Calosoma (Caminara) imbricatum Klug, 1832

Each elytron of imbricatum has 16 intervals, it is to say that only one tertiary interval exists on each sides of a secondary one ("triploïde" type of Jeannel). imbricatum is the only species of Caminara that has this model of sculture of the elytra.
Does exist in South West Africa a population (hottentotum) that seems to be isolated and that is well characterized by its slimmer body shape and unique colour, but in the remaining area in which it is diffused, imbricatum presents a very scarce differentiation, concerning some variation in size and body shape, or the greater or lesser visibility of the sculpture of the elytra.
However, in the north east of the distribution area (Central Asia) we find a population (deserticola) that has a larger and more elongated body with less raised elytral sculpture. Jeannel (1940) considered deserticola as a separated species, that he even included in a different genus (Campalita), and recognized for imbricatum a limited number of subspecies. Later Mandl (1970: 62), considered deserticola as a mere subspecies of imbricatum, and described (1953, 1967, 1970) some more populations of the latter.
In our opinion, except hottentotum and deserticola, the remaining populations of imbricatum, that are widespread from Cabo Verde archipelago through the Sahelian zone to Arabia, Iraq, Iran, Pakistan, do not present significant differences except for a slight variation in size, but in central and northern Pakistan do exist populations with transitional characteristics towards deserticola .
Finally, from further south (Central Indian Peninsula), has been described another population, also of relatively large size, but stouter and with a still more flattened sculpture of elytra (andrewesi), that possibly also could be maintained at subspecies level.

Calosoma (Caminara) imbricatum imbricatum Klug, 1832

Calosoma imbricatum Klug, 1832: pl.XI (Typus: Cap Vert; Museum für Naturkunde Humboldt Universität, Berlin)
Caminara arabica Motschulsky, 1865:304 (Typus: Arabia)
Calosoma (Caminara) imbricatum Breuning, 1927: 221
Caminara (Caminara) imbricata Lapouge, 1932: 410
Caminara (Caminara) imbricata arabica Lapouge, 1932: 410
Caminara imbricatum Jeannel, 1940: 104
Calosoma (Caminara) imbricatum loeffleri Mandl, 1953: 57 (Typus: Iran, Kerman, Rigmati; Zoologisches Forschungsmuseum Alexander Koenig, Bonn)
Calosoma (Caminara) imbricatum linnavuorii Mandl, 1968c: 128 (Typus: Somalia, Hargheissa; Zoologischen Museum Helsinki)
Calosoma (Caminara) imbricatum loeffleri m. rufoappendiculata Mandl, 1967: 44 (Typus: Iran, Sistan and Baluchistan, Bampour)
Calosoma (Caminara) imbricatum ssp. linnavouorii Mandl, 1970: 61

