In salty habitats, on the Mediterranean coast in marshlands (often with Anthrocnemum). Around the Dead Sea and in the Arava Valley in salty wetlands and in date palm plantations (own observations). Nocturnal. Attracted by light.

Teneral individuals in early spring (Cyprus: February), adults are active until approximately November (own observations). The number of eggs laid per females in a laboratory experiment ranges from 3 to 25 (Aydın 2011a), which is relatively low for an insect.

From southern Spain, Morocco and Sardinia to Central Asia (Cassola 1981; Cassola et al. 2014; Franzen 2001b; Franzen and Gigli 2003; Putchkov and Matalin 2003, 2017).

Mediterranean Sea coast of the Sinai Peninsula (eastwards to El-Arish) and close to Haifa (Atlit); Red Sea coast of the Sinai Peninsula (incl. near Eilat); in the Dead Sea area (especially in the swamps south of the Dead Sea) and in the Arava Valley (Franzen 2001b; Nussbaum 1987). Putchkov and Matalin (2003, 2017) list the species for Jordan. Nasir and Katbeh-Bader (2017) cited Putchkov and Matalin (2003), but do not know a record from Jordan. We do not know of any verifiable record from this period. Our record for Jordan: “29.03.2016, Pot Ash City environs, Dead Sea (Tamarisk bushes), saltmarshes, close to the edge of the sink holes in mud clefts” (CGD).

Grammognatha Motschulsky, 1850 is frequently ranked as a subgenus of Megacephala Latreille, 1802, but see Gillett (2009). Darker colored morphs occur frequently in the Dead Sea region and resemble the eastern subspecies armenica (Laporte de Castelnau, 1834) which occurs westwards to Iran. The dark form also occurs on the Mediterranean Sea coast, though it is rare.

Endangered in Israel. The species is sensitive to disturbances (drainage of habitats, cattle grazing, etc.) (Aydın 2011b). The populations found along the Mediterranean coast of Israel are in decline, and there is only one known new record in the last two decades (Atlit, late May 2012, record in collection Aligi Bandinelli). Numerous habitats have been destroyed in the Dead Sea region, but the species can sometimes be found in date palm plantations.

Open habitats with dwarf shrubs and bare ground, mostly on loamy soils (own observations and Chikatunov pers. comm.), also in quarries. Israeli records from about 1000 m a.s.l. upwards, in Lebanon up to about 2200 m a.s.l. Diurnal.

Adults found mainly in May (April to June, own observation, Matalin and Chikatunov 2016). No verifiable records for the long activity period reported by Nussbaum (1987).

Southern Turkey, Lebanon, southwestern Syria, and northern Israel (Deuve 2011).

Some records have been published by Deuve (2011). In Israel the species is only known from two areas: Mount Hermon (in the surrounding of Majdal-Shams) and from Mount Meron (Nussbaum 1987).

The C. campestris Linné, 1758 group in Asia Minor and the Middle East has been the object of recent studies (e.g. Deuve 2011; Franzen 2007; Matalin and Chikatunov 2016), but the taxonomic status of some populations has not yet been completely resolved. We agree with Azadbakhsh and Nozari (2015) that this entire tiger beetle group from South-west Asia needs to undergo revision. We believe that morphometric and molecular studies are necessary to solve the actual taxonomic and systematic problems of this group. Moreover, large amounts of material are needed for studies as the morphometric variability within populations is large (Franzen 2007).

Following Deuve (2011: 136) the specimens from Israel belong to C. javetii azari Deuve, 2011. Two further subspecies occur from Turkey to Lebanon and Syria: C. j. thughurica Franzen, 2007 and the nominate subspecies. The taxon thughurica (Fig. 10) described from southern Turkey, has been also recorded from south-western Syria (Bludan, north-west of Damascus) (Avgin and Wiesner 2009). The given site is close to the Israeli border. The existence of two subspecies of this flight-active species in this small geographic area seems unlikely. Moreover, the elytral pattern, one of the main characters to distinguish the subspecies, is not constant but varies even within a given population clearly (Figs 8, 9).

