Calosoma aethiops (Jeannel, 1940) as a new synonym of Calosoma imbricatum hottentotum Chaudoir, 1852, a new status of Calosoma roeschkei Breuning, 1927, and a revision of the Calosoma senegalense group sensu Häckel, 2012 (Coleoptera, Carabidae, Carabini).

Calosoma aethiops (Jeannel, 1940) as a new synonym of Calosoma imbricatum hottentotum Chaudoir, 1852, a new status of Calosoma roeschkei Breuning, 1927, and a revision of the Calosoma senegalense group sensu Häckel, 2012 (Coleoptera: Carabidae: Carabini). Conducted is a taxonomic revision of the Calosoma senegalense group sensu Häckel, 2012. Placed in the group sensu stricto are four species: Calosoma planicolle Chaudoir, 1869, Calosoma scabrosum Chaudoir, 1843, Calosoma senegalense Dejean, 1831, and Ctenosta strandi Breuning, 1934. Calosoma aethiops Jeannel, 1940 is synonymized with Calosoma imbricatum hottentotum Chaudoir, 1852, and Calosoma roeschkei Breuning, 1927 is newly regarded as a subspecies of Calosoma scabrosum. The taxonomic conclusions are based on morphometry of the holotypes and 10 male and 10 female specimens of each taxon, and on morphology of the aedeagus including inflated endophalus.

Introduction

is the second most speciose genus of the subfamily , with 168 (Lorenz 2005), 128 (Bruschi 2013) or 129 (Häckel 2013) species described from all zoogeographic regions. Most species are excellent fliers widely distributed on all continents and numerous islands, but some are secondarily brachypterous or apterous with narrow distributions (Bruschi 2013, Häckel 2012). Some species inhabit more zoogeographic regions, and some extend to a neighboring continent that belongs in the same zoogeographic region (Häckel 2012). Examples of such distributions are some species of the group ( subgroup sensu Häckel 2013) and of the group ( subgroup sensu Häckel 2013), which inhabit the area of the Horn of Africa. The distributions are probably to some extent responsible for the unsettled situation in the species-level taxonomy of the group (Bruschi 2013, Häckel 2012, 2013, Häckel & Farkač 2012), a part of which this paper attempts to resolve. At the same time it respects the recent supraspecific classification (Häckel 2013), which in light of the known genetic analyses (Su et al. 2005) does not support the traditional subgeneric divisions.

Material and methods

The classification of the group is based primarily on external structural details of the adult, with species-level taxonomy relying also on structural details of the expanded endophalus. The aedeagi were dissected, preserved, studied dry and glued on cards appended beneath the dissected specimens. For study of the endophalus, the aedeagus was soaked 48 hours in 1:1 solution of water and 8% acetic acid, and then the endophalus was inflated using a small Heavy Duty (12V) compressor normally used to inflate tires, set at medium pressure. Fixation of the endophalus morphology was secured by slow drying on a portable electric (220V) single-plate heater, and the whole aedeagus-endophalus preparation was then glued on a paper card. The preparations were photographed by Canon G10 Digital Compact in macrophoto regime with flash. Aedeagi were photographed in the right lateral view, with details of their tips also slanted at an angle.

Inspected and evaluated were the following morphometric parameters of the holotypes and 20 samples (10 males and 10 females) of each species:

a) ,

total length of the adult including mandibles

b) ,

maximum head width including eyes to maximum pronotal width ratio

c) ,

maximum pronotal length to maximum width ratio

d) .

maximum elytral length to maximum width ratio

Subjective evaluations included also differences in termination of the aedeagus (apex) and of the inflated endophalus. Measured were morphometric parameters of 10 males and 10 females of the following taxa: , , , , (all taxa belong to group), PageBreakand from populations inhabiting the Afrotropical region (including Oman and Yemen on the Arab peninsula) and ( group, subgroup). Measured holotypes include (Chaudoir), (Breuning), (Chaudoir) and (Jeannel).

The material examined is housed in the collections listed below:

cMNHN

Muséum national d’Histoire naturelle, Paris, France

cNBCL

National Biodiversity Center, Leiden, Netherlands

cNMP

Národní muzeum, Prague, Czech Republic

cFAR

Jan Farkač collection, Prague, Czech Republic

cHAC

Martin Häckel collection, Hostivice, Czech Republic

cSEH

Rostislav Sehnal collection, Unhošť, Czech Republic

cWRA

David W. Wrase collection, Berlin, Germany

Systematic part

species group (sensu Häckel 2013):

subgroup (= group s. str.)