Length 15-22 mm. What we consider as the typical populations are the ones with individuals of small or medium size having finely punctate head and pronotum, elytra in oval widened, and elytral sculpture with scaly intervals. The upper body is black, with faint bluish reflexes.
C. imbricatum imbricatum continuously spreads in Africa, excluding the area of equatorial forest, from the Canary and Cabo Verte islands, throughout the Sahelian zone, Algeria, Libia, Senegal, Mali, Niger, Chad, to Sudan, Eritrea, Djibuti, Ethiopia, Somalia, Kenya.
Here and there in East Africa it is possible to find single specimens with slightly more wrinkled head and a little stronger elytral sculpture. On this basis Mandl (1968c: 128) described from Somalia the subspecies linnavuorii that he considered intermediate between the typical one and ottenttotum and that would be present throughout the Horn of Africa from Somalia to Kenya. However, the inconsistency of the characters does not seem to justify this subspecies.
In Kenya, the situation is more complicate. In north-eastern Kenya, we find some populations definitely referable to C imbricatum imbricatum and Breuning (1927: 221) reports the nominate subspecies from south west (Taveta, very close to the Tanzanian border). Jeannel (1940: 105) cites C. imbricatum hottentotum from the locality of Sultan Hamud in Makueni Province and Mandl (1970: 61) attributed this citation to his C. imbricatum linnavuorii.
Still from South West Kenya (Amboseli park, mid-way between Sultan Hamud and Taveta), we have seen specimens with more pronounced characteristics of sculpture and color, to the point that could be considered transitional towards C. imbricatum hottentotum. These specimens have bronze lustre of the upper side of the body and rougher elytral sculpture than C imbricatum imbricatum, though, if compared with C. imbricatum hottentotum, they have a stouter shape, a darker colour of the upper body and, slightly less deeply punctate, head and pronotum .
One might conclude with Jeannel (1940: 105) that C. imbricatum hottentotum is present in Kenya (Bruschi, 2013: 76), or consider these specimens as simple individual variations within C imbricatum imbricatum. However, before to definitevely decide if these specimens are part of an omogeneous population and eventually which relationships there are between this population and C. imbricatum hottentotum, remain to explain the citation of Breuning of the typical form from this same region and the strange fact that no captures of specimens of C imbricatum have been published up to now in the vast area between Kenya and South Africa.
Going to East, the roughness of the head and pronotum decreases and the elytral sculpture tends to flatten. We find populations variously named, but that can also be referred to the typical form and that live in the Arabic peninsula: Yemen, Kuwait, Saudi Arabia, Oman, Qatar and Iraq (which all Mandl considers as C. imbricatum arabicum), southern Iran (C. imbricatum loeffleri), up to south western Pakistan. The populations of central and northern Pakistan someway accentuate these characteristic and look as transitional to C. imbricatum deserticola.

Examined specimens and literature’s data
Algeria: Temassinin; Monts Doumergue (Tassili n'Ajjer) (Jeannel, 1940: 105)
Burkina Faso: Gorom-gorom, Essakane (SB)
Cabo Verde (Republic of): São Vicente, São Jago, Brava (Wollaston, 1867: 6); São Antão Archibispo, Fogo (Breuning, 1927: 221); São Nicolau (RMCA); Ilha do Sal (RMCA), Boa Vista, (SB)
Canaries Islands (Spain): Tenerife (Andrewes, 1929: 64).
Djibuti: Djibuti (EM, SB); Ambouli (DP, EM, SB); Obock (RMCA, SB), bay of Tadjoura (Mandl, 1968: 130).
Egypt: Alexandria (Breuning, 1927: 222); Gulf of Solum, El Hamam, Assuan, Gebel Elba (Schatzmayr, 1936: 24), Marsa Halaib, Wadi Haideb, Habu Simbel (Alfieri, 1976: 3).
Eritrea: Wachiro (AC); Massaua (SB).
Ethiopia: Gamo Gofa: Kelam (AVT); Oromia: Langano Lake (SB); 40 km W Konza (SB).
Georgia. Adjara: Gonio (http://molbiol.ru/).
India. Thar Desert (Mandl, 1970: 63).
Irak: Basrah (Andrewes, 1929: 64).
Iran: Abad Geno, 40 km nord Bander Abbas (EM); Rigmati, (paratype of loffleri) (NMB); Sistan and Baluchistan: Bampour, Khasch, Iranshahr, Irafshan; Fars; Lorestan: Firouz Abad (Mandl, 1970: 63), Hormozgan prov: Posht-Koh Village; Minab road (Azadbakhsh & al., 2015: 226)
Kenya. North Eastern Province: El Wak (SB); Marsabit to South Orr (Häckel et al, 2016: 15); Taveta (Breuning, 1927: 221); Rift Valley Province, Amboseli Nat park (SB; sub hottentotum Häckel et al, 2016: 15); Makueni Province: Sultan Hamid (sub hottentotum, Jeannel, 1940: 105).
Kuwait: Sha`ab, Erifjan, Sobiya (sub deserticola Al-Houty, 2004: 62).
Lybia: Fazzan, suburb of Ghat (JCR).
Mali: Timbuctu (Jeannel, 1940: 105).
Niger: Aïr, Tabélot (RMCA); Agades (Breuning, 1927: 221); Nguigmi (Mandl, 1970: 61)
Oman: Jebel Huwarrah (DP); Dhofar prov., Quitbit (NMP)
Pakistan. Baluchistan: Ashtola island (Andrewes, 1929: 64), Khusdar (SB), Quetta (SB), SW Quetta (Mandl, 1970: 63); Punjab: Bahawalpur (NIM), Meyal near Attock (EM); Khyber Pakhtunkhwa: Kohat (NIM); Sindh: Mithi (NIM), Karachi (Andrewes, 1929: 64)
Qatar: Abu Samrah (Abdu & Shaumar, 1985)
Saudi Arabia: Rub al Khati (AMNH), Jidda (Jeddah) (RMCA); Hejaz (Andrewes, 1929: 64); Riyadh (Wittmer & al., 1979: 527); 100 km NE Riyadh, Rawdat Khuraim 550m (SB)
Senegal: Richard Tol (SB)
Somalia: Genale (AC); Hargheissa (sub linnavuorii Mandl, 1968c: 128); 25km NW Berbera, Daragodleh (sub linnavuorii Mandl, 1968c: 128); 25 km S of El Buh (El Haga) (RMCA); El Afweyn (El Afweina) (BMNH; Vigna Taglianti & Bruschi, 1988: 238)
Sudan: Khartum (NMP), Port Sudan (NMP), El Fasher (RMCA); Ambukohl (Breuning, 1927: 221); Wadi Halfa (Schatzmayr, 1936: 24); Kassala, Abend Pass (Mandl, 1968c: 128); Suakin (Mandl, 1970: 63)
Tchad: Nguigmi (Mandl, 1970: 63)
United Arab Emirates: Ras Al Khaimah, Tawi as Saman (SB), Shardza (Al Sharjah) (SB)
Yemen. Is. Socotra: Hadibu plain (BMNH; Vigna Taglianti & Bruschi, 1988: 240), Shuab (SB)

