Therates fruhstorferi W. Horn, 1902

Matalin, Andrey V. & Wiesner, Jürgen, 2023, Revision of the Therates fruhstorferi complex (Coleoptera, Cicindelidae), Zootaxa 5256 (5), pp. 401-433 : 403-406

Therates fruhstorferi W. Horn, 1902

Figs 1–6 , 29–32 , 50–55 , 81–87 , 115–125 , 136–151 .

Therates fruhstorferi W. Horn, 1902: 72 (Type locality— Tonkin, Mouson-Gebirge).

Therates fruhstorferi W. Horn, 1902 — Horn 1908: 194, 1930: 400; Mell 1931: 290; Tan 1987: 539; Cassola 2004: 25.

Therates fruhstorferi fruhstorferi W. Horn, 1902 — Wiesner 1988: 41, 1992: 92, 2020: 95; Cassola 1985: 510; Lorenz 1998: 38, 2005: 117; Matalin 2001: 387, 2014: 300; Werner et al. 2002: 42; Shook & Wiesner 2006: 20; Shook & Wu 2006: 53, 2007: 68; Wu & Shook 2007: 39, 2010: 90; Wu 2011: 31; Putchkov & Matalin 2017: 245; Wiesner et al. 2017: 40 .

Therates fruhstorferi vitalisi W. Horn, 1913 — Sawada & Wiesner 1999c: 307, 2002: 84.

Type material. LECTOTYPE, ♂ (designated here)—“ Tonkin, Montes Mauson [= Mau So’n], April, Mai 2–3000, H. Fruhstorfer“ [printed white label with black border]; “Type!, coll. W Horn” [printed white label with black border]; “ Fruhstorferi , mihi” [handwritten light yellow label with black border]; “ Syntypus ” [printed red label]; “Coll. W. Horn, DEI Eberswalde” [printed white label]; “ fruhstorferi , s. str. W. Horn, det. Wiesner 86” [typewritten white label]; “ DEI Müncheberg, Col—04811” [printed light green label]; “ LECTOTYPE, Therates fruhstorferi W. Horn, 1902 , design. A. Matalin et J. Wiesner, 2016 ” [printed red label] ( SDEI). PARALECTOTYPES (designated here), 4♂♂ ( DEI Müncheberg, Col—04810, 04812, 04816, 04817) 5♀♀ ( DEI Müncheberg, Col—04809, 04813, 04814, 04815, 04818)—“ Tonkin, Montes Mauson, April, Mai 2–3000, H. Fruhstorfer” [printed white label with black border]; “Type! coll. W Horn” [printed white label with black border]; “ Syntypus ” [printed red label]; “Coll. W. Horn, DEI Eberswalde” [printed white label]; “ fruhstorferi , s. str. W. Horn, det. Wiesner 86” [typewritten white label]; “ PARALECTOTYPE, Therates fruhstorferi W. Horn, 1902 , design. A. Matalin et J. Wiesner, 2016 ” [printed red label] (all SDEI).

