Mesobuthus eupeus (C. L. Koch, 1839)

Kovařík, František, Fet, Victor, Gantenbein, Benjamin, Graham, Matthew R., Aydin, Ersen, Yağmur, Šťáhlavský, František, Nikita, Poverennyi & Novruzov, Nizami E., 2022, A revision of the genus Mesobuthus Vachon 1950 with a description of 14 new species (Scorpiones Buthidae), Euscorpius 348, pp. 1-189 : 33-41

Mesobuthus eupeus (C. L. Koch, 1839) , s. str.

( Figures 196–243 , 1155–1156, 1159–1161, 1167, Tables 3, 15) http: //zoobank. org/urn: lsid: zoobank. org: act: 0FB17935-

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Androctonus eupeus C. L. Koch, 1839a: 127–128 , pl. CLXXV, fig. 419.

TYPE LOCALITY AND TYPE REPOSITORY. “Caucasus”; types lost. Neotype (designated here): Georgia, Tbilisi (41.72°N 44.79°E); NMPC.

SYNONYMS:

= Androctonus ornatus Nordmann, 1840: 732 , pl. I, fig. 2 (synonymized by Birula, 1896: 238).

http: //zoobank. org/urn: lsid: zoobank. org: act: 2C6F7E6C-

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( TYPE LOCALITY AND TYPE REPOSITORY: Georgia, Imereti Region [no exact locality; estimated at Kutaisi (42.25°N 42.70°E)], MZHF.

= Buthus cognatus L. Koch, 1878: 58 , pl. I, fig. I (synonymized by Birula, 1896: 238).

http: //zoobank. org/urn: lsid: zoobank. org: act: 88C80B32-

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( TYPE LOCALITY AND TYPE REPOSITORY: Georgia, Samtskhe- Javakheti Region, upper Kura River, environs of Atskuri ( Atskur ) (41.73°N 43.16°E), type lost.

REFERENCES (selected):

Androctonus ornatus: Kessler, 1874: 19 (in part).

Buthus eupeus forma α: Birula, 1896: 239.

Buthus eupeus forma β: Birula, 1896: 240 (in part).

Buthus eupeus: Birula, 1900b: 3 .

Buthus eupeus eupeus (in part): Birula, 1904b: 37; Birula, 1905a: 122, 124–125, fig. 1; Birula, 1905b: 122 (in part); Birula, 1911b: 162–164; Birula, 1917a: 24–39, pl. I; Birula, 1918: 9–10; Birula, 1925: 95.

Buthus (Buthus) eupeus: Birula, 1912: 118–121 .

Buthus eupeus eupeus natio eupeus: Birula, 1917a: 34 .

Buthus eupeus eupeus natio armeniaca: Birula, 1917a: 34.

Mesobuthus eupeus: Vachon, 1950: 152 (in part); Vachon, 1952: 325(in part); Fet & Lowe, 2000: 171–172 (in part; complete references list until 1998); Crucitti & Cicuzza, 2001a: 3–11, figs. 1–6; Crucitti & Cicuzza, 2001b: 231- 232, fig. 5 (in part); Teruel, 2002: 75–76, figs. 1–3 (in part); KarataŞ & KarataŞ, 2003: 1–4, figs. 1–2 (in part); Yağmur et al., 2007: 97–98 (in part); Kaltsas et al., 2008: 219 (in part); Mirshamsi et al., 2010: 2867 (in part); Mirshamsi et al., 2011b: 20 (in part); Gharakhloo et al., 2018: 875 (in part); KovařÍk, 2019: 17, figs. 134–137, 148, 161–163, 171–172.

Mesobuthus eupeus eupeus: Vachon, 1958: 155 (“forme typique”); Fet, 1989: 86–91 (in part; complete references list for the former USSR); Fet, 1994: 527; Fet & Lowe, 2000: 171–172 (in part; complete references list until 1998); Mirshamsi et al., 2010: 2867 (in part); Mirshamsi et al., 2011a: 8, fig. 4 (in part); Yağmur et al., 2012: 16–17, figs. 1–3; Moradi et al, 2015: 12–13 (in part); Navidpour et al., 2019: 4–5 , fig. 55 (in part).