Chaudoir, 1869

Plate 4: Fig. 4

Plate 4.

Aedeagi of species group (sensu Häckel 2012: 30) and (Namibia). a – aedeagus with expanded endophalus in right lateral view, b – same in left lateral view, c – same in anterior view 1 (Oman: Dhofar) 2 (southeastern Kenya: Voi env.) 3 (northeastern Kenya: El Wak) 4 (Namibia: Kavango) 5 (Zimbabwe: Shamva) 6 (Namibia: Okahandja).

Chaudoir, 1869: 369 (type loc. “près du Zambéze”, type in

Harold, 1880: 260 (type loc. “Taita District, Kenya Colony, Ukamba”), syn. sn.

Breuning, 1927: 188. Lapouge 1932: 415;

Jeannel, 1940: 130.

Häckel, 2013: 30.

Material studied.

BOTSWANA. 1♂, 3♀: Ngamiland district, ne. of Maun, Tamalakane (cHAC).

KENYA. 1♂, 1♀: Tsavo, Mtitoanday (cSEH); 1♂, 1♀: Eastern 729, Sosoma, 202 km E of Thika (cSEH).

MOZAMBIQUE. 1♀: Sofala province, 30 km S Caia (cSEH).

Namibia. 1♂: Kavango reg., Okavango river, Rundu, 1050 m (cHAC); 1♂, 1♀: Caprivi reg., Bagani- Popa Falls (cHAC).

Zambia. 1♂: Southern Prov., Livingstone env., Victoria Falls (cHAC).

Zimbabwe. 1♂, 2♀: Midlands Prov., Kwekwe env. 20 km w. Ngezi Park (cHAC); 1♂, 1♀: Matabeleland Prov., 60 km N of Bulawayo, Marapoosa road (cHAC); 2♂: Masvingo province, 95 km NE Beitbridge, Bubi river (cSEH)

Distribution.

Angola, Botswana, Democratic Congo, Ethiopia, Kenya, Lesotho, Madagascar, Malawi, Mozambique, Republic of South Africa, Somalia, Tanzania, Swaziland, Uganda, Zambia, Zimbabwe.

Chaudoir, 1869

Plate 1 : Fig. 1. Plate 2 : Figs 1–2. Plate 4. Fig. 1.

Plate 1.

Type material (habitus in dorsal aspect, male). Scale bar equals 10 mm. 1 (holotype) 2 (holotype) 3 (holotype of Jeannel, 1940) 4 (holotype).

Plate 2.

Material studied (habitus in dorsal aspect, male). 1 (Djibouti: Obock) 2 (Oman: Dhofar, holotype of Häckel, Farkač & Sehnal, 2005) 3 (Kenya: Voi env.) 4 (Namibia: Okahandja). Scale bar equals 10 mm.

Chaudoir, 1843: 745 (type loc. “Kordofan”).

[Kollar in litt.] syn. sn.

Breuning, 1927: 185. Lapouge 1932: 414;

Jeannel, 1940: 128.

Häckel, Farkač & Sehnal, 2005: 2 (type loc. “Oman: Dhofar”), syn. sn.

Häckel, 2012: 57.

Type material.

1♂ labelled “HOLOTYPE / Ex Musaeo Chaudoir / (Chd.) P. Basilewsky vid. 1992 (cMNHN); 1♂ labelled “SW Asia, S Oman, Dzhopar Prov., Al Mughsayi vill. env., 0–50 m a.s.l., VIII.1999, lgt. S. Jákl / HOLOTYPE det. Häckel, Farkač & Sehnal, 2005 / Calsoma det. Häckel, Farkač & Sehnal, 2010” (cHAC).

Other material studied.

Djibouti. 1♂: “Obock” (cNMP).