The very similar species C. herbacea occurs from Lebanon and Syria to Iran including several populations and described subspecies (Deuve 2011; 2012). The separation of these two species may be possible using male genitalia. Deuve emphasized the size and external shape of median lobe of aedeagus (cf. Figs 24–30 in Deuve 2011). Matalin and Chikatunov (2016) also emphasized the length of the median lobe as a distinguishing character. However, we have specimens from northern Lebanon (Les Cedres, Bcharre) in which the median lobe is shorter than indicated by the latter authors. Also the external shape of the median lobe of both species, C. herbacea and C. javetii, varies greatly, even within a population (e.g. Bludan, Antilebanon, 1700–2300m, Fig. 39), and not only between the populations of javetii (cf. Deuve 2011). However, the median lobe of javetii is in general more bent than that of herbacea. Although the ventro-lateral bladders of the median lobes (Matalin and Chikatunov 2016) differ between the two species, they are not a useful character for identification as the procedure of evagination of the internal sac is not feasible for many entomologists. However, clear differences in the ratio internal sac to aedeagus length can usually be seen in embedded median lobes of aedeagus (Fig. 39).

Figure 39. 

Median lobes of the aedeagus of Cicindela species: a, b C. javetii and c C. herbacea.

The body lengths of C. javetii and C. herbacea are not a good diagnostic character as the specimens in our collections show a stronger overlap than expected based in Deuve (2011). However, the proportion of the pronotum as described by Matalin and Chikatunov (2016) seems to be a good character for the identification of the two species.

As C. javetii has recently been recognized as a species, the specimens from Israel are listed under the species names C. campestris or C. herbacea (e.g. Nussbaum 1987; Valdenberg 1983). The specimens from South-west Syria are published under the taxon names herbacea and thughurica (e.g. Avgin and Wiesner 2009). Ptashkovsky (2013) included a photograph of C. herbacea, but it is unlikely that this specimen was collected in Israel.

The correct name of the taxon is javetii (Chaudoir 1861: 1), not javeti (e.g. Deuve 2011; Matalin and Chikatunov 2016).

The species is most likely extinct in Israel, as there have been no new records in the last two decades despite intensive searches on the sites from which the species was previously known. In most cases, the relevant habitats have been destroyed. Populations still exist on the Syrian side of the Hermon, as specimens have been collected there as recently as 2007 (<Syria Occ. Bludan / 40 km west of Damascus / 1700–2300m Antilebano(n) / leg. A. Wrzecionko / 5.5.2007> and same locality, but < … 2200m / Skoupý leg.>; CAL, CGS).

Unknown. Diurnal.

End of March to July (northern Lebanon; CAL, CGS).

From southern Asia Minor to Lebanon, Syria, and Iran (Deuve 2011; 2012). Range overlap with C. javetii (Deuve 2011).

No records, but occurrence possible.

see C. javetii.

Unknown. Most of the records are from mountain areas (Korell 1988).

May (CAL).

From Turkey to Iran (Wiesner 1992).

No records.

Two subspecies are known. The nominate form occurs in Syria (Wiesner 1992).

Sandy beaches (Austin et al. 2008). Larvae inhabit the beach from the high water line to the beginning of dunes (Arndt et al. 2005). Diurnal.

Main activity period of adults from June to August (Arndt et al. 2005).

Along the coasts of the Mediterranean Sea from Aegean Islands to Syria (including Crete, Cyprus, and southern Turkey) (Austin et al. 2008; Franzen 1999).

No records.

The populations from Cyprus, eastern Turkey and Syria belong to the subspecies rouxi (Barthélemy, 1835) (Franzen 1999).

This species is sensitive to disturbances caused by touristic activities on beaches. Even relatively extensive tourism can reduce the activity of adult beetles, and can prevent the development of larvae (Arndt et al. 2005).

On river banks and next to freshwater ponds with sparse vegetation on sandy, sometimes cohesive soil (Arndt 2011; Avgin 2006; Werner 1992). In desert habitats the species can be widespread (cf. Wiesner 1996). Diurnal.

In Turkey, adults from the end of May to the beginning of September (Avgin 2006).

from southeastern Europe to central Asia and India, southwards to Turkey and Syria (Acciavatti and Pearson 1989; Werner 1991; Wiesner 1992). Austin et al. (2008: 22) questioned the occurrence on Cyprus. However, Horn and Roeschkle (1891) list the species for Cyprus, and old records exist in SDEI: <Cyprus, Baudi> (3 specimens in the collections of Kraatz and Rottenberg, Lutz Behne, pers. com.) Therefore, the species is listed correctly by Putchkov and Matalin (2003; 2017: 219) for Cyprus.

The distribution range of the species in the Middle East seems to be incorrectly reported. Despite the fact that numerous authors mention the species from Israel (Avgin 2006; Putchkov and Matalin 2003), we do not know of any verified record from the country. There are no specimens in SMNHTAU, and the species is mentioned neither by Nussbaum (1987) nor by Valdenberg (1983). No verifiable records are known from Jordan (Putchkov pers. comm.). However, the species is still listed for Jordan in the latest version of the Palaearctic Catalogue of Coleoptera (Putchkov and Matalin 2017).

