New data on distribution, biology, and ecology of longhorn beetles from the area of west Tajikistan (Coleoptera, Cerambycidae)

Abstract New data on distribution, biology, and ecology of some little-known cerambycid species, collected in the western part of Tajikistan, are presented. Arhopalus rusticus (Linnaeus, 1758) is recorded in Tajikistan for the first time. New localities of species considered pests or invasive species such as Aeolesthes sarta (Solsky, 1871) and Xylotrechus stebbingi Gahan, 1906 are also given. The list of the taxa collected by the first author during many years of field research in Tajikistan as well as photographs of poorly known species from his collection, including some endemics, are additionally provided. Furthermore, high quality photographs of some extremely rare species that were collected during our expedition, i.e., Turkaromia gromenkoi Danilevsky, 2000 and Ropalopus nadari Pic, 1894, with images of their habitats or feeding grounds are also presented for the first time.


Introduction
The longhorn beetle family (Cerambycidae) is one of the most species-rich groups of beetles (Coleoptera) with approximately 35,000 described species (Švácha and Lawrence 2014). The cerambycid fauna of Tajikistan is represented by only 58 species; however, many of them are endemic to the Central Asia region, including those that only occur on the territory of this country (Löbl andSmetana 2010, Danilevsky 2016).
Tajikistan is located almost entirely within the Pamir-Alay Mountain range although forests only cover 2.6% of the area of the country. That places this country on one of the last positions regarding woodiness among the republics of the former Soviet Union. Some relict species such as Juglans regia, Acer turkestanica, and A. reggeli used to occur in Tajikistan in the past but they are totally extinct as a result of intensive cutting and burning of forests (Rahmonov et al. 2003). Due to a low level of afforestation, cerambycid fauna of Tajikistan is characteristic for the countries of the Central Asia. Species of the subfamily Prioninae which larvae develop in soil feeding on plant roots, as well as many representatives of the tribes of Agapanthiini and Phytoeciini (Lamiinae), whose development takes place in stems and roots of herbaceous plants, dominate in fauna of Tajikistan.
The state of the knowledge of the longhorn fauna of Tajikistan (particularly, of the region of the Pamir Mountains) as well as information about the biology and ecology of some species that are distributed in the region is very poor. Therefore, the present study aims to supplement the knowledge in this field.

Study area and methods
Tajikistan is a relatively small intra-continental country situated at the boundary of the subtropical and temperate climatic zones. It is located in the mountain desert zone of the Eurasian continent in the southern part of Central Asia, where ecosystems such as deserts, steppes, savannoides, conifer forests, mixed mountain forests, highmountain deserts, and glaciers are widely represented. The changeable mountain climatic conditions and hard historical natural processes have promoted the formation of a unique biological diversity in Tajikistan, which counts many relict and endemic species (Safarov 2003).
The entomological expedition, which consisted of four scientists from the Department of Zoology, University of Silesia (Poland), took place at the turn of June and July 2014. During the research, several sampling trips were carried out to various locations in the western part of Tajikistan (Map 1). The most extensive studies were conducted within a radius of 60 km from the capital city, Dushanbe, and in the south-western part of the country along the Afghan border. The research were carried out in several research plots in the villages of Arykboshi, Dushanbe, Ganchi, Garavuti, Gharm, Iskanderkul, Kangurt, Karatag, Kolkhozabad, Kulob, Nurobod, Qurghonteppa, Romit, Sarband, Sarichashma, Shahrinav, Shurroabad, Takob, Tojikobod, and Vose. Tajikistan has a wide variety of habitats that range from gravel deserts in the south through green mountain valleys in the central part of the country to the high mountains with glaciers in the north and east. The area studied includes several different nature ecosystems such as alpine meadows, mesophilic shrubs, various shrub steppes, broad-leaf forests, light forests, and tugay as well as agroecosystems such as gardens, orchards, fields, and pastures.
The most effective, standard methods for collecting beetles such as attracting to artificial light sources, shaking down into an entomological umbrella, sweep netting, and rearing of inhabited material were used during the field research. The beetles were studied using an Optek SZM7045-J4L microscope at 7-90× magnifications. Photographs of the cerambycids in nature, their host plants and habitats were taken with Canon EOS 550D, Canon EOS 600D, and Olympus XZ-1 cameras. Produced images were stacked, aligned, and combined using ZERENE STACKER software (www.zerenesystems.com). Geographical coordinates were read off and recorded using Garmin Oregon 550T 3-Inch Handheld GPS Navigator. For each specimen collected, the exact location (including the GPS coordinates), altitude, date, and the names of the collectors are given. Additionally, information on the general distribution and biology of the species are provided. Some general data that had been collected by the first author during long-term field research were also used. The following abbreviations are used in the text: The nomenclature was adopted from the Catalogue of Palaearctic Coleoptera (Löbl and Smetana 2010) with further remarks (Danilevsky 2016).
The specimens are preserved in the entomological collection of the Department of Natural History of the Upper Silesian Museum in Bytom and in authors' collections.