Notes: Winged, attracted to light at night. Taking into account the wide distribution area, almost entirely included in the equatorial, tropical and sub-tropical zones, the period of activity covers the whole year but varies locally influenced by the precipitation regimes.

Calosoma (Caminara) imbricatum imbricatum
Klug, 1832
Eritrea: Massaua, XII.1972, Boemi lg.
Calosoma (Caminara) imbricatum imbricatum
Klug, 1832
Djibuti II.1990
Calosoma (Caminara) imbricatum imbricatum
Klug, 1832
Somalia, N. Hargheissa,23-28.VI.63. paratype of Calosoma (Caminara) imbricatum linnavuorii Mandl, 1968.
(coll. Naturhistorischen Museum Basel)
Calosoma (Caminara) imbricatum imbricatum
Klug, 1832
Sud Est Persien, Rigmati, West Deghaz, Marian 1950
paratype of Calosoma (Caminara) imbricatum loeffleri Mandl, 1953
(coll. Naturhistorischen Museum Basel)
Calosoma (Caminara) imbricatum imbricatum
Chaudoir, 1852
Djibouti: Ambouli, leg. G.Nazaret (coll. D. Peslier)
Calosoma (Caminara) imbricatum imbricatum
Chaudoir, 1852
Kenya: Amboseli Nat. Park, 25.Jan.2001,Nádai Lázló lg.

Calosoma (Caminara) imbricatum imbricatum
Klug, 1832
Iran: Abad Geno, 40km nordlich von Bander Abbas,
7.IV.1972, F Ressl leg. (coll.E. Migliaccio)
Calosoma (Caminara) imbricatum imbricatum
Klug, 1832
Pakistan: Baluchistan Prov., Khusdar,
23.07.09, Muhammad Akter


Calosoma (Caminara) imbricatum ottentotum Chaudoir, 1852

Calosoma hottentota Chaudoir, 1852: 99 (lectotype: Cap de Bonne Espérance; Muséum National d'Histoire Naturelle, Paris)
Calosoma (Caminara) imbricatum hottentotum Breuning, 1927: 221
Caminara (Caminara) imbricata imbricata hottentota Lapouge, 1932: 410
Caminara imbricatum hottentota Jeannel, 1940: 104

Length 18-20 mm. This population is distinguished from the others because of head and pronotum with large punctures and with deep, transverse, wrinkles and because of the rougher sculpture of the elytra. Moreover, C imbricatum hottentotum in all cases has the upper part with weak bronze reflexes (not bluish as in the other populations).
C imbricatum hottentotum is well known from, and possibly restricted to, Namibia and South Africa.
Coming back to the idea already expressed by Jeannel (1940: 105) and initially shared by Bruschi (2013: 76), Hackel & al. (2016) consider the specimens from South West Kenya (Amboseli park) as C. imbricatum hottentotum, to which they add doubtfully the type of C. aetiops from Niger, that they regard as a synonym for C. imbricatum hottentotum. Thereby the distribution area of this subspecies would be greatly increased and constituted by an array of widely separated populations.

