Calosoma (Caminara) imbricatum Klug, 1832

Each elytron of C. 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). C. imbricatum is the only species of Caminara that has this model of sculture of the elytra.
Does exist in South West Africa a population (hottentottum) that seems to be isolated and that is well characterized by its slimmer body shape and by a lively metallic color, but in the remaining area in which it is diffused, C. 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, which he even included in a different genus (Campalita), and recognized for C. imbricatum a limited number of subspecies. Later Mandl (1970: 62), considered deserticola as a mere subspecies of C. imbricatum, having described (1953, 1967, 1970) some more populations of the latter.
In our opinion, except hottentottum and deserticola, the remaining populations of C. imbricatum, that are widespread from Cabo Verde archipelago through the Sahelian zone to Arabia, Syria, Iraq, Iran, Pakistan, do not present significant differences except for a slight variation in size. However, 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 (described from: Cap Vert); lectotype in Museum für Naturkunde, Berlin (Andrewes, 1929: 64)
Caminara arabica Motschulsky, 1865:304 (described from Arabia); type material: 1 ♂, depository not stated
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 ssp. loeffleri Mandl, 1953: 57 (type locality: Rigmati); holotipe ♂ and 20 paratipes in Naturhistorisches Museum, Wien, Zoologisches Forschungsmuseum Alexander Koenig, Bonn (Ulmen et al. 2010: 10) and in Naturhistorischen Museum Basel [examined]
Calosoma (Caminara) imbricatum loeffleri m. rufoappendiculata Mandl, 1967: 44 (described from Sistan and Baluchistan, Bampour) type material 1 ♂ depository not stated
Calosoma (Caminara) imbricatum linnavuorii Mandl, 1968c: 128 (type locality: Somalia, Hargheissa); holotype ♂ in Zoologischen Museum Helsinki, paratype in Naturhistorischen Museum Basel [examined]
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.
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 hottentottum from the locality of Sultan Hamud in Makueni Province and Mandl (1970: 61) has attributed this citation to his C. imbricatum linnavuorii.
Still from West Kenya (Amboseli park, mid-way between Sultan Hamud and Taveta and here and there along the Tanzanian border in the Rift Valley Province), we have seen specimens with more pronounced characteristics of sculpture and color, to the point that could be considered transitional towards C. imbricatum hottentottum. These specimens have bronze lustre color on the upper side of the body and rougher elytral sculpture than C imbricatum imbricatum, though, if compared with C. imbricatum hottentottum, they have sometimes a stouter shape and a darker colour of the upper body and in any case they have head and pronotum less deeply punctate.
One might conclude with Jeannel (1940: 105) that C. imbricatum hottentottum is present in western Kenya (Bruschi, 2013: 76), or one might consider these specimens as simple individual variations within C imbricatum imbricatum. However, before definitevely deciding if these specimens are part of an omogeneous population and eventually which relationships there are between this population and C. imbricatum hottentottum, remain to explain the presence of specimens of the typical form in this same region and the strange fact that no finding of specimens of C imbricatum has been published up to now in the wide 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), Syria, 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 (Fort Flatters); Tassili n'Ajjer (Mountains Domergue) (Jeannel, 1940: 105).
Burkina Faso. Gorom-gorom, Essakane (SB)
Cabo Verde (Republic of). Ilha de Boa Vista (SB); Ilha Brava (Wollaston, 1867: 6); Ilha do Fogo: (Breuning, 1927: 221), São Filipe (Herbert, 1987: 97); Ilha do Maio (Oromí & al., 2005: 79); Ilha do Sal: (RMCA), Espargo (Herbert, 1987: 97); Ilha de Santo Antão (Breuning, 1927: 221); Ilha de São Nicolau (RMCA); Ilha de Santiago: (Wollaston, 1867: 6), Praia (Herbert, 1987: 97); Ilha de São Vicente: (Wollaston, 1867: 6), São Pedro (Herbert, 1987: 97).
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, Wadi Halfa, Gebel Elba (Schatzmayr, 1936: 24), Marsa Halaib, Wadi Haideb, Abu Simbel, Sallum, Maryut lake (Alfieri, 1976: 3).
Eritrea. Wachiro (AC); Massaua (SB).
Ethiopia. Gamo Gofa: Kelam (AVT); Oromia: Langano Lake (SB); 40 km W Konza (SB).
India. Thar Desert (Mandl, 1970: 63).
Iran. Fars (Mandl, 1970: 63); Hormozgan: Abad Geno, 40 km nord Bander Abbas (EM), 45Km East of Bandar Abbas (SB), Posht-Koh Village, Minab road (Azadbakhsh & al., 2015b: 226), Hasan-Langi (Azadbakhsh & al. 2015a: 45); Kerman: Rigmati, (paratype of loffleri) (NMB); Lorestan: Firouz Abad (Mandl, 1970: 63); Sistan and Baluchistan: Bampur, Khasch, Iranshahr, Irafshan (Mandl, 1970: 63); Tehran: 70km SW Tehran, Yusefabad (Mandl: 1967; 458).
Iraq. Basrah (Andrewes, 1929: 64), Ramadi (Mandl: 1967; 458).
Kenya. North Eastern Province: El Wak (SB); Eastern Province: Laisamis (SB); Marsabit to South Orr (Häckel et al, 2016: 15); Taveta (Breuning, 1927: 221); Rift Valley Province, Amboseli Nat park (SB; sub hottentottum Häckel et al, 2016: 15), Gakong (SB), Oltepesi (SB); Makueni Province: Sultan Hamid (sub hottentottum, 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: Wadi Quitbit (NMP), Taquah env Samahram(SB)
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, Rawdhat Khorim NP 550m (Abdel-Dayem & al., 2017; SB); Riyadh prov. NE of Quwayiyah 728m (SB); Al-Baha prov: El-Hawya, Wadi Turabet Zahran (El-Hawagry & al, 2013).
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)
Syria. as-Suwayda: Al Kafr (SB)
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 ottentottum Chaudoir, 1852