TYPE MATERIAL EXAMINED. Georgia, Tbilisi (41.72°N 44.79°E), 2012, 1♀ (neotype of Androctonus eupeus , designated here) ( Figs. 204–205 , 225–235 , 237, 239 ) , NMPC; Georgia , Imereti Region [no exact locality; estimated at Kutaisi (42.25°N 42.70°E)], 1♀ ( Figs. 196–201 ), leg. A. Nordmann , MZHF (holotype of Androctonus ornatus ) .

OTHER MATERIAL EXAMINED. Armenia, road from Solak to Sevan , 17 June 1981, 3♀, leg. Soukup , FKCP; Little Caucasus Mts., Goght near Geghard (40.15°N 44.79°E), 35 km SE of Yerevan, Azat River valley , 12 June 1988, 2♂ 2♀ 1juv., leg. Z. Jindra , FKCP; Lusashog (39.85°N 44.96°E), 1850 m a. s. l., 21 May 2005, 1juv., leg. S. Snäll , NMPC; Gorovan sands, 39°53'N 44°44'E (39.88°N 44.73°E), 970 m a. s. l., 23 May 2007, 1♂ 2♀ 4juvs., leg. S. Snäll , NMPC; Gorovan, Vedi env., 39°53'46.33"N 44°37'57.63"E (39.53°N 44.37°E), 929 m a. s. l., 17–20 June 2013, 1♀ 1juv. (No. 460), leg. J. RolčÍk , NMPC; Syunik, Meghri , 38°53'805"N 046°15'468"E (38.88°N 46.25°E), 648 m a. s. l., 6♂ ( Figs. 202–203 , 206, 211 – 213214–224, 236, 238, 240–243, Table 3, Nos. 1476, 1477, 1478, 1479) 3♀, 9–10 July 2018, leg. D. Hoferek , NMPC; Tavush, Dilijan, Gosch , 40°49'946"N 044°59'781"E (40.82°N 44.98°E), 1320 m a. s. l., 13 July 2018, 1♀, leg. D. Hoferek , NMPC. Georgia, 1♂ 1juv. (No. 16) , NMPC; Dzhvari (42.71°N 42.03°E), 15 km from Tbilisi, 1juv., 7 August 1988, leg . T. Peš , FKCP; Tbilisi, 1♂, 7 June 1959, leg. J. Niedl , FKCP; 2005, 1♀1♀ im ., NMPC; Vashlovani National Park , 41°13.724'N 046°21.195'E (41.22°N 46.35°E), 322 m a. s. l., 3 June 2019, 2juvs., leg. W. Grosser , NMPC; Karsani env., 41°49.813'N 044°42.109'E (41.82°N 44.70°E), 708 m a. s. l., 25 May 2019, 1♀, leg. W. Grosser , NMPC; Tabakhmela near Tbilisi, 12 June 2013, 1♀ 2juvs., leg. D. Kasatkin , AZMM; Tbilisi, Mt. Mtatsminda , northern slope, 41°41'47.54"N 44°46'46.83"E (41.68°N 44.77°E), 13 June 2013, 1♂ 1juv., leg. D. Kasatkin & I. Shokhin , AZMM. Azerbaijan, Ağstafa District (41.12°N 45.41°E), Poylu Village , 8 June 2013, 1♂, leg. D. Kasatkin , AZMM. Turkey, Ağrı Province, Doğubayazit, Gürbulak Village , 39°25'09"N 44°18'34"E (39.42°N 44.30°E), 1620 m a. s. l., 22 July 2010, 1♂ 1♀, leg. E. A. Yağmur & H. Koç , AZMM; Artvin Province, Yusufeli District , 40°49'N 41°32'E (40.82°N 41.53°E), 740 m a. s. l., 1 July 2008, 1♂ 1♀, leg. A. Avcı , NMPC; Erzurum Province, Horasan, Saç Geçidi, 35 km NWW Eleskirt, SE Horasan (39.80°N 42.66°E), 2315 m a. s. l., 9 May 2002, 1♀ 1juv., leg. P. Kabátek , NMPC; Iğdır Province, 5 km SE Tuzluca (near the border with Armenia ) (40.04°N 43.64°E), 28 May 2005, 1♂ 4♀ 1juv., leg . V. GvoždÍk , NMPC; Aralık District, Yenidoğan Village , 39°47'12.3"N 44°23'36.9"E (39.78°N 44.38°E), 1563 m a. s. l., 16 June 2007, 1♂ 1♀, leg. H. Koç , NMPC; Kars Province, Kars town, Kars Castle 40°37'01.7"N 43°05'28"E (40.61°N 43.09°E), August 2014, 1♂ 1♀ (Nos. 362, 363), leg Ç. Altin , AZMM; Karakurt , 20 km W of Karakurt town (40.17°N 42.59°E), ca 1500 m a. s. l., 14 May 1996, 1♀ 1juv., leg. J. Pitulová , NMPC; 1juv., leg. D. Král , NMPC; Central region, Ani ruins, 40°31'N 43°34'E (40.52°N 43.57°E), 1500 m a. s. l., 20 May 1997, 2♂ 1juv., leg. M. Kaftan , NMPC; Van Province, BaŞkale, Çamlık Village (37.95°N 44.08°E), 1889 m a. s. l., 22 July 2007, 1♂, leg. H. Koç & A. V. Gromov .

