2urn:lsid:arphahub.com:pub:45048D35-BB1D-5CE8-9668-537E44BD4C7Eurn:lsid:zoobank.org:pub:91BD42D4-90F1-4B45-9350-EEF175B1727AZooKeysZK1313-29891313-2970Pensoft Publishers10.3897/zookeys.578.73837383Research ArticleCarabidaeCatalogues and ChecklistsFaunistics & DistributionIdentification keySpecies InventoriesCenozoicEgyptIsraelJordanLebanonSyriaThe tiger beetles (Coleoptera, Carabidae, Cicindelinae) of Israel and adjacent landsMatalinAndrey V.12andrei-matalin@yandex.ruhttps://orcid.org/0000-0002-7790-8709ChikatunovVladimir I.3Education-Scientific Centre Ecology & Biodiversity, Moscow State Pedagogical University, Moscow 129164, RussiaDepartment of Biology, Russian National Research Medical University named after N.I. Pirogov, Moscow 117997, RussiaDepartment of Zoology, Tel-Aviv University, Tel Aviv 69978, Israel
Corresponding author: Andrey V. Matalin (a_matalin@tochka.ru; andrei-matalin@yandex.ru)
Academic editor: A. Casale
201608042016578115160DA5CFFFF-FFCE-6A62-FF8E-5903FF8BF778A1A7FC2B-0E1D-4BC8-8AE4-30CC9478DF7B5793540212201512022016Andrey V. Matalin, Vladimir I. ChikatunovThis 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.http://zoobank.org/A1A7FC2B-0E1D-4BC8-8AE4-30CC9478DF7B
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. Cylinderacontortavaldenbergi and Cicindelajavetiazari 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.
Matalin AV, Chikatunov VI (2016) The tiger beetles (Coleoptera, Carabidae, Cicindelinae) of Israel and adjacent lands. ZooKeys 578: 115–160. doi: 10.3897/zookeys.578.7383
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) Cicindelalittoralisaulicoides. In 1934 Mandl recorded for the Palestine two subspecies of Cicindelalittoralis: 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 Cicindelaaulica (Horn, 1931), Cicindelalittoralisaulicoides (Mandl, 1934) and Megacephalaeuphratica (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 (1983, 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.
Since the beginning of 2000 interest in the Cicindelinae of the Middle East has increased significantly (El-Moursy et al. 2001; Franzen 2001, 2007; Finkel et al. 2002; Wiesner 2002, 2005; Abdel-Dayem et al. 2003; Rittner 2003; Abdel-Dayem 2004, 2012; Chikatunov et al. 2006; Avgin and Özdikmen 2007; Franzen 2007; Avgin and Wiesner 2009; Ptashkovsky 2009; Deuve 2011, 2012; Abdel-Dayem and Kippenhan 2013; Jaskuła and Rewicz 2014). These studies revealed the presence of several species of tiger beetle previously unknown from the area. For example, Habroderanilotica (Dejean, 1825), Hypaethasingularis (Chaudoir, 1876) and Cephalotalittorea (Forskål, 1775) were recorded for the first time in Israel (Chikatunov et al. 2006). However, in the next publications these species were not included (Ptashkovsky 2009).
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)
MPU
Moscow State Pedagogical University (Moscow, Russia)
SIZ
I.I. Schmalhausen Institute of Zoology, National Academy of Sciences of Ukraine (Kiev, Ukraine)
cJW
collection of Jürgen Wiesner (Wolfsburg, Germany)
cIOv
collection of Igor’ Ovsyannikov (Moscow, Russia)
The nomenclature of elytral pattern follows Acciavatti and Pearson (1989); the nomenclature of male internal sac follows Matalin (1998); the chorology follows Vigna Taglianti et al. (1999) with some additions; the regions of Israel and the Sinai Peninsula (Egypt) follow Nussbaum (1987). The similarity of the faunas of tiger beetles was calculated using complete linkage procedure (squared Euclidean distances).
The species included here that are not yet recorded from Israel are marked in the catalogue and in the key with a symbol (○).
Results and discussionCatalogue of the tiger beetles of Israel and adjacent landsFamily Carabidae Latreille, 1802Subfamily Cicindelinae Latreille, 1802Tribe Cicindelini Latreille, 1802Subtribe Cicindelina Latreille, 1802Genus Calomera Motschulsky, 1862AnimaliaColeopteraCarabidae380CB73B-6332-50B1-AC2F-EFEC2CAE5E22Calomeraaulicaaulica(Dejean, 1831)General distribution.
Europe Greece; Asia Lebanon, Israel, Jordan, Syria, Egypt (Sinai), Saudi Arabia, Arab Emirates, Oman, Yemen, Bahrain, Iran, Iraq, Pakistan; Africa: Cape Verde Islands, Senegal, Guinea Bissau, Mauritania, Morocco, Tunisia, Algeria, Libya, Sudan, Chad, Egypt, Somalia, Eritrea, Djibouti.