In Israel and Jordan, the species’ typical habitats, such as river banks in dynamic floodplains or wet pioneer vegetation with patches of bare ground, have mostly been destroyed or are strongly influenced by human activity. Therefore, a recent occurrence of C. fischeri in the Mediterranean part of the southern Levant is unlikely. There is a small chance that the species can be found in wadis or close to water reservoirs in the desert regions (cf. Wiesner 1996).

The nominate subspecies occurs in the northern Levant, while the subspecies elongatosignata (W. Horn, 1922) is found on the Arabian Peninsula (Wiesner 1992).

In riverbeds with gravel banks and stones, or on sandy ground close to water (Werner 2000).

Unknown.

Northeast Africa and the Arabian Peninsula (Werner 2000). Horn (1931) already questioned the validity of the record from Port Said. Not listed by Alfieri (1976) for Egypt.

No record. The nearest known population is found in Wadi Sharis (Abdel-Dayem et al. 2003).

Mainly in salty habitats, such as sea shores and marshlands with salt crusts, or rocky habitats (Abdel-Dayem 2004; Horn 1931; Werner 1991; 2000). Diurnal.

On the Sinai Peninsula from February until October (Abdel-Dayem et al. 2003), in the Dead Sea region from May to December (Matalin and Chikatunov 2016: 120; Nussbaum 1987).

From Senegal through northern Africa and Greece to the Middle East and Pakistan (Acciavatti and Pearson 1989; Arndt 2011; Horn 1931; Werner 2000).

In northern and southern Sinai along the coasts of the Mediterranean and of the Red Sea, and along the Suez Canal. In Israel in the Dead Sea region (Abdel-Dayem et al. 2003; Matalin and Chikatunov 2016; Nussbaum 1987). Rittner (pers. comm.) found a population in the vicinity of Akko on a rocky beach (documented by photographs, see the homepage Israel-nature-site 2017). The only known records from Jordan date back to the 1940s (Matalin and Chikatunov 2016; 4 specimens in SMNHTAU). Now also recent records from Jordan: “JOR-at-Tafila, Hammam Afra, Hot Springs, 08.05.2010” (CGD), “Dead Sea, Wadi ‘Atun, N Wadi Mujib, same date” (CSH).

The coloration can be useful for distinguishing some Calomera species, especially C. aulica, C. diania, C. littoralis and C. aulicoides (Arndt 2011). Nevertheless, C. aulica is extremely variable in color, and the coloration of the elytra ranges from black to bronze or copper with a aditional colors also occurring.

The pale elytral pattern of C. aulica is similar to that of C. aulicoides. Although the tip of the copulatory pieces of median lobe of aedeagus is similar in both species, they can be easily distinguished from each other by the external shape of the aedeagus (Figs 40, 41). A reliable character for differentiating C. aulicoides from related species is the number of teeth on the inner side of the left mandible: C. aulicoides has 4 teeth, while C. aulica and C. littoralis have only 3 (Matalin and Chikatunov 2016) (Figs 51, 52).

Figure 40. 

Median lobes of the aedeagus of species of the Calomera aulica group: a C. aulica b C. diania c C. aphrodisia.

Figure 41. 

Median lobes of the aedeagus of species of Calomera aulicoides (above) and C. littoralis winkleri (below).

Rare and endangered in Israel. Few records exist from recent decades.

Freshwater habitats. In contrast to C. aulica, which can occur on both coastal and inland habitats, C. diania is an exclusive inland species (Naviaux 1983).

End of February to August (Naviaux 1983, ZISP, CGD)

From Iraq to Pakistan and the southern Arabian Peninsula (Wiesner 1992).

No record.

Naviaux (1983) gave an excellent description with which to check any potential records from the Levant.

Rocky habitats in the littoral zone (Austin et al. 2008; Aydın 2011c; Franzen 2001a; Horn 1931). Larval development also occurs in this habitat (Lisa 2002).

May to August (Franzen 2001a; database Gebert; Lisa 2002).

From Sicily and Greece to Turkey and Syria (Wiesner 1992).

The first verifiable record from Israel was found by the cicindelid expert A. Putchkov (pers. comm.). He saw an old specimen from northern Israel (label information: <Izrael: Khaifa env.>), together with an old record from Syria (label information: <N Syrien, Ladyk env.); both specimens are preserved in ZISP. No recent records from Israel, but suitable habitats still exist in northern Israel (e.g. close to Akko and to Hadera).

Declining in Turkey due to touristic activities on the beaches (Aydın 2011c; Aydın et al. 2005). As the species occurs only locally and in habitats which tend to be under strong human pressure, the species should be classified at least as threatened. Data are deficient for the southern Levant.