Calosoma hottentottum Chaudoir, 1852: 99 (described from: Cap de Bonne Espérance); lectotype ♂ designated by Deuve (1978: 249), in Muséum National d'Histoire Naturelle, Paris [examined]
Calosoma (Caminara) imbricatum ssp. hottentottum Breuning, 1927: 221
Caminara (Caminara) imbricata imbricata hottentota Lapouge, 1932: 410
Caminara imbricatum subsp. 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 hottentottum in all cases has the upper part with bronze reflexes (not bluish as in the other populations).
C imbricatum hottentottum is 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 hottentottum, to which they add doubtfully the type of C. aetiops from Niger, that they regard as a synonym for C. imbricatum hottentottum. 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); Erongo: Omaruru (SB), Usakos 900m. (SB), between Omaruru and Uis (EM); Klein Karas Mts. (RMCA); Hardap reg: Maltahohe 1350m. (SB), Kalkrand 1226m (SB), Rehoboth 1400m; Khomas prov: Nauchas 1770m (SB); Otjitambi (Mandl, 1970: 62); Otjozondjupa reg: Otjivarongo; between Gobabis and Aranos (Häckel et al, 2016: 15); Walvis Bay, Great Karas Mts., Kaross, Rehoboth, Kunene reg. (IZICO,
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 (

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

Calosoma (Caminara) imbricatum hottentottum
Chaudoir, 1852
Cap de Bonne Espérance
(lectotype of Calosoma hottentota Chaudoir, 1852)
(Muséum National d'Histoire Naturelle, Paris)
Calosoma (Caminara) imbricatum hottentottum
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 hottentottum
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 (described from: Transcaspia); holotype in Zoological Institute, Russian Academy of Sciences, St. Petersburg (Jeannel, 1940: 116)
Calosoma (Caminara) deserticola Breuning, 1927: 222
Caminara (Caminara) imbricata andrewesi deserticola Lapouge, 1932: 410
Campalita deserticola Jeannel, 1940: 116
Calosoma (Caminara) imbricatum ssp. deserticola Mandl, 1969: 2
Caminara deserticola Basilewski, 1972: 37
Calosoma (Caminara) deserticola ssp. augustasi Obydov, 2005: 5 (type locality: Mongolia, Gobi-Altai aimag), holotype in 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. Kerman: Jussuffabad (Taybad) (Mandl, 1970: 61); Razavi Khorasan: Faruj (SB); Teheran: 50km south-west Teheran (Mandl, 1970: 61)
Kazakhstan. Mangystau reg.: Beyneu env.(SB)
Mongolia. Bayankhongor aimag: 60 km S. Bayankhongor 1640m (sub deserticola augustasi, Obidov, 2005: 7); Gobi-Altai aimak: Alag-Nur Lake (SB); 30 km SW Bugat 1270m; Zahuin Gobi Desert 1090m (Obidov, 2005: 7); (Obidov, 2005: 7); Ömnögovǐ aimag: Gurvan tés (Mandl, 1970: 61); 2-3 km SO Dund gol, (Mandl, 1969: 2; MTTM,
Russia.70 km S. Astrachan' (SB) 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 var. andrewesi Breuning, 1928: 95 (type locality: Assam) holotype ♂ in Naturalis Biodiversity Centre, Leiden (de Boer, 2002: 27)
Caminara (Caminara) imbricata andrewesi Lapouge, 1932: 410
Caminara imbricatum Subsp. 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: 223), 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 correctly identified C. orientale with the species until then called 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
India. Rajasthan: Jaipur (SB), Mount Abu (RS), Bikaner – Jaisalmer (RS); West Bengala: north of Bengal Presidency (Chaudoir, 1869: 368); Assam (holotype NBC, Breuning, 1928b: 95)

Notes: 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.
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.

Calosoma (Caminara) imbricatum andrewesi
Breuning, 1928
India: Rajasthan, Mount Abu, (coll. R. Sciaky)
Calosoma (Caminara) imbricatum andrewesi
Breuning, 1928
India: Rajasthan, Jaipur, VII 1988, Pad. Mal. leg.
updated Oct 14 2019