Distribution of Calomeraaulicaaulica in Israel, Palestine and border areas of Jordan (open circles records before year 1949, half-solid circles records between years 1950–1999, solid circles records after year 2000; map source Eric Gaba Wikimedia Commons user: Sting and Wikimedia Commons user: NordNordWest, URL https://upload.wikimedia.org/wikipedia/commons/7/7c/Israel_relief_location_map.jpg)
Distribution of Calomeraaulicaaulica (red circles) and Calomeralittoralisaulicoides (blue rhombs) in Sinai Peninsula, Egypt (open symbols records before year 1949, half-solid symbols between years 1950–1999, solid symbols records after year 2000; URL map source https://upload.wikimedia.org/wikipedia/commons/5/59/Sinai_relief_location_map.svg).
https://binary.pensoft.net/fig/84318
Jordan, Ma’Daba: Callirhoe, 7.VI.1942, leg. H. Bytinski-Salz 2♀ (TAU).
Egypt (Sinai), Northern Sinai: Sabkhat al Bardawil, 25.VIII.1967, leg. I. Margalit 3♀♀; 24.VIII.1979, leg. A. Valdenberg 1♂ 2♀♀ (TAU); Ismailia (after Alfieri 1976); Zaranik Protectorate (after El-Moursy et al. 2001; Abdel-Dayem et al. 2003; Abdel-Dayem 2004); Sinai Mountains: 20 km NE of Dahab, saline land, 4.VIII.2008, leg. A. Sokolov 4♂♂5♀♀ (MPU); Southwestern Sinai: Suez – 1♂ (ZMUM); Nabeq, 17.VIII.1971, leg. J. Kugler 1♂ 2♀♀; 8.V.1980, leg. A. Valdenberg 4♂♂ 6♀♀; Ras al Tantur, 5.VII.1957, leg. Ch. Lewinsohn 2♂♂ 1♀, 17.VIII.1971, leg. M. Kaplan -1♀ (all TAU); 15 km W Ofira, Golf v. Elat, Straße von Tiran, 3.IV.1981, leg. G. Gerdes 1♂ (cJW); Wadi Gharandal, 20.V.1969, leg. Tsabar 1♂ (TAU); Abu Zenima, Wadi Tayebeh (both after Alfieri 1976); El Tor (after Alfieri 1976; Abdel-Dayem et al. 2003; Abdel-Dayem 2004); Ras Muhammad (after Nussbaum 1987).
Distribution of two subspecies of Calomeralittoralis in Israel, Palestine and border areas of Jordan (red circles – C.l.aulicoides, blue circles – C.l.winkleri, open circles records before year 1949, half-solid circles – records between years 1950–1999, solid circles – records after year 2000; map source Eric Gaba Wikimedia Commons user: Sting and Wikimedia Commons user: NordNordWest, URL https://upload.wikimedia.org/wikipedia/commons/7/7c/Israel_relief_location_map.jpg).
https://binary.pensoft.net/fig/84319
Jordan, Al Balqā’: Al Maghtas, 12.II.1942, leg. H. Bytinski-Salz 1♂ 2♀♀(TAU); Ma’Dabā: Suwayma, Dead Sea, 5.IV.2000, leg. G. & I. Zappi 1♂1♀ (MPU).
Egypt (Sinai), Northern Sinai: Ismailia (after Alfieri 1976); Southwestern Sinai: El Tor (after Alfieri 1976; Abdel-Dayem et al. 2003; Abdel-Dayem 2004).
(Fig. 4). Israel (including State of Palestine), Dead Sea Area: Yeriho, 24.IV.1927, leg. O. Theodor 1♂; ‘Enot Zuqim, 1.II.1994, and 13.III.1994, leg. V. Chikatunov 1♂ 2♀♀; Qalya, 6.V.1980, leg. A. Valdenberg 2♂♂ 3♀♀; Ne`ot HaKikkar, 7.V.1980, leg. A. Valdenberg 4♂♂; 28.IV.1984, leg. E. Shney-Dor 2♀♀; 19.IV.1999, leg. I. Yarom & V. Kravchenko 1♂1♀ (all TAU), 7.V.1980, leg. R. Naviaux 1♀; V.1990, leg. Orbach 1♂1♀ (both cJW).
Distribution of Cephalotazarudnianavartianorum (orange triangles), Cicindelajavetiazari (red circles) and Cylinderacontortavaldenbergi (blue rhombs) in Israel, Palestine and border areas of Lebanon (open symbols records before year 1949, half-solid symbols – records between years 1950–1999; map source Eric Gaba Wikimedia Commons user: Sting and Wikimedia Commons user: NordNordWest, URL https://upload.wikimedia.org/wikipedia/commons/7/7c/Israel_relief_location_map.jpg).
https://binary.pensoft.net/fig/84320Comments.