Oman. 5♀: Dzhophar Prov., Takwa env., 50 m a.s.l. (cFAR, cHAC, cSEH); 1♀: rd. Al Mughsayi – Salalah, ca 3 km from Mughsayi, 20 m a.s.l. (cFAR); 1♀: Dhophar Province, Takwa env., 200 m a.s.l. (cKAL); 3♂, 2♀: Dzhofar prov., Wadi Nashib, 24 km E Salalah (cHAC, cSEH); 5♂: Dzhofar prov., Wadi Nashib, 20 km E Salalah (cHAC, cSEH).

SENEGAL (Niger or Chad probably). 1♀: ”Senegal” (cNMP).

Yemen. 1♂: NW Al Mukhallā: Kawr Saybān Mtn. (cHÄC).

Chad, Djibouti, Eritrea, Ethiopia, Kenya, Niger, Nigeria, Oman, Somalia, South Sudan, Tanzania, Yemen. Data from Burundi, Rwanda and Uganda need confirmation.

Breuning, 1927

Plate 1 : Fig. 2. Plate 2 : Fig. 3. Plate 3 : Fig. 3. Plate 4: Fig. 2.

Plate 3.

Aedeagi of (Namibia) and (Kenya) compared with aedeagus of “” (holotype). 1 (Namibia: Okahandja); a – aedeagus with expanded endophalus in right lateral view, b – the same in left lateral view, c – the same in anterior view 2 (holotype of Jeannel, 1940) aedeagus in left lateral view 3 (Kenya: Voi env.), a – aedeagus with expanded endophalus in right lateral view, b – the same in left lateral view, c – the same in anterior view.

Breuning, 1927: 185 (type loc. “Usambara”).

(partim) Jeannel, 1940: 128 (loc. “Diré-Daoua”);

(partim) Jeannel, 1940: 129 (loc. “Érythrée: Tessenei”).

var.

Culot, 1990: 9.

Häckel, 2012: 57.

Bruschi, 2013: 129.

Breuning, 1927. 1♂ labelled “Usambara”(cNBCL).

Kenya. 1♂, 1♀: E of Garsen, W of Witu (cSEH); 2♂, 1♀: S of Voi (cHAC); 1♂, 1♀: Taita prov. Sagala Hills, Voi env. (cHAC); 1♂: Tsavo East, Voi Lodge, (cWRA); 1♀: NE prov. El Wak (cHAC); 1♂, 1♀: Modo Gashi to Wajir (cHAC); 2♂, 2♀: Coast province, Garissa, N of Bura (cHAC, cSEH), 1♂: Amboseli National Park (cSEH); 2♀: Eastern 729, Sosoma, 202 km E of Thika (cSEH).

Sudan. 1♂, 1♀: Vad Medani (cSEH).

Geographic distribution.

Chad, Kenya, Somalia, Sudan, Tanzania.

Dejean, 1831

Plate 4: Fig. 5

Dejean, 1831: 562 (type loc. “Sénegal”).

Klug, 1853: 247 (type loc. “Téte”).

Motschulsky, 1865: 306.

Breuning, 1927: 187. Lapouge 1932: 415.

Jeannel, 1940: 129.

Häckel, 2012: 58, 64; 2013: 31.

Botswana. 1♂, 1♀: Ngamiland district, ne. of Maun, Tamalakane (cHAC).

Ethiopia. 1♂, 1♀: Gambela region, Gambela env., 400 m (cHAC).

Ghana. 1♂, 1♀: Northern Prov., West Gonja district, Damongo env. (cHAC).

Kenya. 1♂, 1♀: Coast Prov., Taita-Taveta Co., s. of Voi (cHAC).

Madagascar. 1♂, 2♀: Toliara prov., Ampanihy district, Ejeda env. (c FAR, cHAC); 1♂, 1♀: Mahajanga Prov., Ampatika env., Mahajamba river (cHAC).

Namibia. 1♂: Caprivi reg., Bagani- Popa Falls; 1♀: Khomas region, 40 km e. Windhoek (airport) (cHAC).

Senegal. 1♂, 1♀: Thiès region, M’bour department, Saly Portudal (cHAC).

Tanzania. 1♂: Arusha reg., Mto Wa Mbu env. (cHAC); 1♀: Morogoro region, Mikumi (cHAC). Zimbabwe. 1♂: 20 km NE Shamva, Nyagui river (cHAC).