Three subspecies are known, with the nominate form occurring in Turkey and in Syria (Wiesner 1992).

A coastal species which colonizes both sea shores with sandy or with cohesive soils as well as salty marshlands found behind the dunes, especially those covered with salt crusts during the summer and where the vegetation is dominated by Anthrocnemum species and by Tamarix tetragyna. Also found in river mouths and in freshwater habitats (Austin et al. 2008; Nussbaum 1987; Valdenberg 1983; own observations). Diurnal.

Middle of February until November (Matalin and Chikatunov 2016; Nussbaum 1987).

From Greece to Iran and Central Asia; southwards to Israel (Mandl 1981b).

Along the Mediterranean coast from the mouth of Nahal Betzet (=Nakhal Bezet) to the Gaza strip (Matalin and Chikatunov 2016; Nussbaum 1987). It probably also occurs on the northern coast of Sinai Peninsula, but Abdel-Dayem et al. (2003) did not list the subspecies or the nominate form from Egypt. Calomera littoralis winkleri is listed by Puchkow and Matalin (2017) and Nasir and Katbeh-Bader (2017) for Jordan, but not by Matalin and Chikatunov (2016). Nasir and Katbeh-Bader (2017) indicate the species from Ma’in Falls, a typical habitat for C. aulicoides, from where they mention also C. littoralis. We do not know of any verifiable record for Jordan.

The subspecies winkleri can be differentiated from the other subspecies of littoralis using the form of the copulatory piece of the median lobe of the aedeagus (Korell 1988; Mandl 1934; 1981b). Some populations from the eastern part of the distribution range have copulatory pieces which show an intermediate shape between those of nemoralis (Olivier, 1790) and winkleri. These populations are most probably transitional, and likely are hybrid populations. Nonetheless, they are described as a separate subspecies, mandli Mandl, 1934 (Korell 1988; Mandl 1981b).

See also C. aulica for further diagnostic characters.

Not threatened. Although the species lives along seashores which tend to be intensively influenced by touristic activities, the species has not declined as strongly as other littoral tiger beetles (for Greece: Gebert 2013, for Israel: own observation).

On sandy and stony banks close to freshwater (the Jordan River, Sea of Galilee), especially in wadis. Also found in salty habitats close to the Dead Sea. Diurnal.

Throughout the year (records from February to December) (Nussbaum 1987, own observations).

From Egypt and southern Turkey to Iran (Cassola 1999; Korell 1988; Mandl 1981b).

In Israel and Jordan along the Jordan Valley from the Hula Valley and the Sea of Galilee to the Dead Sea region, and in the Arava Valley. In Sinai on the Mediterranean Coast and in South Sinai (Abdel-Dayem et al. 2003; Matalin and Chikatunov 2016; Nussbaum 1987).

In the past most authors ranked this taxon as a subspecies (or even as a form with a rank below the subspecies) of littoralis (or of another taxon of this species group) (e.g. Mandl 1934). However, later Mandl (1981b) ranked aulicoides as a species, though only few authors accepted this ranking (e.g. Korell 1984; Werner 1991), and the majority rank it as a subspecies (Cassola 1999; Matalin and Chikatunov 2016; Putchkov and Matalin 2003, 2017; Wiesner 1992). Pesarini and Monzini (2010: 10) are, to our knowledge, the only authors from the last years, who ranked aulicoides as a valid species. However, the authors seem to have confused it with Calomera aphrodisia (Baudi, 1864), which occurs in Sicily (Brandmayr et al. 2005; Lisa 2002; Vigna-Taglianti 1993), but is not listed by Pesarini and Monzini (2010).

In the southern Levant, both littoraliswinkleri and aulicoides occur. They live in sympatry in the north of Israel (in the Hula Valley: see records for C. littoralis winkleri published by Matalin and Chikatunov 2016, and own records of C. aulicoides from Nahal Guvta (= Wadi al-Hashabi, in some maps indicated as Wadi Guyta; close to the Banias, CAL), while further southwest they are (at least) parapatric with a distance of about 20 km (Tamra – Eilabun) between populations. Although both taxa are flight-active, no intermediate forms are known (in contrast to the form of the copulatory pieces in the littoralis subspecies, see above). Calomera aulicoides has a clear and easily accessible character for differentiation from C. littoralis. While the latter one usually has three teeth on the inner side of the left mandible, C. aulicoides has four of them (Figs 51 and 52; Matalin and Chikatunov 2016). Sometimes specimens of C. littoralis winkleri have a small fourth tooth on the inner edge of the left mandible (Fig. 52). However, these specimens do not represent hybrid populations as the copulatory piece of the median lobe of aedeagus does not show any intermediate characters. The lack of intermediate populations and the sympatric and parapatric distribution ranges in the southern Levant give evidence for the species status of both, C. aulicoides and C. littoralis.