References to Cephalotazarudnianavartianorum (Mandl, 1967) as Cephalotadeserticola (Faldermann, 1836) (Chikatunov et al. 2006; Ptashkovsky 2009) were based on two mis-identified males from Qalya by K. Mandl. According to Franzen and Wiesner (1998)C.deserticola is distributed in the western part of Central Asia, as well as in Iran, Azerbaijan, Armenia and north-eastern Turkey.
Distribution of Cephalotatibialistibialis (red circles), Cephalotalittorealittorea (blue rhombs) and Habroderaniloticanilotica (green triangles) in Sinai Peninsula, Egypt (open symbols records before year 1949, half-solid symbols – records between years 1950–1999; URL map source https://upload.wikimedia.org/wikipedia/commons/5/59/Sinai_relief_location_map.svg).
https://binary.pensoft.net/fig/84321Comments.
The specimen of Cephalotalittorealittorea (Forskål, 1775) with label “Jerusalem” from Zoologisches Museum der Humboldt-Universität (Berlin) is mislabelled (see Gebert 1991). All subsequent records of this species from Israel (Wiesner 1992; Puchkov and Matalin 2003) are in error.
In some publications (Abdel-Dayem et al. 2003; Abdel-Dayem 2004) Cephalotacircumdata (Dejean, 1822) was recorded from the Sinai Peninsula (El Tor). However, the nominotypical subspecies of C.circumdata occurs along the Aegean, Marmora, Black and Mediterranean Sea costs in the Greece, Bulgaria, western Turkey, and, probably Rumania (Franzen 1996; Cassola 1999; Gebert 1999); C.c.cappadocica Franzen, 1996 and C.c.hattusae Franzen, 1996 live along banks of the salt lakes in the central Turkey (Franzen 1996; Cassola 1999; Gebert 1999); C.c.leonschaeferi Cassola, 1970 occupies the Mediterranean sea cost in southern France (including Corsica) and north-western Italia (Gebert 1999); while C.c.imperialis Klug, 1834 records in the Italia (Sardinia and Sicilia), south-eastern Spain (including Balearic Islands), Tunisia and Algeria, but not in the Libya and Egypt (Gebert 1999). Most likely, the aberrant specimen of C.littorea was incorrectly identified as C.circumdata.
Lebanon, Liban-Nord: Bcharré, Les Cèdres, VI. 1997 1♂1♀ (cIOv); Bcharré 1♀ (after Franzen 2007); Tannourine 1♀ (after Deuve 2011).
Comments.
Until recently both these species were recorded from Syria, Lebanon and Israel by several authors as Cicindelaherbacea Klug (Valdenberg 1983; Nussbaum 1987; Wiesner 1992; Puchkov and Matalin 2003; Chikatunov et al. 2006; Franzen 2007; Ptashkovsky 2009). However, according to recent data C.herbacea does not occur in Israel (Deuve 2011, 2012). The nominative subspecies occurs in Lebanon and Syria; C.h.aleppensis Deuve, 2012 is recorded from north-western Syria, while C.h.perreaui Deuve, 1987 and C.h.colasi Deuve, 2011 are found in Turkey Tunceli and Adana Provinces, respectively. On the basis of the shape of pronotum (Figs 38vs 39), white elytral pattern (Figs 54vs 55), size of aedeagus and shape of it apex (Figs 93vs 97), as well as shape of internal sack (Figs 94–96vs 98–100) we consider all studied specimens from Israel to be Cicindelajavetiazari Deuve, 2011 (type locality – Lebanon, Jezzine). It should be noted that the taxonomy of intraspecific forms within the ‘campestris’-group is complex, and additional studies are necessary.
Genus Cylindera Westwood, 1831AnimaliaColeopteraCarabidae4877553B-844C-5EE1-A8AE-F74437ED1F60Cylindera
(Eugrapha)
contortavaldenbergi(Mandl, 1981)General distribution.
Distribution of Lophyraflexuosaflexuosa in Israel and Palestine (open circles records before year 1949, half-solid circles records between years 1950–1999, solid circles records after year 2000; map source Eric Gaba Wikimedia Commons user: Sting and Wikimedia Commons user: NordNordWest, URL https://upload.wikimedia.org/wikipedia/commons/7/7c/Israel_relief_location_map.jpg).
Distribution of Myriochilamelancholicamelancholica in Israel, Palestine and border areas of Jordan (open circles records before year 1949, half-solid circles records between years 1950–1999, solid circles records after year 2000; map source Eric Gaba Wikimedia Commons user: Sting and Wikimedia Commons user: NordNordWest, URL https://upload.wikimedia.org/wikipedia/commons/7/7c/Israel_relief_location_map.jpg).
https://binary.pensoft.net/fig/84323
Jordan, Al Balqā’: Al Maghtas, 23.V.1942, H. Bytinski-Salz 1♀ (TAU).