Additional material. CHINA, Yunnan Prov.: 1♂ — Wenshan, Funing County, Hong Mu Da Shan , 1851 m, 24.VII.2006 ( XX) ; VIETNAM, Ha Giang Prov.: 1♀ —N from Ha Giang , 500 m, 5.7.1963, leg. O. Kabakov ( ZIN) ; Yen Bai Prov.: 1♂ — Yen Bai , 6.2020 ( AA); 1♀ — ibid, 8.2020 ( AA) ; Tuyen Quang Prov.: 2♂♂ — 160 km NNW Ha Noi, NE env. of Na Hang , h = 150–200 m, 5– 10.6.1996, leg. A Napolov & I. Roma ( AN, MSPU); Thai Nguen Prov.: 1♂ 1♀ —mountains 50 km NO Thai Nguen, 300 m, 15.5.1963, 21.6.1963, leg. O. Kabakov ( ZIN); Vinh Phu Prov.: 4♂♂ 3♀♀ —mountains near Tam Dao , 900 m, 15.V.1962, 20.V.1962, 1.VII.1962, 29.VII.1962, 30.VIII.1963, leg. O. Kabakov ( ZIN); 3♂♂ 4♀♀ — Tam Dao , 3– 11.6.1985, leg. Ploka & Viša ( JW); 1♂ — ibid, 2– 11.6.1989, leg. Kubaň ( JW); 1♂ — ibid, 27.5– 2.6.1986, leg. Jan Horák ( JW); 3♂♂ 2♀♀ — ibid, 6– 10.5.1990, leg. Jan Horák ( JW); 3♀♀ — ibid, 12– 24.5.1989, leg. Jan Strnad ( JW); 2♀♀ — ibid, 6– 20.V.1990, leg. M. Kalabza & Z. Koštál ( JW); 1♂ — ibid, 800 m, 12.VI.1994, leg. Sugonjaev ( ZIN); 1♂ 4♀♀ — ibid, 25.V– 10.VI.1995, leg. Gorochov ( ZIN); 1♂ — ibid, 27– 28.06.2019, leg. D. Kasatkin ( MSPU); 2♂♂ 2♀♀ — 70 km NW Hanoi, Tam Dao , 21.27 oN 105.39 oE, 900–1200 m, 9—19.V.1996, leg. Pacholátko & Dembický ( JW); Ha Tay Prov.: 1♂ — 45 km W of Hanoi, Ba Vi Nat. Park , h = 1050– 1100 m, leaf-litter, 21 o 04’N 105 o 21’38’’E, 18.VI.2014, leg. D. Fedorenko ( SIEE) ; Phu Tho Prov.: 11♂♂ 1♀ —~ 90 km W of Hanoi, Xuan Son Nat. Park , h = 400–700 m, 21 o 07’01’’N 104 o 56’39’’E, 6– 15.VI.2014, leg. D. Fedorenko ( SIEE, MSPU); 3♂♂ — ibid, h = 400–470 m, 21 o 07’52’’N 104 o 57’07’’E, 6– 15.VI.2014, leg. D. Fedorenko ( SIEE); 2♂♂ — ibid, h = 300 m, 21 o 07’58’’N 104 o 55’45’’E, river-valley, 6– 15.VI.2014, leg. D. Fedorenko ( SIEE) ; Hoa Binh Prov.: 2♂♂ 1♀ — Yen Thuy distr., Loc Thin , Cuc Phuong Nat. Park , 20 o 18’N 105 o 35’E, 3– 4.V.2002, leg. S. Belokobylsky ( ZIN); 2♂♂ 1♀ — ibid, 20 o 23’N 105 o 34’E, 5– 6.V.2002, leg. S. Belokobylsky ( ZIN) ; Thanh Hoa Prov.: 1♀ — Ban Vin env., Xuan Lien Natn. Park , h ~ 900 m, 19 o 58’45’’N 104 o 58’13’’E, 2– 11.VI.2022, leg. D. Fedorenko ( SIEE) ; Nghe An Prov.: 1♀ — NO Con Cuong , 400 m, 22.04.1962, leg. O. Kabakov ( ZIN); 2♂♂ — Que Phong distr., Pu Hoat National Park , h = 840 m, 19 o 45’19’’N 104 o 47’47’’E, 15– 27.V.2019, leg. D. Fedorenko ( SIEE, MSPU) ; Ha Tinh Prov.: 1♀ — Kim Quang env., Vu Quang Natn. Park , h ~ 440 m, 18 o 16’26’’N 105 o 21’41’’E, 24.V– 1.VI.2022, leg. D. Fedorenko ( SIEE) ; Quang Binh Prov.: 1♂ —NW Dong Hoi , 200 m, 26.II.1963, leg. O. Kabakov ( ZIN); 1♂ — Minh Hoa distr., Yen Hop env., 20.IV.1999, leg. A. Devyatkin ( SIEE); 1♂ 1♀ —Phong Nha-Ke Bang N.P ., Bo Trach , h ~ 400 m, 17 o 25’45’’N 106 o 14’04’’E, 12– 21.V.2022, leg. D. Fedorenko ( SIEE, MSPU) ; LAOS, Boli Kham Xai Prov.: 6♂♂ 7♀♀ — Ban Nape (8 km NE), ~ 600 m, 18 o 21’N 105 o 08’E, 1– 18.V.2001, P. Pacholátko leg. ( JW, MSPU); 1♂ 3♀♀ — Ban Nape—Kaev Nua Pass , alt. 600 ± 100 m, 18 o 22.3’N 105 o 09.1’E ( GPS), 18.4– 1.5.1998, E. Jendek & O. Šauša leg. ( JW; MSPU) .