Examined specimens and literature’s data
Namibia: Mariental 1120 m (SB); Okahandja Gross Barmen (EM); Omaruru Uis (EM, SB); Klein Karas (RMCA); Hardap reg., Maltahohe 1350m. (SB); Erongo prov., Usakos 900m. (SB); Otjitambi (Mandl, 1970: 62); Otjozondjupa reg. Otjivarongo; between Gobabis and Aranos (Häckel et al, 2016: 15).
South Africa: Cap de Bonne Espérance (lectotype) (MNHN); Western Cape: Upington (RMCA), Pniel (Jeannel, 1940: 105); Northern Cape: Kimberley (Breuning, 1927: 223), SW Kimberley, 13 km SW Ritchie (Häckel et al, 2016: 15); Augrabies National Park, Orange river (http://assazhnev.narod.ru/carab_atlas.html).

Notes: Winged, attracted to light at night. Active individuals were captured from January to March

Calosoma (Caminara) imbricatum hottentotum
Chaudoir, 1852
Cap de Bonne Espérance
(lectotype of Calosoma hottentota Chaudoir, 1852)
(Muséum National d'Histoire Naturelle, Paris)
Calosoma (Caminara) imbricatum hottentotum
Chaudoir, 1852
Namibia: Mariental 1120 m., Mariental town, 24°37' S, 17°57'E.,
3-4.III.97, Audisio, Bologna, Zapparoli lg.

Calosoma (Caminara) imbricatum hottentotum
Chaudoir, 1852
Namibia: Mariental 1120 m., Mariental town, 24°37' S, 17°57'E.,
3-4.III.97, Audisio, Bologna, Zapparoli lg.

Calosoma (Caminara) imbricatum deserticola Semenov, 1897

Calosoma deserticola Semenov, 1897: 242 (Typus: Transcaspia; Zoological Institute, Russian Academy of Sciences, St. Petersburg)
Calosoma (Caminara) deserticola Breuning, 1927: 222
Caminara (Caminara) imbricata andrewesi deserticola Lapouge, 1932: 410
Campalita deserticola Jeannel, 1940: 116
Calosoma (Caminara) imbricatum deserticola Mandl, 1969: 2
Caminara deserticola Basilewski, 1972: 37
Calosoma (Caminara) deserticola augustasi Obydov, 2005: 5 (Holotype Mongolia, Gobi-Altai Aimak, Timiryazev State Museum of Biology Moscow)

Length 22-28 mm. Described by Semenov (1897) as a species and maintained as such by Breuning (1927) and Jeannel (1940), it has been considered by Lapouge (1932) and later by Mandl (1969) as a mere subspecies of C. imbricatum. In fact C. imbricatum deserticola is quite similar to the typical C. imbricatum imbricatum and only can be distinguished because of the constantly larger body size, the more flat sculpture of the elytra and by the black colour without bluish or bronze reflections.
The populations of Central Asia mostly have larger body and less expanded elytra, but going east (Mongolia) the individuals became smaller and stouter, although maintaining the same model of elytral sculpture. Obidov (2005: 5) considered the Mongolian population as a distinc taxon (C. deserticola augustasi). The feebles differences that he used in his diagnosys, apart from the lesser size, do not deserve to be taken into consideration in a such variable species as the complex of C. imbricatum is. Thus we can simply conclude that C. imbricatum deserticola in Mongolia is represented by an isolated, but very little differentiated, population. As for the populations of central and northern Pakistan, it is possible that these represent a transition between C. imbricatum deserticola and the typical form.