Egypt (Sinai), Northern Sinai: El Arish, 15.VI.1968, leg. J. Kugler 1♂ 2♀♀ (TAU; including after Abdel-Dayem et al. 2003; Abdel-Dayem 2004); Southwestern Sinai: Ofira, sewage, 2.V.1981, leg. A. Freidberg 1♂ (TAU).
(Figs 8, 9). Israel (including State of Palestine), Northern Coastal Plain: Haifa (after Franzen 2001); ‘Atlit, 4.VI.1979, 1.V.1979, leg. A. Valdenberg 2♀♀; 4.VI.1983, leg. E. Shney-Dor 1♂; 32°42'N, 34°56'E, 17.V.1997, leg. E. Orbach 1♂ 1♀ (all TAU), VI.1989, leg. E. Orbach 1♂ (cJW); V.1989, not far from the coastal line, running to light, leg. E. Orbach 2♂♂ (after Werner 1999);Dead Sea Area: Bet Ha’Arava, 5.IV.1941, leg. O. Theodor 2♂♂ 2♀♀; Jordan River, near Dead Sea, 5.IV.1941, leg. O. Teodor 1♀ (TAU); ‘En Gedi, 24.III.1958, leg. J. Kugler 2♂♂; 15.III.65, leg. K. Yefenof 1♀; Ne`ot HaKikkar, 15.II.1999, 19.IV.1999, leg. I. Yarom & V. Kravchenko 3♂♂ 1♀; Qalya, 11.IV.1958, leg. M. Pener 2.♂♂ 1♀; Sedom, 6.V.1961, at night, leg. J. Wahrman 1♂; Shefekh Zohar, 16.IV.1980, leg. J. Kugler 1♀; Zomet Zohar, 17.IV.1997, leg. L. Friedman 1♀ (all TAU); Nawit Pools (after Nussbaum 1987); Arava Valley: Gerofit, 23.IV.2003, 12.V.2003, 6.VI.2003, leg. D. Utshitel & V. Chikatunov 3♂♂ 2♀♀ (TAU); Southern Negev: Elat, 30.VIII.1959, leg. L. Fishelsohm – 1 ♂ (TAU).
Distribution of Grammognathaeuphraticaeuphratica in Israel and Palestine (open circles records before year 1949, half-solid circles records between years 1950–1999, solid circles records after year 2000; map source Eric Gaba Wikimedia Commons user: Sting and Wikimedia Commons user: NordNordWest, URL https://upload.wikimedia.org/wikipedia/commons/7/7c/Israel_relief_location_map.jpg).
Distribution of Hypaethasingularis (red circles), Myriochilamelancholicamelancholica (blue rhombs), Lophyraflexuosaflexuosa (orange squares) and Grammognathaeuphraticaeuphratica (lilac triangles) in Sinai Peninsula, Egypt (open symbols records before year 1949, half-solid symbols – records between years 1950–1999; URL map source https://upload.wikimedia.org/wikipedia/commons/5/59/Sinai_relief_location_map.svg).
https://binary.pensoft.net/fig/84325
Egypt (Sinai), Northern Sinai: Sabkhat al Bardawil, 23.III.1969, leg. A. Nitzan 1♂♂ 2♀♀ (TAU); Arish (after Abdel-Dayem 2004); Zaranik Protectorate (after El-Moursy et al. 2001; Abdel-Dayem 2004); Sinai Mountains: Dahab (after Nussbaum 1987); Southwestern Sinai: El Tor (after Schatzmayr 1936; Alfieri 1976; Nussbaum 1987; Abdel-Dayem et al. 2003; Abdel-Dayem 2004); Nabeq (after Nussbaum 1987).