References. CHINA, Yunnan Prov.: Yongshan County, 1800 m ( Tan 1987) ; Zhaotong , Daguan County, 1970 m, 3.VII.1991, Lin Su ( Shook & Wu 2007) ; VIETNAM, Lang Son Prov.: Than Moi ( Wiesner 1988) ; Cao Bang Prov.: 1 ex. —vic. Vin Den, Nui Pia Oac Nature Reserve , 22 o 33’53”N 105 o 52’53”E, 900–1300 m, 6–10.V.2013, leg. A. Skale ; 1 ex. —vic. Tinh Tuc, Son Dong, Nui Pia Oac Nature Reserve , 22 o 37’55”N 105 o 52’98”E, 850–1300 m, 9–15.V.2014, leg. A. Skale ; Bac Giang Prov.: 1♂— Tay Yen Tu Nature Reserve , 6 km SW Thanh Son, 21 o 10’49.8”N 106 o 43’25.62”E, 200 m, 18.V.2015, leg. R. Gerstmeier (as T. vitalisi ) ; Phu Tho Prov.: 1♂ 8♀♀ — Xuan Son National Park , 500 m, 18–20.VI.2012, leg. L. Bartolozzi, S. Bambi, F. Fabiano, E. Orbach ; Vinh Phuc Prov.: 2ex.1♀ —vic. Tam Dao Town, Tam Dao National Park , 21 o 27’N 105 o 38’E, 700–1000 m, 2–5.V.2013, leg. A. Skale, A. Weigel ; 2♂♂ 1♀ — Tam Dao National Park , 950 m, 1–4.VI.2013, leg. L. Bartolozzi, S. Bambi, F. Cianferoni, G. Mazza, E. Orbach ; 6♂♂ 1♀ — Tam Dao , 21 o 27’52”N 105 o 38’46”E, 1200 m, 9–22.VI.2014, leg. M. Hauser and N. von Ellenrieder (all Wiesner et al. 2017) ; 1♀ — Tam Dao , 1100 m, 2–6.VI.1999, leg. Ahrens, Jaeger, Fabrizzi ( Werner et al. 2002) ; Ha Noi City: 1♂ 1♀ — Ba Vi Dist., Ba Vi National Park , 700 m, 21–24.VI.2012, leg. E. Orbach ; Hoa Binh Prov.: 1♂ 1♀ — Pa Co Hang Kia Nature Reserve , 900–1200 m, 9–12.VI.2013, leg. L. Bartolozzi, S. Bambi, F. Cianferoni, G. Mazza, E. Orbach ; Ninh Binh Prov.: 1♂— Cuc Phuong National Park , 20 o 17’34.32”N 105 o 40’03.12”E, 270 m, 22.V.2015, leg. R. Gerstmeier (all Wiesner et al. 2017) ; Nghe An Prov.: Vinh ( Wiesner 1988) ; Thua ThienHue Prov.: 3♂♂— Bach Ma National Park , 1400 m, 16 o 12’00”N 107 o 51’00”E, 23–28.V.2014, leg. A. Bandinelli, L. Bartolozzi, S. Bambi ( Wiesner et al. 2017) .

Diagnosis. Most specimens of T. f. fruhstorferi are clearly distinguished from other congeners by a complete humeral lunule, distinct basal dot, medium-sized or narrow apical dot as well as by partly pale hind tibiae and tarsi ( Figs 4, 5 , 83 ). In some specimens the humeral lunule is divided on the basal and apical portions ( Fig. 85 ) or present only by the basal portion ( Figs 1, 3 , 81, 82, 84 ). Dark-coloured specimens of T. fruhstorferi ( Figs 6 , 86, 87 ) are differentiated from the similarly coloured T. ziyardamensis sp. n. ( Fig. 12 ) and T. pseudovitalisi sp. n. ( Figs 9–11 ) by the shape of aedeagus ( Figs 115–125 , 136–151 vs. Fig. 131 and Figs 132–135 , respectively) as well as by the longer hind tarsi, especially in males ( Figs 175–178 ).