Examined specimens and literature’s data
Afghanistan. Herat: Bala Murghab; Maimana: Ali Gul; Wadi-ye Namak Shor; Badghis: Mangan (Mandl, 1970: 61)
Iran. North Khorasan, Faruj (SB); 50km south-west Teheran (Mandl, 1970: 61); Jussuffabad (Taybad) (Mandl, 1970: 61)
Kazakhstan: Mangystau reg.: Beyneu env.(SB)
Mongolia: South Gobi (Mandl, 1970: 61); Gobi-Altai aimak: Alag-Nur Lake (SB); Gobi-Altai Aimak, 30 km SW Bugat 1270m, Zahuin Gobi Desert 1090m; Bayankhongor Aimak: 60 km S. Bayankhongor1640m (Obidov, 2005: 7)
Turkmenistan: Nebit-Dag (SB); Kopet Dag, Karakata vill. (SB); Repetek Nat. reserve (SB); Merw (SB); Karakum desert, 50 km. SE Chardjew (SB); Kizilkum desert (Jeannel, 1940: 117); Kisil Arwat (EM); Ljutfabad (EM); Türkmenbaşy (Krasnovodsk) (Jeannel, 1940: 117), Tedschen (Mandl, 1970: 61)
Uzbekistan: Taskent (SB); Khorezm reg., Karakapakstan rep. (Khamraev, 2003: 47); Kizil Kum desert (Mandl, 1970: 61)

Notes: C. imbricatum deserticola is winged and inhabits sandy semidesertic steppes, sandy semideserts and sand deserts, where it hides in the sand during the day. At night it hunts and is an excellent climber. Active individuals were captured in spring from the middle of April to the beginning of July

Calosoma (Caminara) imbricatum deserticola
Semenov, 1897
Turkmenistan: Merw, juin 1896
Calosoma (Caminara) imbricatum deserticola
Semenov, 1897
Turkmenistan: Repetek (Nat. reserve),
15-30.IV.1990, Sendek lgt.
Calosoma (Caminara) imbricatum deserticola
Semenov, 1897
SW Mongolia: Gobi-Altai aimak, Dzhungarian Gobi,
Alag-Nur Lake 1300m. (near Ajlyn-Tsan-Khuduk),
1-2.VI.2011 Yakovlev lgt.

Calosoma (Caminara) imbricatum andrewesi Breuning, 1928

Calosoma (Caminara) imbricatum andrewesi Breuning, 1928: 95 (Typus: Assam; Naturalis Biodiversity Centre, Leiden)
Caminara (Caminara) imbricata andrewesi Lapouge, 1932: 410
Caminara imbricatum andrewesi Jeannel, 1940: 104

Length 20-22 mm. Chaudoir (1869: 368) first described a specimens of a Calosoma coming from India and tentatively identified it as the Calosoma orientale previously named by Hope. Later Breuning (1927: 222), once ascertained the presence of a population of C. imbricatum in India, kept the name Calosoma imbricatum orientale for it and so endorsed the misunderstanding of Chaudoir. Only more later Breuning (1928: 95), when recognized the identity of C. orientale con C. squamigerum renamed the new subspecies C. imbricatum andrewesi.
C. imbricatum andrewesi consists of populations of a relatively large size, with a stout body and flattened elytral sculpture. The colour of the upper body is a dull, not metallic, black.
It has been found in few localities of northern India (Rajasthan, Bengal, Assam), where it seems quite rare.

Examined specimens and literature’s data
Rajasthan, Jaipur (SB); West Bengala, north of Bengal Presidency (Chaudoir, 1869: 368); Assam (holotype NBC, Breuning, 1928b: 95)

Notes: The subspecies has been named after Herbert Edward Andrewes (1863 - 1950) an English entomologist, specialist in Carabidae, that worked at the Indian Forest Service and later at the Natural History Museum in London.
As the other populations of the same species, C. imbricatum andrewesi is winged and attracted to light at night. Active individuals were captured in July.

Calosoma (Caminara) imbricatum andrewesi
Breuning, 1928
India: Rajasthan, Jaipur, VII 1988, Pad. Mal. leg.

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