An identification key to the tiger beetles of Israel and adjacent lands
1(2)
Anterior angles of pronotum projected towards the margin of prothorax (Figs 10, 45); the fourth joint of maxillary palpus shorter than the third one
Anterior angles of pronotum not projected towards the margin of prothorax (Figs 12–14); the fourth joint of maxillary palpus longer than the third one
Cicindelini 3
3(4)
Proepisterna prominent on pronotum so pronotopleural suture clearly visible dorsally (Fig. 42); anterior margin of pronotum with row of flat white setae
Proepisterna not prominent on pronotum so pronotopleural suture not visible dorsally (Figs 30–41, 43–44); anterior margin of pronotum glabrous
5
5(6)
Labrum with four submarginal setae (Fig. 29); middle and hind femora with numerous hooked setae along posterior margin, hind femora with sparse hooked setae (Fig. 64)
Myriochila (s. str.) (Myriochila(s. str.)melancholica melancholica (Fabricius, 1798))
6(5)
Labrum with at least six submarginal setae, except aberrant specimens with 3–5 setae (Figs 15–26, 28); femora without hooked setae along posterior margin
7
7(8)
Genae pilose (Figs 11, 14)
9
8(7)
Genae glabrous (Figs 12–13)
13
9(10)
Clypeus glabrous, anterior and posterior margins of each eye with group of white decumbent setae; labrum with 10 submarginal setae in a single row (Fig. 25); fourth antennomere of males with penicillus (Fig. 14); white elytral pattern with complete humeral lunule, long sinuate middle band and apical lunule coupling together via marginal and sutural bands (Fig. 56)
Clypeus pilose, anterior and posterior margins of each eye glabrous; labrum with several rows of numerous submarginal setae (Figs 15–18); fourth antennomere of males glabrous (Fig. 11); white elytral pattern without marginal and sutural bands (Figs 46–49)
Calomera 11
11(12)
Elytra dark brown with purple-bronze or green reflection (Fig. 46); pronotum 1.05–1.15 times as wide as long with straight parallel or slightly convergent lateral sides (Fig. 30); aedeagus straight, with long thin basal portion, apical lobe with distinct lateral flanges and small hook, without central groove (Figs 73, 77), ventro-apical bladder of internal sac short, right and left basi-lateral bladders very large (Figs 77, 81)
Calomeraaulicaaulica (Dejean, 1831)
12(11)
Elytra green sometimes with bronze or blue reflection (Figs 47–49); pronotum 1.15–1.35 times as wide as long with rounded distinctly convergent lateral sides (Figs 31–33); aedeagus curved, with short basal portion, apical lobe without lateral flanges and hook, but with clear central groove (Figs 74–76, 82–84), ventro-apical bladder of internal long, right and left basi-lateral bladders as small acicular areas (Figs 78–80, 82–84)
Calomeralittoralis (Fabricius, 1787) 12a
12a(12b)
Left mandible with four teeth distal to apical molar (Fig. 16); pronotum narrow, 1.15–1.2 times wider than long (Fig. 31); aedeagus with small distinct bulge on the dorsal surface (Fig. 74); ventro-apical bladder of internal sac long and curved towards and on the left, apex of medial tooth blunt (Figs 74, 78, 82)
Left mandible with three teeth distal to apical molar (Fig. 17–18); pronotum wide, 1.2–1.35 times wider than long (Figs 32–33); aedeagus without bulge on the dorsal surface (Figs 75–76); ventro-apical bladder of internal sac straight and not curved, apex of medial tooth sharp (Figs 75–76, 79–80, 83–84)
12c
12c(12d)
Labrum wider, 2.6–2.65 times as wide as long (Fig. 32), base of medial tooth of internal sac with one rarely two small additional spikes (Figs 75, 79, 83)
Calomeralittoraliswinkleri (Mandl, 1934)
12d(12c)
Labrum narrower, 2.35–2.45 times as wide as long (Fig. 33), base of medial tooth of internal sac smooth, without additional spikes (Figs 76, 80, 84)
(○) Calomeralittoralisnemoralis (Olivier, 1790)
13(14)
Labrum tridentate with distinctly prominent apical teeth; mandibles with two teeth distal to apical molar (Fig. 26); scapus covered by numerous white decumbent setae (Figs 13, 26), fourth antennomere of males with penicillus (Fig. 13); posterior margin of each eye with group of white decumbent setae; white elytral pattern with basal dot and incomplete sutural band (Fig. 57)
Lophyra (s. str.) (Lophyra(s. str.)flexuosa flexuosa (Fabricius, 1787))
14(13)
Labrum unidentate (Fig. 20–24, 28), in some species tridentate but with not or slightly prominent apical teeth only (Fig. 19); mandibles with three teeth distal to apical molar (Fig. 19–24); scapus glabrous (Figs 19–22, 28) or only with several sparse setae except apical ones (Figs 23–24), fourth antennomere of males glabrous (Fig. 11); posterior margin of each eye glabrous; white elytral pattern without basal dots and sutural band (Figs 50–55, 59)
15
15(16)
Head glabrous; scapus with apical setae only (Fig. 19–22, 28); lateral side of pronotum pilose (Figs 34–37, 43); white elytral pattern with long marginal band and long sinuate middle band (Figs 50–53, 59)
17
16(15)
Frons and vertex with long soft hairs, scapus with several setae except apical ones (Figs 23–24); lateral side of pronotum with soft sparse setae (Figs 38–39); white elytral pattern without marginal band and only with short slightly curved middle band (Figs 54–55)