Redescription. TL = 9.2–12.0 mm (mean = 10.5 mm, n = 45) in males, 9.8–12.8 mm (mean = 11.5 mm, n = 43) in females.

Head shining black with distinct violet reflection; orbital plates indistinctly or finely striated in posterior third; frons smooth, frontal sulci deep and slightly convergent in anterior 2/3, shallow and sharply divergent in posterior third. Mandibles brown or light-brown underside, pale-yellow in basal half of topside, with light-brown teeth and apical molar. Labial palpi pale-yellow except brown apical palpomere; maxillary palpi brown except yellowish palpomeres 1 and 2. Antennae do not extend posteriorly to the shoulders, scape pale anteriorly, brownblack posteriorly, in some females entirely dark-brown or black; antennomeres 9 and 10 with protruding downward anterior lower margins, more distinct in males ( Figs 29–32 ). Labrum varies from sub-quadrate to slightly transverse, LW/LL = 0.94–1.14 (mean = 1.05, n = 88), from dark-brown to black with large or medium-sized yellow central spot usually not extending to the border of the central apical teeth in males but to the border in females ( Figs 50–54 ), in some females central labral spot small and dark brown ( Fig. 55 ).

Pronotum as wide as long, PW/PL = 0.92–1.1 (mean = 1.03, n = 88), shining black-blue with violet reflection, apical lobe slightly wider in middle part; thorax black-blue with violet reflection.

Fore femora pale-yellow with brown violet-tinged posterior side and apices, middle femora pale-yellow with brown anterior side and light violet tinge, hind femora bicoloured, pale-yellow or light brown in basal half or third and black with blue tinge in apical half or 2/3; tibia often pale-yellow with brown-black apices or apical half, sometimes light brown, dark brown or even entirely black with light blue tinge; tarsomeres 1 and 2 or only tarsomere 1 of all legs in most specimens pale-yellow with brown apices, other tarsomeres dark-brown or black, in some specimens two basal tarsomeres brown with dark-brown apices and brown-black other tarsomeres, sometime only first fore and middle tarsomeres or only fore-tarsomere dark-brown while others including all hind tarsomeres black, rarely all tarsomeres black; HTbL/HTaL = 1.15–1.33 (mean = 1.22, n = 77) ( Figs 175, 176 ).

Elytra indistinctly divergent towards apex; EL/EW = 1.86–2.09 (mean = 1.98, n = 71); EL/PL = 3.14–3.57 (mean = 3.37, n = 45) in males, 3.14–3.75 (mean = 3.48, n = 42) in females; brown-black or black with violet or blue-violet reflection; punctuation deep and regular between basal hump and central dot, then shallow and sparse, apical quarter virtually smooth; scutellum dark-brown or black with blue tinge, apical margin shortly cut. Elytral pattern presented by relatively large or dramatically small yellow or light brown humeral lunule complete in one specimens ( Figs 4, 5 , 83 ) or divided on basal and apical portions in others ( Fig. 85 ), sometimes apical portion undeveloped ( Figs 1, 3 , 81, 82, 84, 86, 87 ), by yellow basal dot well developed in most specimens ( Figs 1, 3–5 , 81–85 ) and barely visible to absent in others ( Figs 6 , 86, 87 ), by transverse slightly oblique central dot ( Figs 1, 3–6 , 81–87 ), as well as by narrow apical dot along elytral margin ( Figs 3–5 , 83, 85 ), in some specimens only by thin yellow, yellow-brown ( Figs 1 , 81, 82, 84, 87 ) or rarely dark-brown ( Figs 6 , 86 ) apical edge.

Aedeagus in most specimens with moderately long and narrow slightly sinuate rounded apex ( Figs 115–118, 120–125 , 136, 137, 139, 141–149, 151 ), in some specimens with very thin ( Fig. 150 ) or dramatically short blunt apex ( Figs 119 , 138, 140 ) without lateral carinae; AL = 2.4–3.0 mm (mean = 2.67, n = 40), EL/AL = 2.19–2.68 mm (mean = 2.47, n = 40).