Results
During the one-month expedition, a total number of 12 species (approximately 20% of the Tajik cerambycid fauna) belonging to three subfamilies (Prioninae 1 sp., Cerambycinae 10 spp., and Lamiinae 1 sp.) were recorded. The list of recorded taxa along with the new localities, general characteristics, and remarks on the species biology and ecology follow. Although the species is distributed also in Turkmenistan and Tajikistan, most of the known specimens were collected in the Samarkand region in Uzbekistan. In Tajikistan, it was only observed in the north-western part of the country as far as the southern slope of the Gissar Mountain ridge (Danilevsky 2010). The locality in Sarichashma, which is presented for the first time, extends the species range approximately 200 km to the south-east and is one of the first in the country.
Adults are active from early May to late July. In the hilly area of the Katagurgan environs (Uzbekistan), Danilevsky (2014) observed the mass flight of numerous males in the middle of a hot day (between 11 am and 4 pm) on 12 June 1992. This huge number of males was attracted by females standing motionlessly on the ground. Evidently, the daily activity of this species appears to be a unique behaviour among the representatives of the subfamily Prioninae, which is probably correlated with the small size of the eyes in this species in both males and females (Danilevsky 2014). The larvae develop in soil, where they probably feed on roots of various plants. According to Danilevsky (2014), the larva found in Uzbekistan feeding on roots of Taraxacum kok-saghyz and described as P. turkestanicus by Švácha and Danilevsky (1987) certainly belongs to another species common in that region: Psilotarsus hirticollis hirticollis Motschulsky, 1860. Therefore, the larval stage of P. turkestanicus has not been described yet.
According to Danilevsky (2010), species of the genus Psilotarsus Motschulsky, 1860 are often characterised by a very high degree of individual variability. This was also confirmed by us during examining of the collected specimens, which differed, inter alia, in details of elytra, punctuation of the pronotum, shape of the scutellum and size of the spikes on the pronotum.
In the Sarichashma environs, the specimens were collected in a semi-ruderal plant community (Fig. 3B, C) characterised by a variety of plants, including single trees. We observed flights of single males at about noon, although most of the specimens moved on the ground where they were fighting with each other and seeking females. Flying specimens sometimes became the prey of the European roller Coracias garrulous, which is a very common species in some regions of Tajikistan. The beetles emit characteristic, audible sounds that make it possible to detect their presence. We noticed many circular exit holes in the ground, which probably belong to this species. It is also noteworthy that the culmination of the occurrence of this species probably took place shortly before or during our first visit to this plot on 25 June. However, when we came back to this location on 3 July, we did not find even a single specimen.
Because this site is located directly on the Tajik-Afghan border, it can be expected that this species will also occur in Afghanistan. On the other hand, the bordering Panj River may form a natural migration barrier, particularly for the females, which are probably flightless. It is noteworthy that two other related Prionini species, Miniprionus pavlovskii (Semenov, 1935) and Pogonarthron semenovianum (Plavilstshikov, 1936), were also recorded in the immediate vicinity of the plot mentioned above (Danilevsky 1999, Lorenc unpublished data).
The authors feel compelled to state that this plot is located in a strongly guarded zone just a few kilometres from the Afghan border. No foreigners are allowed to enter this area without the proper permits. Staying in this zone (especially at night) can have serious consequences from both the Tajik authorities and Taliban fighters from Afghanistan due to the large drug route in the region.

Cerambycinae Latreille, 1802
Trichoferus campestris (Faldermann, 1835) Trichoferus campestris is considered an invasive species, which has rapidly increased its range in recent years. Its presence in Europe has recently been confirmed inter alia in Romania (2003) (Dascălu and Serafim 2011), the Czech Republic (2006), Slovakia (2007) (Sabol 2009), andPoland (2009) (Kruszelnicki 2010). However, the species appears to be native to China, Japan, the Korean peninsula, Mongolia, Russia, and Central Asia, including Tajikistan (Grebennikov et al. 2010, Dascalu et al. 2013. The larvae are polyphagous on both deciduous and coniferous trees, although the species seems to prefer light deciduous forests and especially orchards. The adults are active at night from June to August and are often attracted to artificial sources of light (Kadyrov 2007). The city longhorn beetle A. sarta is a species widely distributed throughout the Palaearctic and the Oriental region. It is believed that it originated in Pakistan and Western India from which it spread to Afghanistan, Iran, and to Central Asia (Orlinskii et al. 1991).