Cicindela (s. str.) 25
17(18)
Anterior portion of apical lunule long, extending basal transverse portion of middle band (Fig. 59)
Anterior portion of apical lunule short, extending only apical portion of middle band (Figs 50–53)
Cephalota (Taenidia) 19
19(20)
Labrum tridentate, relatively short, no less than 2.3 times as wide as long (Fig. 19); pronotum 1.2–1.4 times wider than long (Fig. 34); mesepisternum entirely covered by white setae, densely in males and sparsely in females; white elytral pattern with relatively broad marginal band coupling with humeral and apical lunule as well as with middle band (Fig. 50), apical margin of elytra in sexes wide rounded, subtend practically right angle with sutural tooth (Figs 65–66); aedeagus with long thin gradually curved basal portion (Fig. 85)
Labrum unidentate, relatively long, no more than 2.3 times as wide as long (Figs 20–22); pronotum 1.1–1.25 times wider than long (Figs 35–37); mesepisternum covered by white setae only along posterior margin and on the base; white elytral pattern usually with narrow marginal band or without it so in some specimens humeral lunule distinctly separated (Figs 51–53), apical margin of elytra at least in males subtend acute angle with sutural tooth (Figs 67–72); aedeagus with short thin basal portion (Figs 87, 89, 91)
21
21(22)
4–11th antennomeres dark brown; elytra bright purple, 1.5–1.6 times as long as wide (Fig. 51), apical elytral margin in females narrowly rounded and subtend small right angle with sutural tooth (Fig. 67–68); aedeagus with broad blunt apex (Figs 87–88)
4–11th antennomeres light brown or yellowish; elytra greenish or greenish-blue sometimes with distinct golden-purple reflection, no less than 1.65 times as long as wide (Figs 52–53), apical elytral margin in both sexes subtend acute angle with sutural tooth (Figs 69–72); aedeagus with arrow-shaped apex (Figs 89–92)
23
23(24)
Labrum shorter, 2.0–2.2 times as wide as long (Fig. 21); lateral side of pronotum straight, slightly convergent to large posterior angles (Fig. 36); humeral lunule separated or narrowly coupled with marginal band (Fig. 52); aedeagus larger, with relatively long thin basal portion (Fig. 89) and short tapered apex (Fig. 90)
Labrum longer, 1.6–1.7 times as wide as long (Fig. 22); lateral side of pronotum slightly rounded, distinctly convergent to small posterior angles (Fig. 37); humeral lunule coupled with middle band via marginal band (Fig. 52); aedeagus smaller, with short thin basal portion (Fig. 91) and long tapered apex (Fig. 92)
Pronotum with convex lateral sides gradually convergent to posterior angles, anterior margin same length or slightly longer than posterior one, notopleural suture looks like smooth border (Fig. 39); mesepisternum of female with small shallow pit and deep all along coupling sulcus, mesepimeron with groove along anterior margin (Fig. 63); middle band of white elytral pattern without oblique strip between transverse basal and circled apical portions, basal portion of apical lunule small (Fig. 55); aedeagus shorter, no more than 0.55 times as long as elytra (Fig. 97); basal and right ventro-lateral bladders of internal sac short (Figs 98–100)
Cicindela(s. str.)javeti azari Deuve, 2011
26(25)
Pronotum with straight lateral sides sharply convergent to posterior angles, anterior margin clearly longer than posterior one, notopleural suture looks like cut border (Fig. 38); mesepisternum of female with deep apically but shallow and indistinct basally coupling sulcus only, mesepimeron without groove along anterior margin (Fig. 62); middle band of white elytral pattern with distinct oblique strip between transverse basal and circled apical portions, basal portion of apical lunule large (Fig. 54); aedeagus longer, no less than 0.6 times as long as elytra (Fig. 93); basal and right ventro-lateral bladders of internal sac long (Figs 94–96)
Head and pronotum of males, left lateral view: 10Grammognathaeuphraticaeuphratica11Calomeraaulicaaulica12Cicindelajavetiazari13Lophyraflexuosaflexuosa14Habroderanilotica; aa pr – anterior angle of pronotum; pc – penicillus. Scale bars: 1 mm.
Labrum and mandibles of males, dorsal view: 15Calomeraaulicaaulica16Calomeralittoralisaulicoides17Calomeralittoraliswinkleri18Calomeralittoralisnemoralis19Cephalotatibialistibialis20Cephalotalittorealittorea21Cephalotazarudnianavartianorum22Cephalotadeserticoladeserticola. Scale bars: 1 mm.
Labrum and mandibles of males, dorsal view: 23Cicindelaherbaceaherbacea24Cicindelajavetiazari25Habroderaniloticanilotica26Lophyraflexuosaflexuosa27Hypaethasingularis28Cylinderacontortavaldenbergi29Myriochilamelancholicamelancholica. Scale bars: 1 mm.
Left elytron of males, dorsal view: 46Calomeraaulicaaulica47Calomeralittoralisaulicoides48Calomeralittoraliswinkleri; 49Calomeralittoralisnemoralis50Cephalotatibialistibialis51Cephalotalittorealittorea52Cephalotazarudnianavartianorum53Cephalotadeserticoladeserticola54Cicindelaherbaceaherbacea55Cicindelajavetiazari56Habroderaniloticanilotica57Lophyraflexuosaflexuosa58Hypaethasingularis59Cylinderacontortavaldenbergi60Myriochilamelancholicamelancholica61Grammognathaeuphraticaeuphratica. Scale bars: 1 mm.
