The tiger beetles (Coleoptera, Carabidae, Cicindelinae) of Israel and adjacent lands

Abstract Based on field studies, museums collections and literature sources, the current knowledge of the tiger beetle fauna of Israel and adjacent lands is presented. In Israel eight species occur, one of them with two subspecies, while in the Sinai Peninsula nine species of tiger beetles are now known. In the combined regions seven genera from two tribes were found. The Rift Valley with six cicindelids species is the most specious region of Israel. Cylindera contorta valdenbergi and Cicindela javeti azari have localized distributions and should be considered regional endemics. A similarity analysis of the tiger beetles faunas of different regions of Israel and the Sinai Peninsula reveal two clusters of species. The first includes the Great Rift Valley and most parts of the Sinai Peninsula, and the second incorporates most regions of Israel together with Central Sinai Foothills. Five distinct adult phenological groups of tiger beetles can be distinguished in these two clusters: active all-year (three species), spring-fall (five species), summer (two species), spring-summer (one species) and spring (one species). The likely origins of the tiger beetle fauna of this area are presented. An annotated list and illustrated identification key of the Cicindelinae of Israel and adjacent lands are provided.


Introduction
The first data about tiger beetles of Palestine were published in the first third of the XXth century. In 1913 Sahlberg described from Wadi El Nawaime (modern Wadi en Nu'eima) Cicindela littoralis aulicoides. In 1934 Mandl recorded for the Palestine two subspecies of Cicindela littoralis: C. l. winkleri and C. l. aulicoides. The first species list of Palestinian Coleoptera including five species of tiger beetles was published by Bodenheimer in 1937. Around the same time, the first information about cicindelids of the Sinai Peninsula appeared and Cicindela aulica (Horn, 1931), Cicindela littoralis aulicoides (Mandl, 1934) and Megacephala euphratica (Schatzmayr, 1936) were recorded. Unfortunately, detailed locality data and collecting dates for specimens of these species were often incomplete.
A second wave of tiger beetles studies in the Levant was completed in the last third of XXth century. Alfieri (1976) published the catalogue of Egyptian Coleoptera with information about 11 species of tiger beetles, six of which were recorded for the Sinai Peninsula. The first data about Cicindelinae of Israel were published by Valdenberg (1983Valdenberg ( , 1985 and Nussbaum (1987). It should be noted that these papers also contained information about tiger beetles of the Sinai Peninsula. In all eight species were recorded from Israel and seven species for the Sinai. Unfortunately, in the paper by Nussbaum (1987) data about localities for several species given in the text and on the maps do not coincide.
During the last decade, new information about the distribution of tiger beetles in different parts of the Levant has accumulated, and we include these new records here.

Material and methods
Specimens and data for this report come from the following museums and private collections:

TAU
Tel Aviv University (Israel); ZMUM Zoological Museum of Moscow State University (Moscow, Russia);

Distribution
With these current records, eight species of tiger beetles, one of them with two subspecies, belonging to seven genera of two tribes are known from Israel (  Fig. 7), while L. flexuosa flexuosa is the second most widespread species of tiger beetles absent only from northern (Galilee, Golan Heights) and central (Samaria, Judea) regions (Table 1, Fig. 6). Lophyra flexuosa (Fabricius, 1787) reaches the eastern limit of its distribution in Israel.
The nominative subspecies of Cylindera contorta (F.-W., 1828) is widely distributed in Central Asia, some regions of Cis-and Transcaucasia as well as in the northern and western sides of the Black Sea from southern Russia to Romania (Wiesner 1992;Cassola 1999;Puchkov and Matalin 2003), however it is not known from Anatolia (Corel 1988;Cassola 1999;Puchkov and Matalin 2003;Avgin and Özdikmen 2007), Syria (Wiesner 1992;Puchkov and Matalin 2003;Avgin and Wiesner 2009;Jaskuła and Rewicz 2014), Jordan (Wiesner 1992;Puchkov and Matalin 2003), Iraq (Ali 1978;Wiesner 1992;Puchkov and Matalin 2003) and Saudi Arabia (Wiesner 1992;Cassola and Schneider 1997;Puchkov and Matalin 2003;Al Ahmadi and Salem 1999). The populations of C. contorta valdenbergi inhabit the Mediterranean coast from 'Akko (Northern Coastal Plain) to Bat Yam (Central Coastal Plain) in Israel (Nussbaum 1987; our data) as well as between Ras El Bar and Abu Qir in north-eastern Egypt (Alfieri 1976;Abdel-Dayem et al. 2003) are distinctly scattered and bound the southwestern limit of the distributional area of C. contorta as a whole.  Cicindela javeti azari has a restricted distributional area and now is known only from southern Lebanon (Deuve 2011), southwestern Syria (Avgin and Wiesner 2009) as well as northern regions of Israel: Upper Galilee and Golan Heights (Nussbaum 1987; our data). Among three known subspecies (Deuve 2011) C. javeti azari inhabits the southern part of the species range area.
Cephalota zarudniana vartianorum lives from south-eastern Iran across Iraq and Syria to Jordan and Israel (Wiesner 1992;Puchkov and Matalin 2003). The Dead Sea Area is the western border of the distributional area both for this subspecies as well as for the species as a whole.
It should be noted that the three mentioned above subspecies were recorded in Israel only during XX century (Fig. 4), and the latest records are dated from the late 80's to the early 90's.
The Sinai Peninsula is the most diversity of tiger beetles region from all neighbouring territories by Israel because nine species live here, and C. tibialis tibialis, C. littorea littorea, H. singularis and H. nilotica nilotica are never really observe in Israel (vs Chikatunov et al. 2006). Among them C. tibialis tibialis is an endemic of Egypt and occurs along Mediterranean Sea coast in the Governorates Matrouh, Alexandria, Kafr el-Sheikh, Damietta, Port Said and North Sinai (Gebert 1991;Abdel-Dayem et al. 2003;Abdel-Dayem 2012). Moreover, C. littorea littorea is an regional near-endemic living along Red Sea coast in Egypt and Saudi Arabia (Gebert 1991;Cassola and Schneider 1997;Abdel-Dayem et al. 2003). Arabian-African Hypaetha singularis lives along Red Sea coast in Egypt, Sudan, Eritrea and Yemen, and on the shore of Gulf of Aden in Djibouti, Somalia and Yemen (Wranik et al. 1991;Werner 2000;Wiesner 2002Wiesner , 2005 as well as on the littoral of Arabian Sea in Oman (Cassola and Rihane 1996). The Sinai localities are limited the northern border of the distribution area of this species. African Habrodera nilotica nilotica is widely distributed in Afrotropical Region (Wiesner 1992;Werner 2000). Two known localities from Sinai Mountains (Alfieri 1976;Abdel-Dayem et al. 2003;Abdel-Dayem 2004) are limited the distribution range of this species to the east.
According to the analysis of the similarity between faunas of tiger beetle of natural regions of Israel and the Sinai Peninsula two large clusters are recognized (Fig. 101).