Angola, Benin, Botswana, Burkina Faso, Burundi, Cameroon, Cabo Verde Islands, Chad, Congo, Côte ďIvoire, Democratic Congo, Eritrea, Ethiopia, Gabon, Gambia, Ghana, Guinea, Guinea-Bissau, Guinea Equatorial, Kenya, Lesotho, LiPageBreakberia, Madagascar, Malawi, Mali, Mauritania, Mozambique, Namibia, Niger, Nigeria, Republic of Central Africa, Republic of South Africa, Rwanda, Senegal, Sierra Leone, Somalia, Swaziland, Tanzania, Togo, Uganda, Zambia, Zimbabwe.

Breuning, 1934

Plate 4: Fig. 3.

Breuning, 1934: 38 (type loc. “Masaua”).

Jeannel, 1940: 130.

Häckel, 2013: 31.

Kenya. 8♂, 6♀: North-Eastern Prov., El Wak. (cFAR, cHAC, cSEH); 2♂, 4♀: Eastern Prov. Marsabit to South Horq (cFAR, cHAC, cSEH).

Eritrea, Ethiopia, Kenya, Somalia.

species group (sensu Häckel 2013)

subgroup

Klug, 1832

Klug, 1832: pl. IX. (type loc. “Cap Vert”).

Motschulsky, 1865: 304.

Mandl, 1953: 57.

m.

Mandl, 1970: 61.

Kenya. 1♂, 1♀: Marsabith to South Orr (cHAC).

Oman. 8♂, 8♀: Wadi Qitbit, 150m (cFAR, cHAC, cSEH).

Senegal. 1♀: Senegal (cHAC).

Sudan. 1♂: Port Sudan (cHAC).

Algeria, Cabo Verde Islands, Canarian Islands, Chad, Djibuti, Egypt, Eritrea, Ethiopia, western India, Iran, Iraq, northern Kenya, Kuwait, Libya, Mali, Niger, Oman, Pakistan, Saudi Arabia, Senegal, Somalia, Sudan, United Arab Emirates, Yemen.

Chaudoir, 1852

Plate 1 : Figs 3–4. Plate 2 : Fig. 4. Plate 3 : Figs 1–2. Plate 4. Fig. 6.

Chaudoir, 1852: 99 (type loc. “Cap de bonne-Espérance”).

Breuning, 1927: 221.

Lapouge, 1932: 410.

Jeannel, 1940: 127 (type loc. “Azbin, à 20 km. ďAgadès, dans ľAïr”),

Häckel, 2013: 24.

1♂ labelled “LECTOTYPE / Ex Musaeo Chaudoir / Lectotype Chaud. 1852 Th. Deuve det,. 1978” (cMNHN); 1♂ labelled PageBreak“MUSEUM PARIS AZBIN (AIR) REG. de TINTABORAC 20 K E ď AGADÈS CAPde POSTH 1908 / HOLOTYPE / n. sp. Jeannel det.” (cMNHN).

Kenya. 1♂, 1♀: Amboseli National Park (cSEH).

Namibia. 2♂, 2♀: Omaruru (cHAC); 5♂, 3♀: Okahandja, Gross Bamen (cHAC, cSEH); 1♀: Otjivarongo (cSEH); Gobabis-Aranos (cSEH); 1♂: Maltahohe (cSEH).

Republic of South Africa. 1♂, 3♀: Northern Cape Province, SW Kimberley, 13 km SW Ritchie (cHAC).

Southern Kenya, Namibia, Tanzania, Republic of South Africa.

Comments on classification

Our study shows the following:

1. Termination (apex) of the aedeagus. We have found no difference in shape of the apex among species or subspecies within the same group, the shape is distinct only among species belonging to different groups. The apex of (narrower PageBreakand sharper, Plate 3: Figs 1–2. Plate 4: Fig. 6) differs from those in all species of the group (more blunt apex, Plate 3: Fig. 3. Plate 4: Figs 1–5). The apex in Jeannel’s holotype of corresponds to that in the group (Plate 3), which does not support the opinion of Bruschi (2013) that is a synonym of .

2. Shape of inflated endophalus. We have not found an apparent difference either among species or subspecies belonging to the same group, or among species belonging to different groups (Plate 4: Figs 1–5). This result does not support the opinion that shape of the endophalus can be used to indicate assignments to to species or species groups.