It is possible that both taxa occur sympatrically in Jordan as well (Ma’in Falls, Nasir and Katbeh-Bader 2017). We do not know any population of C. littoralis from Jordan.

The external shapes of the median lobes of the aedeagi of both taxa do not differ strongly from each other (Fig. 40), though the copulatory pieces are strongly differentiated (Fig. 42). The sharp spines of the copulatory piece in these taxa may even act in a similar way to the spines of the endophallus in closely related Carabus species of the subgenus Ohomopterus, which seem to be an example of lock-and-key genitalia (Sota and Kubota 1998). This is in contrast to most other animals, as in depth discussed by Eberhard (2010; 1985). Caution is necessary in postulating lock-and-key-functions for the genitalia in tiger beetles as we lack empirical evidence.

Figure 42. 

Copulatory pieces (arrows) of the median lobes of aedeagi of Calomera aulicoides (above) and C. littoralis winkleri (below).

Figure 43. 

Median lobes of the aedeagus of Cephalota species: a C. littorea b C. tibialis c C. circumdata d C. vartianorum.

Moreover, we have to be cautious to establish species ranks solely based on the external shape of the median lobe of aedeagus. In this respect, clearly differentiated taxa of ground beetles can have an excessive geneflow (e.g. Matern et al. 2011).

The change in the taxonomic rank of C. aulicoides stat. rest. indicates the need for a revision of the entire C. littoralis group. As a recent phylogeographic study from southeastern Europe reveals, molecular approaches can help to solve the taxonomic chaos in this group with its overlooked or neglected taxa (Jaskuła et al. 2016).

See also C. aulica for further diagnostic characters.

Not threatened in Israel or Jordan. Still widely distributed in the Sea of Galilee region and in the wadis around the Dead Sea, also in strongly grazed habitats.

Sandy and stony river banks and on the shore of lakes (Werner 2000). See also Habrodera nilotica.

In Africa throughout most of the year (Werner 2000).

From Senegal to Sudan and Ethiopia (Werner 2000). Probably does not occur in Egypt (see discussion in Abdel-Dayem et al. 2003).

No record.

Described from Ambukol (= Ambukohl) which belongs today to Sudan (Abdel-Dayem et al. 2003).

Freshwater habitats (Abdel-Dayem 2004). Sandy and stony banks of rivers and lakes, especially on yellow sand. In Africa it frequently occurs together with C. fimbriata (Werner 2000). Diurnal and nocturnal. Attracted by light.

In Africa and in the Middle East throughout almost the entire year (Matalin and Chikatunov 2016; Werner 2000). In Egypt records from January, June, August and November (Alfieri 1976).

Widely distributed in Africa: From Senegal to Somalia and from Egypt to South Africa (Werner 2000).

Wadi Isla (southern Sinai) (Abdel-Dayem et al. 2003; Alfieri 1976). Chikatunov et al. (2006) published records from Israel. We could not find any verifiable record from this survey in SMNHTAU. Therefore, the occurrence of this species in Israel is highly questionable.

On seashores and in marshland habitats. Diurnal and nocturnal. Attracted by light (Abdel-Dayem et al. 2003; Cassola 1972; Nussbaum 1987).

May to September (Abdel-Dayem et al. 2003; Nussbaum 1987).

From southern Spain to the Arabian Peninsula and Northeast Africa (Gebert 1991).

Only in southern Sinai (Abdel-Dayem et al. 2003; Alfieri 1976; Gebert 1991; Nussbaum 1987). No record from Israel or Jordan (Putchkov and Matalin 2017), but populations still exist not far from the border to both countries (<Bir Suweir / Sinai 30.4.2016 / A. Gera> SMNHTAU, CAL).

Only the nominate subspecies occurs in the southern Levant (Gebert 1991). Cephalota littorea and C. tibialis have long been confused (e.g. Mandl 1935). The revision of Gebert (1991) revealed the species status of both taxa, and described their variability (incl. genitalia and pale coloration pattern on elytra). The subspecies C. littorea alboreductata (Horn, 1934) occurs south of the distribution range of C. littorea s.str. (Gebert 1991).