Phenology
According to the literature data (Alfieri 1975;Nussbaum 1987;Abdel-Dayem et al. 2003) and the results of our own study some aspects of the phenology of tiger beetles both in Israel and on the Sinai Peninsula are discussed. The period of activity of the beetles but not the breeding period was analysed first of all. As a result, five groups of the tiger beetles were obtained (Table 2). Three species with the longer period of activity from January to November or from February to December belong to the all-year group. Five species, including two subspecies of Calomera littoralis (F., 1787), characterized by the prolonged period of activity from February to October-November, from March-April to November or from March to December and form the richest springfall group. Two species recorded only on the Sinai Peninsula with the period of activity from May to August-September are composed the summer group. At last, both the spring group (activity from February to May) and the spring-summer group (activity from February to August) contain a single species each.
It should be noted that the period of activity of some studied species does not correspond with the data of previous studies in Israel (Nussbaum 1987) and on the Sinai Peninsula (Alfieri 1975;Abdel-Dayem et al. 2003), as well as in the other parts of the distribution area . For example, the activity of C. aulica aulica, C. littoralis aulicoides, C. zarudniana vartianorum, C. contorta valdenbergi and G. euphratica euphratica start one-two months earlier, while the activity of C. aulica aulica, C. littoralis aulicoides, M. melancholica melancholica and G. euphratica euphratica finish one-three, and in the case with L. flexuosa flexuosa even six months later comparing with the data of Nussbaum (1987). On the other hand, Nussbaum (1987) indicated longer period of activity of C. littoralis winkleri and C. tibialis tibialis as well as the later finish of the activity of C. contorta valdenbergi and C. javeti azari (Table 2).
Similarly, the periods of activity of C. aulica aulica, L. flexuosa flexuosa and M. melancholica melancholica in the central and southern Levant as well as on the Sinai Peninsula are appreciably longer than in the Maghreb region. So, in Tunisia C. aulica aulica records only in June and July , while in Israel it active from March to December and on the Sinai Peninsula from February to October (Table 2). Both in Tunisia and Morocco the period of activity of L. flexuosa flexuosa lasts from March-April to July  but in Israel it continues from February to December (Table 2).
On the contrary, in Tunisia the activity of G. euphratica euphratica begins in March and ends in July  that is similar with the period of activity in Israel and on the Sinai Peninsula (Table 2), while in Morocco it takes only three months -from June to August . The same situation is observed for different subspecies of Cephalota littorea (Forskål, 1775) as well as C. littoralis. In Tunisia C. littorea gouditii (Dejean, 1829) is active from May to October  while the period of activity of C. littorea littorea on the Sinai Peninsula lasts from May to September (Table 2). The activity of C. littoralis littoralis in Morocco is observed from April to October  and in Tunisia from March to August , while the activity of C. littoralis aulicoides in Israel and on the Sinai Peninsula as well as C. littoralis winkleri in Israel occurs from February to October and from February to November, respectively (Table 2).
However, we must remember that the obtained data are compilative. The differences in the time and the density of sampling, the collection technics as well as the frequency of visit of the particular localities and habitats could really distort the real pattern.