3. Morphometric parameters.

A. . In three measured holotypes (or lectotypes) are the values within the minimum and maximum intervals found in corresponding populations and sexes, whereas in the male holotype of the value is outside of the interval. The TL value in the holotype is closest to the values found in males of , and lies within the interval found in females of that subspecies (all types are males, see Table 1a, Table 2). This fact supports our opinion that the holotype of is an extremely large male of .

Table 1a.

Total length, intervals.

Taxon Holotype (HT), lectotype (LT)	Total length including mandibles in millimeters (TL). Interval of minimum and maximum value of TL measured in 10 specimens of the same sex is in parentheses. Yes (Y) – TL value of type is within interval. No (N) – TL value of type is outside of interval.
Calosoma scabrosum scabrosum Chaudoir, 1843 (HT ♂)	25.5 (23.1–26.2 ♂♂)	Y
Calosoma scabrosum roeschkei Breuning, 1927 (HT ♂)	25.0 (22.0–28.5 ♂♂)	Y
Calosoma imbricatum hottentotum Chaudoir, 1852 (♂ LT Deuve 1978)	21.0 (17.5–21.3 ♂♂)	Y
Calosoma aethiops (Jeannel, 1940) (HT ♂)	21.5 (Calosoma imbricatum hottentotum ♂♂ 17.5–21.3 ♀♀ 19.2–23.0)	N

Table 2.

Intervals of all measurements in each taxon (intervals), in TL and LE/WE for both sexes separately and total.

Calosoma scabrosum scabrosum	23.1–26.2	23.1–26.2	23.4–25.7	1.35–1.63	1.33–1.67	1.38–1.56	1.44–1.56	1.38–1.53
Calosoma scabrosum roeschkei	22.0–28.5	22.0–28.5	24.1–27.9	1.29–1.65	1.38–1.77	1.33–1.59	1.39–1.57	1.33–1.59
Calosoma senegalense	23.0–30.0	23.0–29.3	24.9–30.0	1.22–1.62	1.30–1.73	1.45–1.56	1.45–1.55	1.45–1.56
Ctenosta strandi	25.2–30.3	25.2–29.0	26.4–30.3	1.32–1.44	1.46–1.62	1.44–1.64	1.50–1.64	1.44–1.64
Calosoma imbricatum imbricatum	17.0–22.0	17.0–20.4	18.2–22.0	1.33–1.43	1.41–1.68	1.26–1.57	1.39–1.53	1.26–1.57
Calosoma imbricatum hottentotum	17.5–23.0	17.5–21.3	19.2–23.0	1.33–1.54	1.43–1.84	1.33–1.57	1.33–1.50	1.38–1.57

Total length including mandibles

B. . The WP/WH in two measured holotypes (or lectotypes) is within the minimum and maximum intervals found in the pertinent populations and sexes. In the third taxon the WP/WH value of the holotype is outside of the interval in both sexes. In the holotype of is the WP/WH value within the interval found in the corresponding sex (males) of and also within the interval found in females. Overall the WP/WH values found in the measured taxa is very variable in both species and sexes (Table 1b, Table 2), and in our opinion thus cannot be used as a criterion in species-level taxonomy.

Table 1b.

Pronotal width to head width ratio, intervals.

Taxon Holotype (HT), lectotype (LT)	Maximum pronotal width to maximum head width inc. eyes ratio (WP/WH). Interval of minimum and maximum value of WP/WH measured in 10 specimens of the same sex is in parentheses. (Y/Y) – value of type is within interval. (N/N) – value of type is outside of interval. (N/Y) – value of type is outside of interval in males but within interval in females.
Calosoma scabrosum scabrosum Chaudoir, 1843 (HT ♂)	1.50 (1.34–1.44 ♂♂) (1.35–1.48 ♀♀)	N/N
Calosoma scabrosum roeschkei Breuning, 1927 (HT ♂)	1.50 (1.29–1.63 ♂♂) (1.41–1.65 ♀♀)	Y/Y
Calosoma imbricatum hottentotum Chaudoir, 1852 (♂ LT Deuve 1978)	1.45 (1.33–1.50 ♂♂) (1.23–1.54 ♀♀)	Y/Y
Calosoma aethiops (Jeannel, 1940) (HT ♂)	1.32 (Calosoma imbricatum hottentotum) (Calosoma scabrosum scabrosum) (Calosoma scabrosum roeschkei)	N/Y N/N Y/N

Maximum pronotal width to maximum head width including eyes ratio

. The WP/LP value in two measured holotypes (or lectotypes) is with exception of males of within the minimum and maximum intervals found in the pertinent populations and sexes. In the holotype of is the WP/WH value within the interval found in all compared species of both sexes. Overall the WP/LP values found in the measured taxa are quite non-specific in both species and sexes (Table 1c, Table 2), and in our opinion thus cannot be used as a criterion in species-level taxonomy.