DNA DATA. Mesobuthus eupeus (s.str.) was represented in our survey by five DNA sequences from Armenia and Turkey ( Table 16); no DNA markers from Georgia and Azerbaijan are yet available. The same species was also represented, in our opinion, among the populations studied by Mirshamsi et al. (2010) in their DNA survey of Iranian Mesobuthus (two specimens from the West Azerbaijan Province). We did not examine any specimens of M. eupeus , s. str., from Iran. Another species in the dataset of Mirshamsi et al. (2010) from West Azerbaijan, in our opinion, is M. persicus (see below) ( Table 16).

DIAGNOSIS. Total length of adults 39 mm (male) to 60 mm (female). Trichobothrium db on fixed finger of pedipalp situated between trichobothria est and esb, near to est or with level with est. Male with fingers proximally more twisted than in female. Pedipalp chela length/width ratio 3.2–3.5 in male and 3.7–3.9 in female. Pectinal teeth number 21–25 in male, 18–21 in female. Chelicerae yellow, without reticulation. Pedipalps and metasoma very sparsely hirsute. Carapace and tergites reddish brown, black pigmented, dark spots and stripes usually present; metasoma, telson, pedipalps and legs reddish brown, only part of metasomal segment V black; metasomal ventral carinae usually dark colored. Femur of pedipalp with 4–5 granulated carinae. Patella with 8 granulated or smooth carinae. Chela lacks carinae. Movable fingers of pedipalps with 11–12 cutting rows of denticles and 5 terminal denticles. Central lateral and posterior lateral carinae of carapace not joined to form a continuous linear series of granules to posterior margin. Sternite VII smooth, with 4 well marked smooth or granulated carinae. Metasomal segment I with 10 carinae; segments II to IV with 8 carinae, other two carinae are indicated by incomplete row of denticles on metasomal segments II and III; segment V with 5 carinae. Metasoma IV with lateromedian carina smooth to absent in male but present and usually granulated in female, all other carinae granulated. Intercarinal surfaces on metasoma I laterally usually smooth. Length to depth ratio of metasoma III 1.2–1.35; metasoma IV 1.45–1.70 in both sexes. Telotarsus III ventral setation represented by short and strong spiniform setae. Tarsi hirsute, in adults with 7–10 retroinferior macrosetae on basitarsus III. Tibial spur on both legs III and IV moderate. Telson elongated. Anal lobe divided into three parts. Tergites III–IV with three straight, parallel carinae.

