored in micro-vials containing glycerol and pinned under the specimen. Habitus images were taken by Cannon 5D mark IV attached to Zeiss Axio-Zoom V-16 stereomicroscope with diffuse LED lighting, stacked in Zerene Stacker 1.04 and subsequently edited in Adobe Photoshop CC.
Identifications were carried out using original species descriptions, identification keys (e.g., Ermisch 1956Ermisch , 1963bErmisch , 1969aErmisch , 1977 or by comparison with the type material. Each specimen was labelled with identification data containing full species name, name of the identifier and year when the identification was made. Examined specimens are deposited in the following collections: In the section "New records", we provide the data on examined material concerning only those countries from which the species are reported for the first time. The "Distribution" section contains the list of countries from which the species have been previously reported, followed by the citation of its first appearance in the published sources. In the "Remarks" section, we provide information about habitat preferences, host plants and collecting circumstances (if available).

Results
In the list below, we present 22 new country-level records of 16 Mordellidae species. The list is based on 201 examined specimens from thirty localities in the Western Palearctic. We also provide information about distribution and bionomy for each species. (Franciscolo, 1951)   Distribution. Albania (Franciscolo 1951), Montenegro (new country record). Remarks. Variimorda caprai was described based on three male specimens from Shkodër, Albania and since then no other record has been published. In 2011, the first author collected 83 specimens in the environment of Bar in Montenegro on the slopes with xerothermophilous grassland vegetation (Fig. 2). Specimens were found feeding on the flowers of Helichrysum. The immature stages and their host plants remain unknown.
Remarks. Variimorda mendax inhabits various grassland and ruderal habitats. Adults are usually found feeding on the flowers of Daucus carota. Larvae and host plants remain unknown.
Remarks. This species inhabits xerothermophilous and mesophilous grasslands and ruderal vegetation where the adults feed on flowers of herbaceous plants, e.g., Chrysanthemum, Cirsium, Achillea (Borowiec 1996). The larva was described by Odnosum (1983Odnosum ( , 2010) and reported to develop in stems of Artemisia absinthium and A. vulgaris. The first author reared adults from stem of Apiaceae plant infested by larvae in southern Slovakia.

Mordellistena
Remarks. This species has a wide distributional range in Europe, North Africa and the western parts of Asia. It inhabits xerothermophilous grassland habitats where adults feed on flowers of various herbaceous plants (e.g., Daucus carota, Tithymalus). The larvae described by Odnosum (2010) feed on the stems of Asteraceae plants (e.g., Arctium, Centaurea salonitana).
Remarks. Adults have been reported feeding on flowers of Thapsia villosa in Spain (Plaza 1983). The immature stages are unknown.
Remarks. This species inhabits steppes and xeric grasslands where adults feed on flowers of Apiaceae and Euphorbiaceae (e.g., Seseli, Tithymalus). The larvae and host plants remain unknown. Specimens examined for the present study were compared with the type series deposited in SNSD.
Remarks. The first author collected adults in Mediterranean xeric grasslands and ruderal vegetation along roadsides and field margins in Bulgaria, Montenegro (Fig. 2) and Cyprus (Fig. 3). The specimens were feeding on flowers of Daucus carota. The immature stages and host plants remain unknown. Specimens examined for the present study were compared with the type specimens deposited in SNSD.

Mordellistenochroa fallaciosa
Remarks. Up to present, M. fallaciosa is known only from several localities in Italy, Hungary and Switzerland (Ermisch 1969c;Merkl and Németh 2008;Horák 2008). Although this species was originally described from Italy by Ermisch (1969c), we provide here another record of this rare species, increasing the information about its distribution in the country. The immature stages and their host plants remain unknown. Ermisch, 1965 Fig. 5A Mordellistenula longipalpis Ermisch, 1965: 256-259 [type locality: Belgrader wald, Turkey].
Remarks. This species is known from the Balkans and western Asia. It inhabits dry grasslands where adults feed on flowers of herbaceous plants. The first author observed adults on Mediterranean xeric grassland along the seashore on flowers of Helichrysum in Montenegro (Fig. 2), and in ruderal vegetation along roadsides on flowers of Daucus carota in Bulgaria. The immature stages and their host plants remain unknown.
Remarks. Mordellochroa milleri usually occurs in beech, oak or floodplain forests, and on its margins, but it was also reported from coniferous forest with spruces and pines in Bialowieza National Park, Poland (Kubisz 2000). Adults were found feeding on flowers of Apiaceae (e.g., Libanotis montana) and Brassicaceae (Cardaria draba) plants. Larvae feed in dead wood.

Discussion
The Palearctic fauna of Mordellidae consists of approximately 700 described species (Horák 2008;Odnosum 2009;Takakuwa 2010;Horák et al. 2012;Ruzzier and Kovalev 2016;Tsuru 2017;Selnekovič and Kodada 2019). The most recent comprehensive cata-logue of Palearctic Mordellidae was provided by Horák (2008). Since then several new country-level records were published in the catalogues focused on smaller geographical areas (Abdul-Rassoul 2010;Serrahima 2011;Horák et al. 2012;Ruzzier 2013;Ruzzier et al. 2017) or as single-species records (Horák and Háva 2008;Ferenca and Tamutis 2009;Odnosum 2009;Viñolas et al. 2009;Diéguez Fernández 2010;Sanchez et al. 2015;Viñolas et al. 2016;Selnekovič and Kodada 2019). Although the distribution of Mordellidae species has been studied mostly in the western Palearctic, the recent fieldwork combined with efforts to re-examine the material deposited in major European collections have revealed 22 new country-level records for 16 species. Such results suggest that our knowledge of the distribution of Mordellidae species can be still considered poor and that continuous sampling effort will lead to further new and interesting records.
Furthermore, huge gaps exist in our knowledge of the bionomy of these beetles. Information about habitat preferences and host plants is scarce. Most of the larval descriptions and host plant records were provided by V. K. Odnosum (e.g., Odnosum 1983Odnosum and Litvin 2009). Despite his and other authors' great effort, the number of species for which larval stages are described is relatively low. The general lack of information makes it extremely difficult to identify the larvae based solely on their morphology. The use of DNA barcoding should simplify the species identifications in the future and reveal new information about host-plant relationships and habitat preferences.
Each published record represents a small step to better understanding the distribution and bionomy of mordellid beetles, and their role in the ecosystems. Such information is becoming still more critical, especially in the current age of global environmental changes and loss of natural habitats.