Table 1c.

Pronotal width to length ratio, intervals.

Taxon Holotype (HT), lectotype (LT)	Maximum pronotal width to maximum pronotal length ratio (WP/LP). Interval of minimum and maximum value of WP/LP measured in 10 specimens of the same sex is in parentheses. (Y/Y) – value of type is within interval. (N/N) – value of type is outside of interval. (N/Y) – value of type is outside of interval in males but within interval in females.
Calosoma scabrosum scabrosum Chaudoir, 1843 (HT ♂)	1.65 (1.33–1.56 ♂♂) (1.52–1.67 ♀♀)	N/Y
Calosoma scabrosum roeschkei Breuning, 1927 (HT ♂)	1.67 (1.38–1.77 ♂♂) (1.47–1.74 ♀♀)	Y/Y
Calosoma imbricatum hottentotum Chaudoir, 1852 (♂ LT Deuve 1978)	1.74 (1.43–1.74 ♂♂) (1.55–1.84 ♀♀)	Y/Y
Calosoma aethiops (Jeannel, 1940) (HT ♂)	1.55 (Calosoma imbricatum hottentotum) (Calosoma scabrosum scabrosum) (Calosoma scabrosum roeschkei)	Y/Y Y/Y Y/Y

Maximum pronotal width to its maximum length ratio

D. . The WP/LP value in two measured holotypes (or lectotypes) is in both subspecies of within the minimum and maximum intervals found in the pertinent populations and sexes. In both sexes of the value is outside the interval. In the holotype of the value is within the interval found in both subspecies of . Overall the WP/LP values found in the measured taxa can be regarded as variable, namely in . In our opinion they cannot be used as a criterion in species-level taxonomy (Table 1d, Table 2).

Table 1d.

Elytral length to width ratio, intervals.

Taxon Holotype (HT), lectotype (LT)	Maximum elytral length to maximum elytral width ratio (LE/WE). Interval of minimum and maximum value of LE/WE measured in 10 specimens of the same sex is in parentheses. (Y/Y) – value of type is within interval. (N/N) – value of type is outside of interval. (N/Y) – value of type is outside of interval in males but within interval in females.
Calosoma scabrosum scabrosum Chaudoir, 1843 (HT ♂)	1.44 (1.44–1.56 ♂♂) (1.38–1.53 ♀♀)	Y/Y
Calosoma scabrosum roeschkei Breuning, 1927 (HT ♂)	1.52 (1.39–1.57 ♂♂) (1.33–1.59 ♀♀)	Y/Y
Calosoma imbricatum hottentotum Chaudoir, 1852 (♂ LT Deuve 1978)	1.54 (1.33–1.50 ♂♂) (1.38–1.57 ♀♀)	N/Y
Calosoma aethiops (Jeannel, 1940) (HT ♂)	1.39 (Calosoma imbricatum hottentotum) (Calosoma scabrosum scabrosum) (Calosoma scabrosum roeschkei)	Y/Y N/Y Y/Y