Aedeagus of Calomera spp., right lateral view: 73C.aulicaaulica74C.littoralisaulicoides75C.littoraliswinkleri76C.littoralisnemoralis; mt – median tooth. Scale bar: 1 mm.
Internal sack of Calomera spp.: 77, 81C.aulicaaulica78, 82C.littoralisaulicoides79, 83C.littoraliswinkleri80, 84C.littoralisnemoralis77–80 right lateral view 81–84 dorsal view; BLR basi-lateral right bladder; BLL basi-lateral left bladder; VA – ventro-apical bladder; mt – median tooth. Scale bar: 1 mm.
Similarities between tiger beetle faunas of different regions of Israel and the Sinai Peninsula (Complete linkage procedure, squared Euclidean distances).
https://binary.pensoft.net/fig/84337Distribution
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 (Table 1). The Rift Valley, including Jordan Valley, Dead Sea area and Arava Valley, with six cicindelids species is the most speciose region. The Coastal Plain is the second richest region with five species. The species richness gradually decreases from Northern (fife species) through Central (four species) to Southern (three species) Coastal Plain. In the central densely populated areas of Israel, such as Samaria and Judea, the least number of tiger beetles species are recorded. Among all M.melancholicamelancholica is the most common species observed in all regions of the country (Table 1, Fig. 7), while L.flexuosaflexuosa 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). Lophyraflexuosa (Fabricius, 1787) reaches the eastern limit of its distribution in Israel.
The distribution of tiger beetles in different regions of Israel and the Sinai Peninsula.
Species/Subspecies
Choro-types
Israel
Siani (Egypt)
Galilee
Golan Heights (including Mt. Hermon)
Coastal Plain
Judea
Rift Valley
Negev
Northern Siani
Central Siani Foothills
Sinai Mountains
Southwestern Sinai
Upper
Lower
Northern
Central
Southern
Judean Desert
Judean Hills
Jordan Valley
Dead Sea Area
Arava Valley
Northern
Central
Southern
Calomeraaulicaaulica
SSS
●
●
●
●
●
●
Calomeralittoralisaulicoides
NAA
●
●
●
●
●
●
Calomeralittoraliswinkleri
SAT
●
●
●
●
●
Cephalotazarudnianavartianorum
INP
●
Cephalotatibialistibialis
EGYP
●
Cephalotalittorealittorea
NAA
●
●
Cicindelajavetiazari
LEVC
●
●
Cylinderacontortavaldenbergi
CPND
●
●
Habroderaniloticanilotica
AFT
●
Hypaethasingularis
NAA
●
Lophyraflexuosaflexuosa
WMA
●
●
●
●
●
●
●
●
●
●
●
Myriochilamelancholicamelancholica
AMC
●
●
●
●
●
●
●
●
●
●
●
●
●
●
●
Grammognathaeuphraticaeuphratica
SMS
●
●
●
●
●
●
●
Total for localities
3
2
3
5
4
3
1
1
3
6
5
2
2
3
5
1
4
7
4
3
5
2
6
3
9
Total for regions
8(9)
9
Chorotypes: AFT
– Afrotropical
, AMC
– Afrotropical-Mediterraneo-Centralasiatic
, INP
– Irano-Palestinian
, NAA
– NE-African-Arabian
, SAT
– S-Anatolia-Turanian
, SMS
– S-Mediterraneo-Sindian
, SSS
– Saharo-Sahelo-Sindian
, WMA
– W-Mediterraneo-N-African
, CPND
– Coastal Plain-Nile Delta endemic
, EGYP
– Egyptian endemic
, LEVC
– C-Levntian endemic
.
Three subspecies, C.contortavaldenbergi, C.javetiazari and C.zarudnianavartianorum, are characterized by a restricted distribution in Israel (Fig. 4). The first two first subspecies should be considered as regional endemics.
The nominative subspecies of Cylinderacontorta (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.contortavaldenbergi 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.
Cicindelajavetiazari 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.javetiazari inhabits the southern part of the species range area.
Cephalotazarudnianavartianorum 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.tibialistibialis, C.littorealittorea, H.singularis and H.niloticanilotica are never really observe in Israel (vsChikatunov et al. 2006). Among them C.tibialistibialis 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.littorealittorea 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 Hypaethasingularis 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 2002, 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 Habroderaniloticanilotica 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). First of them includes the faunas associated with southern part of the Great Rift Valley (Arava valley and Dead Sea area) and most part of the Sinai Peninsula, while the second combine most Israeli regions as well as Central Sinai Foothills. The last cluster diverges on the four groups. The fist combines assemblages of tiger beetles of the Mediterranean coastal habitats within the Northern and Central Coastal Plains. The communities typical for the arid habitats of the Negev Desert and the Central Sinai Foothills as well as for coastal habitats of the Southern Coastal Plain form the second group. The third group includes assemblages of the northern not seashore habitats of the Jordan Valley, Lower Galilee and Golan Heights. The last group is artificial, because the fauna of tiger beetles of Judea should be most similar to the fauna of the Dead Sea Area or the Northern Negev, while the fauna of tiger beetles of the Galilee, Jordan Valley and Golan Heights should be the most similar to each other. First of all, this discrepancy is due to a lack of data about tiger beetles of the central regions of Israel.