Aeolesthes (Aeolesthes) sarta (Solsky, 1871)
The species is polyphagous with a wide range of host plants and it primarily attacks tree trunks. For this reason, the species is considered a serious pest in the countries in which it occurs. In many cities of Central Asia, poplars and willows have been destroyed as well as plane trees, acacias, and ashes in Dushanbe (Kadyrov 2007). It takes two years for the larvae to develop. The adults overwinter in a pupal cell and emerge in the following spring. The flight period of adults begins in the second part of April and lasts more or less until mid-July. Imagines are generally active in the evening and night and very often are attracted to sources of light (Kadyrov 2007).
It was observed that A. sarta attacks both maximally exposed and shaded trees. The larval feeding grounds (Fig. 3D) of this species were additionally found in two other locations: Garavuti env. [37°35'N, 68°31'E] and Shahrinav env. [38°36'N, 68°19'E]. In above-mentioned Romit area, in addition to the imagines that were collected, about 20 larvae of A. sarta (Fig. 3E) were also found on a dead trunk of Prunus sp. (Fig. 3F). It is distributed in several countries of Central Asia, Iran, Afghanistan, and western China (Kadyrov 2007, Danilevsky 2016. It is a common species in Tajikistan and occurs everywhere its host plant ferule (Ferula spp.) grows (Kadyrov 2007). In Tajikistan, the genus Ferula consists of nearly 40 species (Safarov 2003). According to Plavilstshikov (1940), larvae of N. scapularis may also develop in species of the genus Scorodosma. The larvae primarily feed on the rhizomes and roots of these plants and their development usually takes a year, sometimes even two (Plavilstshikov 1940). Pupation takes place in calcareous cocoons in the soil (Švácha and Danilevsky 1988). The adults appear and feed on the flowers or stems of the host plants from the end of April to July, depending on the local altitude (Kadyrov 2007).

Neoplocaederus scapularis (Fischer von
The beetles are probably active in the evening and at night. We only observed adults on the ferule (Fig. 3G, H) in the early morning, due to the fact that the beetles still had not managed to hide after the night. Moreover, during the research, imagines were often attracted to an artificial light source. The genus Turkaromia Danilevsky, 1993 was quite recently separated by Danilevsky (1993) and includes two species, Turkaromia pruinosa (Reitter, 1903) and T. gromenkoi, which are distributed in the region of Central Asia. According to , T. gromenkoi is distributed in the western part of the Gissar Mountain ridge in Uzbekistan and Tajikistan. The species was described from four specimens: one male and two females from Kaltakol (Uzbekistan) and one female from Iskanderkul (Tajikistan). All specimens were observed in July. The biology and ecology of species as well as the stages of the larvae and pupae are unknown.

Turkaromia gromenkoi Danilevsky, 2000
In the environs of the Iskanderkul Lake, we observed one female on a flower (Apiaceae) in a biotope near a river valley that had been overgrown by willows (Salix spp.) and shan birches Betula tianschanica (Fig. 4A). The larvae probably develop in the living wood of willows similar to the related species T. pruinosa. In the immediate vicinity of the area where the beetle was collected, we found sawdust-like waste on the outside of the trunk of a middle-aged willow (Fig. 4B), which was probably the result of the larval feeding of T. gromenkoi.
It is noteworthy that only one specimen was found despite a few hours of examining the plot using various methods. The presence of only a single female may indicate the end of the period of the occurrence of this species. It appears that T. gromenkoi is endemic to the Gissar Mountains. This species occurs in Kyrgyzstan, Tajikistan, and Uzbekistan (Danilevsky 2016) and is endemic to the region of western Tian Shan Range (Kadyrov 2007). R. nadari is a polyphagous species, which usually inhabits growing wild fruit trees in the upper zone of deciduous forests (Fig. 4D). This species most frequently inhabits trunks and boughs of the wild apple tree Malus sieversii. Its larval development usually takes two years (Plavilstshikov 1940, Kadyrov 2007. Adults are found from June to July. After they emerge, the imagines are unwilling to fly and generally stay on the host plant and only visit flowers of Ferula spp. and Prangos spp. occasionally (Kadyrov 2007).