Although the taxon goudotii (Dejean, 1829), which occurs along the coasts of the western Mediterranean, is currently ranked as a subspecies of C. littorea, it is probably a valid species. Cephalota littorea s.str. and goudotii do not occur parapatrically as their ranges are separated from each other by a gap which is partially filled by the distribution range of tibialis. Moreover, the differences in the median lobe of the aedeagus (shape, internal sac) may support the species status of both goudotii and littorea (but see C. aulicoides for discussion of genital structures as characters to delineate species).

The correct spelling of the author name is Forskål (Forskål 1775) and not Forsskål (Putchkov and Matalin 2017).

Shorelines of salt lakes and ponds (Austin et al. 2008; Jaskuła and Rewicz 2015; Lisa 2002), sandy beaches (Nussbaum 1987). Nocturnal (Abdel-Dayem et al. 2003). Attracted by light (Nussbaum 1987).

February to September (Abdel-Dayem et al. 2003).

From Tunisia to Egypt (Gebert 1991).

along the Mediterranean coast of the Sinai Peninsula (Gebert 1991). No record from Israel (Matalin and Chikatunov 2016; Nussbaum 1987).

Only the nominate subspecies in the southern Levant, the other two subspecies in northern Africa and on Cyprus (Gebert 1991). See also C. littorea.

On salty habitats which have very sparse vascular plant vegetation. Often found on salty crusts of lagoons and ponds behind the coastal dunes (Lisa 2002), but can also be found in similar habitats farther inland (Cassola 1970; Franzen 1996). Diurnal and nocturnal species. Attracted by light.

In Italy from June to October with activity maximum in June and July (Lisa 2002).

A Mediterranean species from Spain and Algeria to Turkey (Cassola 1970; Lisa 2002).

No verified population. – The occurrence in El Tor (southern Sinai) has been questioned by Horn and Roeschke (1891). Schatzmayr (1936) could not examine specimens from there or from anywhere else on the peninsula. Alfieri (1976) and Abdel-Dayem et al. (2003) and Abdel-Dayem (2004) list the species for southern Sinai, while Nussbaum (1987) and Matalin and Chikatunov (2016) do not. Horn et al. (1990) report that Alfieri’s beetle collection has been incorporated into the collection of Frey, which is now preserved in the natural history museum in Basel. However, no verifiable specimens of C. circumdata from Sinai are preserved in the Frey collection, and only few tiger beetle individuals from Alfieri’s collection are found in Basel (Sprecher-Uebersax, pers. comm.). We do not know of any verifiable record from the Sinai. As all other populations are known from areas with a typical Mediterranean climate, we believe that C. circumdata is not found in the Sinai (cf. Matalin and Chikatunov 2016).

Cephalota circumdata has several subspecies which are mainly characterized by the elytral pale patterns. However, Franzen (1996) reported strong pattern variability within some populations.

Saline habitats with sparse vegetation and salt crusts during summer. Diurnal and nocturnal. Attracted by light (Korell 1984).

Spring, records from February to June (Gebert 2016; Matalin and Chikatunov 2016; Nussbaum 1987).

Israel, Syria to Iran (Gebert 2016).

In the Dead Sea region of Israel (Gebert 2016; Matalin and Chikatunov 2016; Nussbaum 1987). We do not know any verifiable record from Jordan. This is in agreement with the distribution indications of Puchkov and Matalin (2003) and Wiesner (1992), but it is in disagreement with Puchkow and Matalin (2017). The country indications for Saudia-Arabia, Yemen and Jordan have not been verified (Matalin pers. comm. to Jörg Gebert on November 26, 2017).

While in older publications this taxon is listed as a subspecies of C. zarudniana (Tschitschérine, 1903), Gebert (2016) elevated it to full species rank. Cephalota vartianorum differs from C. zarudniana by slightly slender habitus as well as shape of the median lobe of aedeagus, and in the complete lack of white setae on the genae.

Critically endangered in Israel. Israel has a national responsibility for the worldwide conservation of the taxon.

Tiger beetles of coastal habitats are often sensitive to touristic use of beaches (Aydın et al. 2005; U.S. Fish and Wildlife Service 2009). Most of the Israeli beaches known to host this species are intensively used as recreational areas. With high probability at least some of the (local) populations have become extinct. Matalin and Chikatunov (2016) stated that the most recent records date from the late 80’s to the 90’s of the last century. Our most recent records are from 1990 in Israel (Neot HaKikkar = Neot Hakikar, 13. May 1990, leg. E. Orbach, COQ, CAL) and from 2000 in Syria (Euphrates, database Gebert). All other 34 entries in the database Gebert date back to the late 1980s and 1990s. Intensive searches, including use of light traps at night, of several sites in Israel such as the Enot Tsukim Reserve (= Enot Zuqim = Enot Zukim = Einot Zukim = En Fescha) from where populations have been previously recorded, have yielded no new records. The lowering of the water table and changes in land use in the Dead Sea region have strongly impacted many habitats, both freshwater and saltwater. Therefore, at further studies of the Dead Sea region, both on the Israeli and the Jordanian side, are needed to establish wether or not populations still exist.