HISTORY OF STUDY. A renowned German arachnologist Carl L. Koch (1839a; not 1838 as often cited) was the first to describe a species of the future Mesobuthus , Androctonus eupeus , which also became the type species of this genus. The single type specimen, possibly a female, came from the Nuremberg collection of Jacob Sturm (1771–1848), a famed engraver and insect collector. The provenance of the type was listed by C. L. Koch (1839a) only as “Caucasus”, and the type is long lost (Birula, 1917a: 35).

Next year, Alexander von Nordmann (1840), a Swedish zoologist who worked in Helsingfors (now Helsinki , Finland, then in the Russian Empire) published his extremely brief, and largely unreliable, description of Androctonus ornatu s from the Imereti Region of Georgia, based on a single female collected in 1836. The holotype still exists in the University of Helsinki Zoological Museum , Finland ( MZHF), and was examined by us through a kind loan by Pedro Cardoso ( Figs. 196–201 ) .

The first review of scorpions of the Russian Empire was published in 1874 by the famous German-Russian zoologist Karl Kessler (1815–1881). Kessler (1874) used the name Androctonus ornatus for a common Caucasian buthid scorpion but conflated under it two independent and very different species, now called Mesobuthus eupeus and Olivierus caucasicus ( Nordmann, 1840) . The name Androctonus ornatus has been in use by Russian zoologists for years until Birula (1896) corrected it to its senior synonym, then Buthus eupeus . Since it was published in Russian language, Kessler’s review remained largely unknown to the European zoologists.

In 1878, Carl Koch’s son Ludwig Koch (1825–1908), also a famed German arachnologist, published a description of Buthus cognatus from Georgia ( Koch, 1878: 58, Taf. I, Fig. 7 ). This specimen was collected in Atskhur (Imereti) on the upper Kura River in 1875 by the German geographer Oscar Schneider (1841–1903) during his six-month expedition, which provided rich zoological materials, mainly from the Caucasus but also from Transcaspia (modern Turkmenistan). Koch noted (p. 59) that this new species was close to Buthus eupeus , described also from the Caucasus by his father. The species Androctonus ornatus was not mentioned by L. Koch (1878) who therefore likely was not aware of the publications by Nordmann (1840) or Kessler (1874). Birula (1905a: 122; 1917a: 37) examined the type of Buthus cognatus and deemed it identical with the “typical race” of Buthus eupeus eupeus .

Birula (1896) was the first to synonymize both Androctonus ornatus Nordmann, 1840 and Buthus cognatus L. Koch, 1878 with Buthus eupeus (C. L. Koch, 1839) . He also started a longterm tradition of establishing subspecies within this ‘umbrella’ species. In his definitive review of the Caucasian scorpiofauna, Birula (1917a: 24–39) provided a detailed redescription and discussion of Mesobuthus eupeus (then Buthus eupeus ) from the Caucasus, based on a nominotypical subspecies. Birula (1917a: 35) considered Tiflis (now Tbilisi, Georgia) a “typical locality” for this species; however, this designation remained informal as no neotype was ever fixed. In Birula’s view, this taxon was an extremely widespread (from Turkey to China, with the highest diversity in Iran) “polymorphic species” forming more than a dozen of allopatric subspecies.

This opinion was accepted by all scorpiologists for the next 100 years (Vachon, 1958; Fet, 1989, 1994; Fet & Lowe, 2000). Recently, Mirshamsi et al. (2011a: 15) split all Iranian Mesobuthus eupeus into two species, elevating M. phillipsii to the species rank. The splitting trend was continued by KovařÍk (2019) who recognized 12 valid species of Mesobuthus .

COMMENTS. Here, we limit M. eupeus s. str. to populations from the Caucasus. See Fet (1989) for a detailed list of localities from the former USSR (modern boundaries of Armenia, Georgia, and Azerbaijan).