Ropalopus
Only a single female was observed despite a 24-hour monitoring of the trees on the plot. This may either be related to the end of the period of the occurrence of the species or its hidden life in the treetops. This species is distributed in the countries of Central Asia and the Xinjiang region of China (Danilevsky 2016). It inhabits the upper zone of deciduous forests and, less frequently, valleys. Although T. pilosum is polyphagous on deciduous trees (Kadyrov 2007), the larvae can also feed on conifers (Danilevsky 2001a). The species inhabits dry twigs and stems (Fig. 4E). Its development usually takes two years with pupation in spring. The adults fly from April to August (Danilevsky 2001a, Kadyrov 2007.
Mating of this species seems to start at the end of June. We observed adults flying into wooden components and actively moving on trunks of the wild apple tree Malus sieversii, where they were also mating. T. pilosum was recorded sympatrically with R. nadari in the same habitat (Fig. 4D). This is a widely distributed species, whose origin is not clear (Cocquempot and Lindelöw 2010). It probably originally came from the region of northern India (Himalayas, Tibet). In recent years, this invasive species has spread to and become acclimated in the Middle East, the Mediterranean region (Sama 2002), and possibly in Central Asia. In Europe, it was recorded for the first time in Italy in 1990 (Dioli and Viganò 1990). The larvae are polyphagous on broad-leaved trees. Its life cycle lasts two years. Adults are usually encountered between May and November (Sama 2002, Ali et al. 2015.

Xylotrechus (Xylotrechus) stebbingi Gahan, 1906
It was observed that this species appears to be strongly synanthropic in Tajikistan: adults were found in various anthropogenic environments such as backyards and orchards, where they willingly flew to artificial light sources. This species occurs in Kyrgyzstan and Tajikistan (Danilevsky 2016), where it is widely distributed in both mountain areas and valleys. It is polyphagous on deciduous trees. The larvae feed subcortically on recently dead twigs or branches. Larva overwinters and then it pupates in the wood in the following spring; sometimes the imago overwinters. The adults fly from April to June and feed on flowers, especially those of Cerasus, Prunus, Malus, Pyrus, Rosa, Cotoneaster, Atraphaxis, and Exochorda (Kadyrov 2007).  This species is distributed from the Caucasus to Central Asia (Danilevsky 2016) where it mainly inhabits floodplains and riparian forests (tugay) (Fig. 4F, G). The larvae feed on the dead wood of various deciduous trees. They were recorded inter alia on Elaeagnus, Halimodendron, Caragana, and Robinia. The life cycle of this species usually takes two years with pupation in spring and early summer. The adults feed on various flowers from April to July (Svácha and Danilevsky 1988, Kaliuzhnaja et al. 2000, Kadyrbekov and Tleppaeva 2004. This species is distributed in the Caucasus, the Far East, Central Asia, and Oriental region (Danilevsky 2016). It occurs in valleys as well as in mountain areas up to 2500 m. Like the previous species, it is rather common in a tugay habitat. It is polyphagous on deciduous trees, mostly on poplars and willows (Kadyrov 2007), although it also inhabits tamarisks Tamarix, oleasters Eleagnus, elms Ulmus and pears Pirus Danilevsky 1988, Shapovalov 2012). Furthermore, its larvae also develop in wooden structures, which makes Ch. faldermanni one of the most serious pests of timber in Central Asia. Its life cycle lasts one or two years. Its flight period is from May to September Danilevsky 1988, Kadyrov 2007). The adults frequently visit flowers, especially Apiaceae (Kadyrov 2007).

Lamiinae Latreille, 1825
Agapanthia ( This species occurs in Tajikistan, Uzbekistan, Kyrgyzstan, and Kazakhstan (Danilevsky 2016). It is common in alpine meadow at altitudes of between 1000 and 3000 m (Fig. 3G). The larvae feed on the stems and rhizomes of Prangos spp. The adults  feed on host plants from May until the end of July, although they were also observed on mallows Malva spp. The imagines usually stay on the stems of host plants, where they copulate and supplementary feed during summer. After that, the females lay eggs in the incisions in the stems. A female usually lays only one egg on each stem. In 8 to 12 days, the larva hatches and bites into the core of the stem where it moves towards the root. The larva forms a pupal cell and overwinters in the lower part of stem or the upper part of root. Pupation occurs in the spring (Kadyrov 2007).
A massive mating of this species was observed in the Takob environs between 8 and 10 July. The beetles performed characteristic slow flights during the day. It is noteworthy that no more species of the genera Agapanthia or Phytoecia were caught during the entire expedition, despite the very frequent use of the sweep-netting method in appropriate habitats (e.g. alpine meadows) in various parts of the country. This species seems to occur much later or longer than the other related species.