Faunogenesis
The tiger beetle fauna of Israel as well as the Levant as a whole is complex. In geological time these areas were settled by species from different Mediterranean, African and Asiatic regions.
Unfortunately, the information about fossil Cicindelinae is extremely scant (Nagano et al. 1982). At present time South American Oxycheilopsis cretacicus Cassola & Werner, 2004 (Lower Cretaceous ca. 125 Ma) is the oldest known fossil tiger beetle (Cassola and Werner 2004). Three samples of fossil cicindelids are known from the northern Europe Baltic Amber (Oligocene ca. 23-34 Ma). Despite the identification ambiguity of the species, the genera were interpreted as the recent ones (Nagano et al. 1982;Röschmann 1999) as most known fossil Carabidae and other Coleoptera (Alekseev 2013). All other fossil records of the tiger beetles from the Europe and northern America (USA and Canada) are dated from the Quaternary period from Pleistocene to Holocene, and all other species are interpreted as recent (Nagano et al. 1982).
By analogy with other groups of carabid beetles (Kataev 1984(Kataev , 2011Casale and Vigna Taglianti 1999;Ruiz et al. 2012), we can assume that the genesis of the ancestral taxa of most recent cicindelids in the Mediterranean region began in late Paleogeneearly Neogene (on the border of Oligocene -Miocene). According to data of DNA analysis the divergence processes of taxa of subtribe Cicindelina began ca. 15-25 Ma with most intensity between 2-10 Ma (Barraclough and Vogler 2002;Pons et al. 2004;Tsuji et al. 2015). For example, the diversification of the species within Cicindela hybrida group started ca. 2 Ma (Cardoso and Vogler 2005), while the separation of the genus Cosmodela Rivalier, 1961 from other Cicindelinae took place ca. 2.2-5 Ma (López-López et al. 2015;Tsuji et al. 2015). Based on the fossil material we could be argued that at least 60,000-70,000 yrs. BP the recent species of tiger beetles were already presented both in the North America and in the Eurasia (Nagano et al. 1982).
The continental drift of the Arabian and Anatolian Plates, their collision and, as the result, closing the Neotethys Ocean during Oligocene-Miocene were the most important processes forming the Mediterranean Sea and the genesis of the terrestrial Mediterranean fauna. The Eurasian-African land-bridge formed during late Burdigalian -middle Serravallian ca. 12.5-18 Ma (Rögl 1998) initiated the species change/exchange between the Europe, Asia and Africa (Koufos et al. 2005). The territory of the Sinai Peninsula and the Levant free from the sea formed the first transit corridor. However, it was interrupted at least twice in Langhian (ca. 16-16.4 Ma) and in early Serravallian (ca. 13-13.3 Ma), while in Tortonian (ca. 11.6 Ma) the final connection of Arabian and Anatolian plates and isolation of the Mediterranean Sea took place (Rögl 1998(Rögl , 1999. Because the Central and Southern Levant as well as the Sinai Peninsula were the part of the Arabian plate connected with the African continent (Rögl 1998;Popov et al. 2004;Robertson et al. 2012;Berra and Angiolini 2014) the African species G. euphratica, H. nilotica, M. melancholica and L. flexuosa could have colonized these territories before the other species. The sharp decrease of the level of the Mediterranean Sea in Messinian (ca. 5.5-6 Ma) caused the formation of both numerous shallow enclosed saline basins and the land-bridges between Southern Europe and Northern Africa (Rögl and Steininger 1983). In our opinion during this time the active divergence and dispersion of such halophilic genera as Cephalota, Calomera and Hypaetha as well as the species of the subgenus Eugrapha occurred. All of them are arisen in the saline landscapes along the seashores of Para-and Neotethys in the Southern Russland as well as Central and Western Asia. From these regions the ancestors of the recent taxa probably dispersed through the Middle East, Arabian Peninsula and Anatolia to the Levant and the Sinai Peninsula, and some of them to Northern Africa. The second stream of the migration was possible along the Mediterranean coast of Southern Europe. Following this some species reached the Iberian Peninsula, and then the western regions of Northern Africa. In contrast G. euphratica, M. melancholica, L. flexuosa could be populated Southern Europe (Garcia-Reina et al. 2014), Western and Central Asia as well as Sind and some regions of South-Eastern Asia. Finally, possible during the last Glacial Period, the ancestors of C. javeti and C. herbacea dispersed into the Levant from the Anatolia, a region characterized by a higher level of diversity of the species of the Cicindela campestris group (Cassola 1999;Franzen 2007;Deuve 2011Deuve , 2012our unpublished data).
This proposed version of the biogeographical genesis of the fauna of tiger beetles of the Levant should be considered an initial hypothesis. Molecular analysis and more detailed paleontologic information are necessary to robustly reject or validate it.