Distribution. CHINA: Yunnan Province; VIETNAM: Lang Son, Cao Bang, Ha Giang, Yen Bai, Bac Giang, Tuyen Quang, Thai Nguen, Vinh Phu, Ha Tay, Phu Tho, Ha Noi City, Hoa Binh, Thanh Hoa, Ninh Binh, Nghe An, Ha Tinh, Quang Binh, Thua Thien-Hue Provinces; LAOS: Bolikhamsai Province ( Fig. 184 ).

Figures

FIGURES 184. Distribution of Therates spp.: dark blue solid circle—T. vitalisi vitalisi; dark blue half-solid circle—T. vitalisi ida; lilac solid circle—T. hunnanensis sp. n.; orange solid circle—T. biserratus; pink solid circle—T. motoensis; red open circle—T. sauteri; red solid quadrat—T. fruhstorferi fruhstorferi; red half-solid quadrat—T. fruhstorferi australis ssp. n.; blue solid quadrat—T. ziyardamensis sp. n.; green solid quadrat—T. pseudovitalisi sp. n. (nominotypical subspecies); green halfsolid quadrat—T. pseudovitalisi tibetana ssp. n.; L in the symbols—according only to the literature sources. Map source— https://commons.wikimedia.org/wiki/File:East_Asia_topographic_map.png.

FIGURES 1–14. Therates spp., habitus: 1–6—T. fruhstorferi fruhstorferi (1–3—Vietnam, Montes Mouson; 4—Vietnam, Na Hang; 5—Vietnam, Tam Dao; 6—Laos, Ban Nape); 7, 8—T. fruhstorferi australis ssp. n. (Vietnam, Kon Ka Kinh); 9, 10—T. pseudovitalisi sp. n. (9—Laos, Bang Son Cha; 10—Laos, Ban Bun Neua); 11—T. pseudovitalisi tibetana ssp. n. (China, Xia- Cha-Yu); 12—T. ziyardamensis sp. n. (Myanmar, Zi Yar Dam); 13, 14—T. motoensis (China, Hanmi); 1, 2, 4, 5, 7, 9–13— males; 3, 6, 8, 14—females; 1, 2—lectotype; 3—paralectotype; 7, 9, 11, 12—holotypes; 8, 10—paratypes; 1, 3–14—habitus; 2—type labels.

FIGURES 29–49. Therates spp., left antennomeres 6–11: 29–32—T. fruhstorferi fruhstorferi (29, 30 – Vietnam, Na Hang; 31, 32 – Laos, Ban Nape); 33, 34—T. fruhstorferi australis ssp. n. (33—Vietnam, Kon Ka Kinh N.P.; 34—Vietnam, Pak Khe River); 35—T. ziyardamensis sp. n. (Myanmar, Zi Yar Dam); 36, 37—T. pseudovitalisi pseudovitalisi ssp. n. (36—Laos, Ban Song Cha; 37—Laos, Ban Bun Neua); 38, 39—T. pseudovitalisi tibetana ssp. nov. (China, Xia-Cha-Yu); 40, 41—T. motoensis (China, Hanmi); 42, 43—T. hunnanensis sp. n. (China, Shunhuangshan); 44—T. biserratus (China, Fanjing Shan-Kuaichang); 45, 46—T. vitalisi vitalisi (Vietnam, Xuan Son N.P.); 47—T. vitalisi ida (China, Fukien); 48, 49—T. sauteri (48—China, Taiwan, Taipei Hsien; 49—China, Taiwan, Ilan Hsien); 30, 32, 34, 35, 37, 39, 41, 43, 44, 46, 49—males; 29, 31, 33, 36, 38, 40, 42, 45, 47, 48—females; 33, 35, 37, 39, 41—holotypes; 34, 36, 38, 42—paratypes. Without scales.