Checklist of the Cerambycidae of Tajikistan
The followed list is based on Danilevsky (2016). Species collected by the first author over many years of research are marked with an asterisk (*). Endemic taxa are marked with a letter (E).

Discussion
Knowledge about the fauna of Tajikistan, particularly invertebrates, is still poor. Species of beetles that are new to science are increasingly being described, for example, Kytorhinus kergoati Delobel & Legalov, 2009 (Chrysomelidae) (Delobel and Legalov 2009 Longhorn beetles arouse great interest among beetle families, and therefore the current state of knowledge on Tajik cerambycids appears to be better than that for other groups. However, the knowledge about the species that occur in this country is still insufficient; thus, some new taxa have been described in recent years: Turkaromia gromenkoi , Pogonarthron petrovi petrovi (Danilevsky 2004), P. petrovi ivanovae (Pak and Skrylnik 2014), and Agapanthia ustinovi .
The distribution of species within the individual subfamilies is characteristic for Central Asia and it is presented as follows: Prioninae (9), Lepturinae (2), Spondylidinae (1), Apatophyseinae (2), Cerambycinae (20), and Lamiinae (25). Species of the subfamily Prioninae, whose larvae develop in soil and feed on plant roots, as well as many representatives of the tribes of Agapanthiini and Phytoeciini (Lamiinae), whose development take place in the stems and roots of herbaceous plants, dominate the fauna of Tajikistan. Representatives of the subfamily Lepturinae are few in number possibly due to the lack of trees and, consequently, small amounts of deadwood, which is quite a normal situation in Central Asia. For example, the total number of Lepturinae species is: 5 (Uzbekistan), 6 (Turkmenistan), 9 (Kyrgyzstan), and 4 (Afghanistan).
Because of the many geographical barriers in Tajikistan, a high level of endemism occurs in this group of beetles. Currently among the 60 taxa, as many as eleven taxa occur exclusively in Tajikistan. The endemics include Mesoprionus zarudnii, Pogonarthron semenovianum (all known specimens were only collected in Tajikistan, near the Afghan border (Danilevsky 2004), but according to Danilevsky (2016) this species occurs also in Afghanistan), P. bedeli, P. petrovi petrovi, P. petrovi ivanovae, Miniprionus pavlovskii, Cleroclytus gracilis, Agapanthia ustinovi, Mallosiola regina, Oberea ruficeps muchei, and Phytoecia kubani.
Due to its climatic conditions, Tajikistan is a very unique place to collect insects. The flight period of various groups of longhorn beetles is quite diverse. Some species only occur in spring, for example the genus Phytoecia, which is represented here by as many as 13 species. On the other hand, some species of the subfamily Prioninae begin to fly in late summer. There are also many nocturnal species that lead very cryptic life-styles. For these reasons, comprehensive studies that include the entire growing season are needed to obtain a reasonably full and true picture of the composition of the species of longhorn beetles in this country.
Today, the industrial and economic activities of humans are the most important factors that influence ecosystems. Such anthropogenic activities cause important changes in fauna and flora that lead to the simplification of biogeocenosis structures and to decrease in the differences between landscape zones. Some agrotechnical measures, such as ploughing desert regions of a country, deforestation, intensive irrigation, chemical use, and the development of industry in large parts of southeast Central Asia have induced the development of specific fauna in the anthropogenic landscapes. Among the most important forms of human activity are ploughing and managing soils that had not been used earlier, which causes the formation of secondary biocoenosis and agrocoenosis, which in turn leads to disturbances of the ecosystems and, as a result, to the disappearance of many vulnerable species. Planting trees and shrubs in disturbed regions is extremely important due to their role as an ecological corridor (Rahmonov et al. 2003).
Other than the extensive research activities of the first author (e.g. Kadyrov 1989Kadyrov , 2007 and Danilevsky (e.g. 2001a, b), we do not know of any other studies on Cerambycidae in Tajikistan, with the exception of the activity of beetle collectors. In spite of the fact that access to the region of the Pamir Mountains is severely limited and probably requires special permits for entry, it seems to be particularly interesting in the context of research on longhorn beetles. We still do not know enough about the biology of the local endemic species and there are probably some species that are as yet undiscovered, and therefore new expeditions to this region are quite desirable.