Maximum elytral length to its maximum width ratio

The above data lead us to conclude that there are no convincing morphological differences between and . The only exception may possibly be the somewhat higher WP/LP ratio (Table 2) and lighter coppery coloration in most specimens of . Since specimens of both taxa have never been found together, we assume that they belong to allopatric populations of one species that have yet to reach the state of full speciation. Therefore, we lower the status of (sensu Bruchi 2013) to a subspecies of . occupies mainly the southern part of distribution of the species. The present data show the north – south distribution of both subspecies to have a virtually disjunct character (Map 1). But it is in our opinion important to realize that in no area have the described subspecies been found to occur together. It is therefore likely that the northern Sudan – Ethiopia borderland will continue to produce . The describer (Breuning 1927: 186) of wrote: [“This form () is due to its more robust head, less bulging eyes, wider, toward base more right-angled pronotum with more rounded hind angles, shallower basal pits, somewhat flatter, at shoulders broader and terminally more abruptly slanted elytra and dark to brownish bronze dorsal coloration with light brown, coppery-rimmed margins and foveae in primary intervals so PageBreakconspicuous that I originally intended to describe it as a separate species. However, some individuals of this form () are clearly transitional to the typical form (), with coloration remaining as the only constant character. Differences (of separate populations) from the nominotypical form show step-like transitions, for which reason I presently regard as a subspecies of ” (original in German)].

Map 1.

Geographic distribution of the and species subgroups (sensu Häckel 2012). T – the type locality. Dark blue discs – s. str. Chaudoir, 1843. Light blue discs – Breuning, 1927. Green discs – Breuning, 1934.

Jeannel (1940: 128) did not see Breuning’s subspecies but treated it as a variety and remarked: [“Breuning’s variety rather appears to be another (separate) species closely related to ]”; he did not further comment on the taxon and placed specimens corresponding to Breuning’s description among African populations of “” (Jeannel 1940: 128). In this connection, Jeannel and subsequent authors such as Rougemont (1976: 247) and Vigna Taglianti and Bruschi (1986: 21) solved taxonomic uncertainties by incorrect determination of specimens from eastern PageBreakAfrica as Chaudoir, 1869 (Häckel 2012: 57.). The holotype of is from Bengal (today either Bangladesh or West Bengal in India) and other specimens of the type series are from Coimbatore in the vicinity of Madras (today Tamilnadu State in southern India). The name was therefore synonymized with (Jeannel 1940: 128).

Jeannel nevertheless realized that African populations identified as (=) most likely belong to another species, coined for them a new taxon, Jeannel, 1940, and included in the distribution of this taxon also populations corresponding to Breuning’s (“Diré Daoua”, Jeannel 1940: 128). The only exception was a population from Eritrea (“Tessenei” 1940: 129), which he continued regarding as . In the description of Jeannel (1940: 127–128) wrote: [“if we regard with its gular and labial setae as belonging to the genus , we can see that in reality it is a transitional species combining characters of and , an important species attesting to its assignment to the genus , which differs by reduction of the said setae and the type of sculpture placing it near the - lineage. The male mesotibiae have the setal brush prolonged as in and similar to that present in . The ventral side of the fourth male protarsomere is smooth]”.

Evident from Jeannel’s text are the difficulties he had in placing the new species in his system and in defining the “genera”. More recently some authors (Culot 1990, Bousquet et al. 2003, Bruschi 2013) regarded the genera as subgenera, but in our opinion Jeannel’s criteria do not allow to distinguish them. For instance Bruschi in the key does not adhere to Jeannel’s criteria (setae, elytral microsculpture) and originally separates from on the hind pronotal angles. In the key he (2013: 29) states: “13(14) always perceptible hind angles of pronotum – /; 14(13) very small and pointed hind angles of pronotum, in some cases quite obliterated”. In our opinion Jeannel’s genera are not valid, which is supported also by their discord with results of DNA analyses. With only one exception, all the subgenera were synonymized with the genus s. str. (Häckel 2012: 56, 2013: 12). Jeannel’s difficulties in separating the “genera” and (according to us two close species groups of the genus ) reflect the confused composition of the type series of his . Without examination of Jeannel’s holotype of , most subsequent authors regarded all African populations similar to , with golden bronze coloration, as (Vigna Taglianti and Bruschi 1986: 21, Culot 1990: 9, Lorenz 2005: 68).

Only Bruschi, first on the internet and later in print (2013: Plate 17: Figs 8, 9), published photos of the holotypes of (Tanzania: Usambara) and (Niger: Azbin). The holotype of (Usambara, see Map 1) clearly is a species belonging to the group, and we concur with Bruschi that in compliance with the priority principle the taxon must be ascribed to Breuning (the name has priority over , if the two are the same species).