FIGURES 50–57. Therates fruhstorferi ssp., labrum: 50–55—T. fruhstorferi fruhstorferi (50, 51—Vietnam, Montes Mauson; 52—Vietnam, Na Hang; 53—Vietnam, Tam Dao; 54, 55—Laos, Ban Nape); 56, 57—T. fruhstorferi australis ssp. n. (56—Vietnam, Kon Ka Kinh N.P.; 57—Vietnam, Pak Khe River); 50, 52, 54, 56—males; 51, 53, 55, 57—females; 50, 51—paralectotypes; 56—holotype; 57—paratype.

FIGURES 81–98. Therates spp., left elytron: 81–87—T. fruhstorferi fruhtorferi (81, 82—Vietnam, Montes Mauson; 83— Vietnam, Na Hang; 84—Vietnam, Pu Hoat N.P.; 85—Vietnam, Tam Dao; 86—Laos, Ban Nape—Kaev Nua Pass; 87—Laos, Ban Nape); 88, 89—T. fruhstorferi australis ssp. n. (88—Vietnam, Kon Ka Kinh N.P.; 89—Vietnam, Pak Khe River); 90– 93—T. pseudovitalisi pseudovitalisi ssp. n. (90, 93—Laos, Ban Song Cha; 91—Laos, Ban Bun Neua; 92—Laos, Oudom Xai); 94, 95—T. pseudovitalisi tibetana ssp. n. (China, Xia-Cha-Yu); 96—T. ziyardamensis sp. n. (Myanmar, Zi Yar Dam); 97, 98—T. motoensis (China, Hanmi); 81, 83, 84, 87, 88, 90, 91, 94, 96, 97—males; 82, 85, 86, 89, 92, 93, 95, 98—females; 81, 82—paralectotypes; 88, 90, 94, 96—holotypes; 89, 91–93, 95—paratype.

FIGURES 115–135. Therates spp., aedeagus: 115–125—T. fruhstorderi fruhstorferi (115, 116—Vietnam, Vietnam, Montes Mouson; 117–121—Vietnam, Xuan Son N.P.; 122, 123—Vietnam, Tam Dao; 124—Laos, Ban Nape; 125—Laos, Ban Nape— Kaev Nua pass); 126–130—T. fruhstorferi australis ssp. n. (126—Vietnam, Kon Ka Kinh N.P.; 127—Vietnam, Gia Lai; 128— Vietnam, Ngok Boc I Mt.; 129, 130—Vietnam, song Thanh N.P.); 131—T. ziyardamensis sp. n. (Myanmar, Zi Yar Dam); 132– 134—T. pseudovitalisi pseudovitalisi ssp. n. (132—Laos, Ban Song Cha; 133, 134—Laos, 17 km NEE Oudom Xai); 135—T. pseudovitalisi tibetana ssp. n. (China, Xia-Cha-Yu); 115—lectotype; 116—paralectotype; 126, 131, 132, 135—holotypes; 127–130, 133, 134—paratypes; 117—dorsal view; 115, 116, 118–135—left lateral view. Scale bars in mm.

FIGURES 136–151. Therates fruhstorferi fruhstorferi, variability of aedeagus, left lateral view: 136–143—Vietnam, Phu Tho Province, ~90 km W of Hanoi, Xuan Son Nat. Park, h = 400–700 m, 21o07’01’’N 104o56’39’’E, 6–15.VI.2014; 144–151— Vietnam, Vinh Phu Province, Tam Dao, different years (144, 145—3–11.VI.1985; 146—27.V–2.VI.1986; 147—2–11.VI.1989; 148—6–10.V.1990; 149—12.VI.1994; 150, 151—9–19.V.1996). Scales in mm.

FIGURES 175–178. Morphometric features of Therates spp: 175, 177—relation of the proportions EL/PL and HTbL/HTaL (green rhombus—T. pseudovitalisi sp. n.; blue circles—T. fruhstorferi; open symbols—males; solid symbols—females); 176, 178—relation of the proportions HTbL/HTaL (small quadrates—medians; rectangles—25–75 percentiles; whiskers—min–max; significantly different samples are marked with different lower letters, according to Mann–Whitney U-test, p <0.05 (176—U = 112.0, z = 5.08, p = 0.000; 178—U = 23.5, z = 3.37, p = 0.0007)); 175, 176—all specimens; 177, 178—males.