In open forests and in grasslands, mainly on sandy ground in higher elevations (Avgin 2006; Korell 1988).

Spring species (Korell 1988).

Turkey and Syria (Franzen 2003; Wiesner 1992).

No records.

In the southern Levant found exclusively in sandy shore habitats along the Mediterranean Sea. Larval development takes place a few centimeters above the mean sea level, just above where most of the waves break (Valdenberg 1983), though larval holes are occasionally flooded by seawater. The nominate subspecies is found in both coastal and inland habitats (Cassola and Jaskuła 2004). Attracted by light (own observation).

Adults from the beginning of May until mid-November (Valdenberg 1983). Few adults also from March onwards (Matalin and Chikatunov 2016; Nussbaum 1987). The larvae overwinter mostly as second and third instars (and rarely also first instars). Egg laying begins immediately with the appearance of the adults in spring, and in certain years a few individuals may complete an entire life-cycle in the same summer, though most do not (Valdenberg 1983).

The nominate form is found from south-eastern Europe (Romania and Ukraine) and Asia Minor to Central Asia and China. The subspecies valdenbergi is patchily located in a small area along the Mediterranean coast from western Egypt to northern Israel (Horn 1931, Mandl 1981). The nominate form does not occur along the Mediterranean coast (Wiesner 1992).

In Egypt around Abu Qir, Maadia and Ras el Bar (Abdel-Dayem et al. 2003; Alfieri 1976; Horn 1931; Mandl 1981a; Nussbaum 1987). In Israel from Bat Yam to Akko, though the Bat Yam population is thought to be extinct since several decades (Valdenberg 1983; own observation).

Mandl (1981a) described the taxon valdenbergi from Ma’agan Michael as a subspecies. It is characterized by the excessive pale elytral pattern (Figs 31, 32), though there is some variation in the pattern between individuals. However, the pale elytral coloration differs even within the nominate form strongly. Some populations from the Caspian Sea show similar elytral pattern as valdenbergi (cf. Mandl 1981a; Werner 1992). Mandl mentioned in his description small differences in the copulatory pieces of the median lobe of aedeagus. As we believe this character is still not sufficiently studied. But the material we studied let us assume that valdenbergi differs from the nominate subspecies at least in the proportion and shape of the pronotum and the elytra (Fig. 29). We recommend conducting a detailed study, including morphometric and molecular methods, to clarify the status of both taxa. Ptashkovsky (2013) includes a photograph of “Lophyra contorta valdenbergi”, but the shown specimen belongs to the nominate subspecies of Cylindera contorta (cf. Fig. 29), and with all probability the photographed specimen was not collected in Israel.

Figure 44. 

Median lobes of the aedeagus of Cylindera species: a C. contorta s.str. b C. rectangularis.

Critically endangered in Israel. Tiger beetles of coastal habitats tend to be very sensitive to touristic use of beaches (Aydın et al. 2005; U.S. Fish and Wildlife Service 2009). Most of the Israeli beaches which are known to host this species are intensively used as recreational areas, and therefore at least many of the populations have gone extinct. As a metapopulation structure may be possible in this species, the decrease in some populations can have a tremendous effect on the survival of the entire subspecies. Matalin and Chikatunov (2016) stated that the latest records date from the late 80s to the 90’s of the last century, and our most recent records are from 2003 (Shefeh Na’aman Nature Reserve, COQ). As the entire world population of this taxon is located in Israel (the majority of the known populations) and in Egypt, these countries bear the responsibility for the worldwide preservation of this subspecies.

Banks of freshwater in wadis (Abdel-Dayem and Kippenhan 2013), especially on loamy soil (Werner 2000). In Saudi Arabia it co-occurs with Calomera aulica, C. alboguttata and Myriochila melancholica (Abdel-Dayem and Kippenhan 2013).

March, but more frequently in June (Abdel-Dayem and Kippenhan 2013).

From Central Africa to Sudan and Saudi Arabia (Abdel-Dayem and Kippenhan 2013).

No record.

Margins of both stagnant and running freshwater bodies, including artificial water reservoirs, and in salty habitats (Austin et al. 2008; Jaskuła and Rewicz 2015; Lisa 2002). Diurnal. Attracted by light (Abdel-Dayem et al. 2003).

Figure 45. 