However, in our opinion the specimen from Azbin (Jeannel’s holotype of ) looks different and does not belong to the group (=Jeannel’s genus PageBreak). Our comparisons of types and their aedeagi show that the holotype of corresponds in shape, size and sculpture of the elytra, shape of the legs, and termination of the aedeagus to Chaudoir, 1852. It belongs to another group ( group, subgroup sensu Häckel, 2013: =Jeannel’s genus ), which partially overlaps the distribution of the group. Our conclusions are based chiefly on the different aedeagal morphology unequivocally shown by the photos (Plate 3, Figs 1, 2 versus Fig. 3). Futher documentation of morphological characters is not needed.

sensu lato is by a number of authors understood as a species with an extremely wide distribution reaching from Canary and Cape Verde Islands through the African Sahel belt, subsaharan Africa, Arabia, Iran and Pakistan to India and Bagladesh (Breuning 1927: 221–223, Jeannel 1940: 104–106, Mandl 1970: 61–63, see Map 2). The cited authors identify southern African populations of as the subspecies , whose holotype comes from the Cape province (Chaudoir 1852: 99). is usually regarded as cofined to southern Africa, and Mandl (1970: 61, 63) even named another subspecies, , for populations from eastern Africa (northern Kenya, Somalia), which is a transitional form differing from the northern (Sahel-Arabian) nominotypical form () by wider pronotum. Populations from southern Kenya and northern Tanzania with more lighter coppery coloration approaching rather the southern African populations (). Jeannel (1940: 105) is the only author who regarded also the Kenya population as . Mandl (1970: 63) commented these occurrences as “Wahrnscheinlich gehören die folgende Orte zu dieser Subspecies []: Kenya Sultan Hamid zwischen Voi und Nairobi, die Jeannel für angibt”, and Bruschi (2013) countered: „It seems that, contrary to the opinions of Mandl (1970: 61) that attributed this citation to his C , Jeannel is right since in south western Kenya C is really present“. In our opinion it is evident that the northern limit of is vague and hosts a number of forms transitional to the nominotypical . It is therefore possible that Jeannel’s type of from Azbin (today northern Niger) belongs to one of the transitional populations and the locality is in fact correct (see Map 2). In our opinion Jeannel‘s type series of contains the holotype that we regard as (most likely the subspecies ) and specimens from other populations belonging to some of the subspecies of , mostly to .

Map 2.

Geographic distribution of the species subgroup (sensu Häckel 2013) compared to Mandl‘s data (1970) and Chaudoir, 1852. T – the type locality, ● (full disc) – recent records, ⚬ (empty disc) – Mandl‘s data 1–4 s. str. Klug, 1832 5–7 Chaudoir, 1852 1 s. str. (sensu Mandl 1970) 2 (Motschulsky, 1865) sensu Mandl 1970 3 Mandl, 1953 sensu Mandl 1970 4 Mandl, 1970 sensu Mandl 1970 5 (holotype of Jeannel, 1940) 6 ( sensu Mandl 1970) 7 . sensu Mandl 1970.

Comparison of the types of and in our opinion also confirms the original Breuning’s idea of one species with two terminal forms and a number of transitional forms between them (Plate 1: Figs 1–2). Another taxonomical inaccuracy was caused by Häckel et al. (2005: 2), who believed that populations newly discovered on the Arabian peninsula (Oman, Yemen, see Map 1) represent a different species (, Plate 2. Fig. 2). Eventually, after comparison with the type of it became clear that the Arabian specimens agree with the type (Häckel et al. 2010: 11), and today they are included in the distribution of the nominotypical subspecies s. str. (Müller 1977, Häckel 2012: 57).

Below we present morphometric tables comparing populations of (hitherto labeled as ) with specimens of the Horn of Africa (Djibouti) and the Arabian peninsula (Oman, Yemen). The tables also compare the noted populations of with specimens of from southern and eastern Africa and from Afrotropical Region. In this connection we consider it important that no known locality has produced sympatrically living and (see Map 1). However, at at least one locality (Kenya: North-Eastern Province, El Wak, see Map 1) has produced (ssp. ) together with (ssp. ) and another species of the group ( Breuning). PageBreakOur conclusion therefore is that Breuning’s (1928: 185) original idea is valid, and consequently we demote sensu Bruschi (2013: 129) to a subspecies of .

The subgeneric placement of , in the subgenus follows the recently proposed classification supported by results of DNA analyses (Su et al. 2005, Häckel 2012, 2013).