Two local “races” ("Natio”) (armeniaca and talyschensis) from the modern Armenia and Azerbaijan, respectively, were described as infrasubspecific names of the subspecies Buthus eupeus eupeus by Birula (1917a: 34); these names are not available according to the ICZN and do not enter into synonymy.

Presence of M. eupeus within Russia (in the North Caucasus) was noted by Fet (1989: 90) only in North Ossetia – Alania (Ordzhonikidze, now Vladikavkaz). Several other new localities are reported north of the Georgian border in North Ossetia – Alania (K. Lotiev, pers. comm). A detailed further study of these populations is needed.

We restrict M. eupeus from Turkey to Ağrı, Artvin, Erzurum, Iğdır, Kars, and Van Provinces. We also describe three new species of Mesobuthus from Turkey: M. rahsenae sp. n., M. turcicus sp. n., and M. yagmuri sp. n. (see below) within the geographic range earlier reported for M. eupeus . In addition, we confirm a wide presence of M. mesopotamicus ( Penther, 1912) , stat. n. in southeastern Turkey; these populations were previously identified as M. eupeus phillipsii or M. phillipsii .

DISTRIBUTION. Armenia, Azerbaijan, Georgia, Iran (West Azerbaijan Province), Russia, Turkey (Figs. 1155–1156).

Figures

Figures 196–201. Mesobuthus eupeus, female holotype of Androctonus ornatus Nordmann, 1840. Dorsal (196) and ventral (197) views. Pedipalp chela dorsal (198). Chelicerae, carapace and tergites I–III (200) and sternopectinal region (201). The label say, in Latin: top: “Androctonus (Liurus) ornatus N-[ordma]nn Trans-Caucasus Nordm[ann]”; bottom (both in A. Birula´s hand): left, “Buthus eupeus eupeus 1♀ det. A. A. Birula”; right, “specimen typicum Androctonus (Lejuri) ornati Nordm.? AB”. Scale bar: 10 mm (196–197)

Figures 202–205. Mesobuthus eupeus. Figures 202–203. Male from Armenia, Syunik, Meghri, NMPC, dorsal (202) and ventral (203) views. Figures 204–205. Female neotype, dorsal (204) and ventral (205) views. Scale bars: 10 mm (202–203, 204–205).

Figures 206–213. Mesobuthus eupeus. Figures 206, 211–213. Male from Armenia, Syunik, Meghri, NMPC, telson lateral (206), metasoma and telson, lateral (211), dorsal (212), and ventral (213) views. Figures 207–210. Female neotype, telson lateral (207), metasoma and telson, lateral (208), dorsal (209), and ventral (210) views. Scale bars: 10 mm (208–210, 211–213).

Figures 214–235. Mesobuthus eupeus, pedipalp. Figures 214–224. Male from Armenia, Syunik, Meghri, NMPC, chela, dorsal (214), external (215), and ventral (216) views. Patella, dorsal (217), external (218) and ventral (219) views. Femur and trochanter, internal (220), dorsal (221), and ventral (222) views. Movable (223) and fixed (224) fingers. Figures 225–235. Female neotype, chela, dorsal (225), external (226), and ventral (227) views. Patella, dorsal (228), external (229) and ventral (230) views. Femur and trochanter, internal (231), dorsal (232), and ventral (233) views. Movable (234) and fixed (235) fingers. The trichobothrial pattern is indicated in Figures 214–218, 220–221.

Figures 236–243. Mesobuthus eupeus. Figures 236, 238, 240–243. Male from Armenia, Syunik, Meghri, NMPC, chelicerae, carapace and tergites I–III (236), sternopectinal region and sternites III–IV (238), left legs I–IV, retrolateral aspect (240–243). Figures 237, 239. Female neotype, chelicerae, carapace and tergites I–IV (237) and sternopectinal region and sternites III–V (239).

Figures 6–8: Mesobuthus afghanus, female lectotype. Figures 6–7. Dorsal (6) and ventral (7) views. Figure 8. Metasoma V and telson ventral view. Scale bar: 10 mm (6–7).