Median lobes of the aedeagus of Lophyridia species: a L. flexuosa b L. hilariola c L. histrio.

Figure 46. 

Median lobes of the aedeagus of Myriochila species: a M. melancholica b M. orientalis.

March to December (Abdel-Dayem et al. 2003; Matalin and Chikatunov 2016; Nussbaum 1987).

From southern Europe to southern Africa and from the Cape Verde Islands to China (Wiesner 1992).

Widespread in Sinai, Israel and Jordan (Matalin and Chikatunov 2016; Nasir and Katbeh-Bader 2017; Wiesner 1992).

Not endangered, it is abundant even in habitats strongly influenced by human activity (e.g. on intensively grazed sites or on wet fallow land close to Tel Aviv and Amman).

Unknown.

Unknown.

From Turkey and Syria to China (Wiesner 1992).

No record.

Semi-desert and savanna habitats (Werner 2000).

Unknown.

Southern Sahel zone from Mauritania and Senegal to Sudan (Werner 2000). Listed by Horn and Roeschke (1891) also for Egypt. Horn (1931) knew of four specimens labelled “Egypt”, all from different collections. Therefore it seems unlikely that all records are mislabelled. Nonetheless, the records may refer to Egypt in its historical sense which includes parts of modern-day Sudan (cf. Alfieri 1976).

No record.

Sandy seashores (Abdel-Dayem et al. 2003).

Recorded in Egypt from May to August (Abdel-Dayem et al. 2003, Matalin and Chikatunov 2016). In SMNHTAU are also specimens collected in September (own observation).

Figure 47. 

Median lobe of the aedeagus of Hypaetha singularis.

Figure 48. 

Head in lateral view: Calomera alboguttata (above) and Habrodera nilotica (below).

Figure 49. 

Metafemora, lateral view on lower side: C. aulica (left), C. aulicoides (right).

Figure 50. 

Head of Calomera species in dorsal view: C. diania (left), C. aulica (right).

Figure 51. 

Head of Calomera species in dorsal view: C. aphrodisia (left), C. aulicoides (right).

Figure 52. 

Head of Calomera littoralis winkleri: with regular form of left mandible (left) and a small fourth tooth on the inner side of left mandible (arrow, right).

Figure 53. 

Tibial spurs and 1^st tarsal segment of Calomera littoralis winkleri (left) and Calomera aulicoides (right).

Figure 54. 

1^st antennal segment: with only one erect distal seta (Calomera alboguttata; above) and with one erect distal seta and additional recumbent white setae (Lophyra histrio; below).

Figure 55. 

Forebody of cicindelid species: Hypaetha singularis (above, left), Habrodera nilotica (above, right), and Calomera alboguttata (below).

Figure 56. 

Overview of the main functions of the application TIGER BEETLES ID. This stand-alone application for mobile Android devices (smartphones, tablets) can be freely downloaded at https://doi.org/10.3897/zookeys.734.21989.suppl1.

From Egypt east of the Nile to the Arabian Desert (Oman, Yemen), also found in tropical Africa (Werner 2000; Wiesner 1992).

Only in southern Sinai (Abdel-Dayem et al. 2003; Nussbaum 1987), from where the SMNHTAU records also originate (Matalin and Chikatunov 2016). Ptashkovsky (2013) recorded the species for Israel, but no verifiable records are found in SMNHTAU, which includes the collection of Ptashkovsky.

Eurytopic species found on sea shores, in saltmarshes, in dune depressions, on river banks, in oases and in palm plantations, not restricted to coastal habitats (Abdel-Dayem et al. 2003; Jaskuła and Rewicz 2015; Lisa 2002; Nussbaum 1987).

Throughout most of the year, from February to December (Matalin and Chikatunov 2016; Nussbaum 1987).

From Morocco to Israel (Wiesner 1992).

Numerous records from the Mediterranean coast in Israel and northern Sinai, southwards to the Negev and central Sinai (Abdel-Dayem et al. 2003; Horn 1931; Matalin and Chikatunov 2016; Nussbaum 1987).

Not threatened. A widespread species which also can be found in highly disturbed habitats.

On sparsely vegetated escarpments along rivers (Franzen and Bischoff 1995).

Poorly known, records from April and May, but may have a longer activity period (Franzen and Bischoff 1995).

From Turkey to Iran (Wiesner 1988).

No record.

A table found in Franzen and Bischoff (1995) can be used to differentiate L. hilariola from L. flexuosa.

On beaches, in salt marshlands and in freshwater habitats; can be found together with C. fischeri (Wiesner 1996).

February to September (Wiesner 1996).

From the Arabian Peninsula to India (Wiesner 1992).

No record.