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 Calomeralittoralis (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 spring-fall 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.
The phenology of tiger beetles in Israel (grey – our data; pink – after Nussbaum 1987) and in the Egypt (green – after Alfieri 1975; blue – after Abdel-Dayem et al. 2003).
Cephalotazarudnianavartianorum
Spring
Grammognathaeuphraticaeuphratica
(Si)
Spring-summer
(Si)
Hypaethasingularis
Si
Si
Summer
Si
Si
Cephalotalittorealittorea
Si
Si
Si
Si
Si
Si
Si
Si
(Si)
(Si)
(Si)
Cephalotatibialistibialis
Si
Si
Si
Spring-fall
Si
Si
Si
Si
Si
Si
Si
Si
Si
Si
Si
Si
Si
Si
Calomeralittoralisaulicoides
Si
Si
Calomeralittoraliswinkleri
Cylinderacontortavaldenbergi
Cicindelajavetiazari
Myriochilamelancholicamelancholica
Habroderaniloticanilotica
Si
All-yaer
Calomeraaulicaaulica
(Si)
(Si)
(Si)
(Si)
Si
(Si)
Si
(Si)
(Si)
(Si)
(Si)
(Si)
(Si)
(Si)
(Si)
(Si)
Lophyraflexuosaflexuosa
(Si)
(Si)
(Si)
I
II
III
IV
V
VI
VII
VIII
IX
X
XI
XII
Notes. Si – records only on the Sinai Peninsula, (Si) – records including the Sinai Peninsula. The density of the grey color corresponds with the frequency of the records of species (subspecies):
1–3
4–6
7–9
10–12
13–15
16–18
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 (Jaskuła and Rewicz 2015; Jaskuła et al. 2015). For example, the activity of C.aulicaaulica, C.littoralisaulicoides, C.zarudnianavartianorum, C.contortavaldenbergi and G.euphraticaeuphratica start one-two months earlier, while the activity of C.aulicaaulica, C.littoralisaulicoides, M.melancholicamelancholica and G.euphraticaeuphratica finish one-three, and in the case with L.flexuosaflexuosa even six months later comparing with the data of Nussbaum (1987). On the other hand, Nussbaum (1987) indicated longer period of activity of C.littoraliswinkleri and C.tibialistibialis as well as the later finish of the activity of C.contortavaldenbergi and C.javetiazari (Table 2).
Similarly, the periods of activity of C.aulicaaulica, L.flexuosaflexuosa and M.melancholicamelancholica 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.aulicaaulica records only in June and July (Jaskuła and Rewicz 2015), 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.flexuosaflexuosa lasts from March-April to July (Jaskuła and Rewicz 2015; Jaskuła et al. 2015) but in Israel it continues from February to December (Table 2).
On the contrary, in Tunisia the activity of G.euphraticaeuphratica begins in March and ends in July (Jaskuła and Rewicz 2015) 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 (Jaskuła et al. 2015). The same situation is observed for different subspecies of Cephalotalittorea (Forskål, 1775) as well as C.littoralis. In Tunisia C.littoreagouditii (Dejean, 1829) is active from May to October (Jaskuła and Rewicz 2015) while the period of activity of C.littorealittorea on the Sinai Peninsula lasts from May to September (Table 2). The activity of C.littoralislittoralis in Morocco is observed from April to October (Jaskuła et al. 2015) and in Tunisia from March to August (Jaskuła and Rewicz 2015), while the activity of C.littoralisaulicoides in Israel and on the Sinai Peninsula as well as C.littoraliswinkleri 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 Oxycheilopsiscretacicus 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, 2011; Casale 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 Paleogene – early 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 Cicindelahybrida 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, 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 Cicindelacampestris group (Cassola 1999; Franzen 2007; Deuve 2011, 2012; our 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.
Acknowledgements
We are very grateful to Dr. Laibale Friedman (Tel Aviv, Israel), Dr. Alexander Puchkov (Kiev, Ukraine), Mr. Jürgen Wiesner (Wolfsburg, Germany), Mr. Igor’ Ovsyannikov, Mr. Alexander Sokolov, Mr. Pavel Udovichenko and Mr. Eugeny Shankhiza (all Moscow, Russia) who kindly has given material and information for our study. Special thanks to Prof. David L. Pearson (Arizona State University, USA) for revising the English text. For the first author the study received financial support from the Ministry of Education and Science of the Russian Federation (Project No 6.632.2014/K).
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