Notes on the Genera Graphidessa Bates, 1884, Pararondibilis Breuning, 1961, Paratimiola Breuning, 1965, and Tuberenes Breuning, 1978 (Coleoptera, Cerambycidae, Lamiinae)

Simple Summary

The genera Graphidessa Bates, 1884 (presently consisting of four species and one subspecies), Pararondibilis Breuning, 1961 (presently consisting of five species), Paratimiola Breuning, 1965 (presently consisting of one species), and Tuberenes Breuning, 1978 (presently consisting of four species) are small groups of the subfamily Lamiinae. The genera Graphidessa and Tuberenes have been studied well, but we found that some species of it should be transferred to the genera Pararondibilis and Paratimiola. In this paper, we provide a taxonomic review of some species of these four genera.

Abstract

Taxonomic reviews of some species of Graphidessa Bates, 1884, Pararondibilis Breuning, 1961, Paratimiola Breuning, 1965, and Tuberenes Breuning, 1978 are provided herein. The following new combinations are proposed: Graphidessa variegata Hayashi, 1974, Tuberenes minuta (Pic, 1925), and Tuberenes sikkimensis Breuning, 1978 are transferred to Pararondibilis; Tuberenes vietnamensis Breuning, 1972 is transferred to Paratimiola. The replacement name Pararondibilis rubina nom. nov. is proposed for Pararondibilis sikkimensis (Breuning, 1978). Updated species checklists for Graphidessa, Pararondibilis, Paratimiola, and Tuberenes are provided. Keys to species of Graphidessa and Pararondibilis are provided. Tuberoenes Breuning, 1972 is a nomen nudum of Tuberenes.

1. Introduction

Bates established Graphidessa for Graphidessa venata Bates, 1884 [1], and Liu et al. described Graphidessa jinfoensis (from China) and made a key to all species [2]. Graphidessa presently consists of four species and one subspecies from East Asia [3]. Breuning established Pararondibilis for Pararondibilis sikkimensis Breuning, 1961 (from India) [4]; subsequently, Pararondibilis acrosa Holzschuh, 2003 (from Nepal), P. eluta Holzschuh, 2003 (from Nepal), P. macularia Holzschuh, 2003 (from India), and Pararondibilis pinicola Holzschuh, 2018 (from Nepal) were described [5,6]. Pararondibilis presently consists of these five species [3]. Breuning established Paratimiola for Paratimiola rondoni Breuning, 1965 (from Laos) [7], which is presently the only described species of the genus [3]. Breuning established Tuberenes for Eryssamena robustipes Pic, 1939 (from China); he also described Tuberenes sikkimensis (from India) and transferred Eryssamena minuta Pic, 1925 (from Vietnam) and Tuberoenes vietnamensis Breuning, 1972 (from Vietnam) to Tuberenes [8]. Tuberenes presently consists of these four species [3].

We found some issues suggesting that the taxonomic status of Graphidessa variegata Hayashi, 1974, Tuberenes minuta (Pic, 1925), Tuberenes sikkimensis Breuning, 1978, and Tuberenes vietnamensis Breuning, 1972 are doubtful. Therefore, this paper reviews these issues.

2. Materials and Methods

The specimens examined in this study are deposited in the following institutional and private collections:

BPBM Bernice Pauahi Bishop Museum, Honolulu, HI, USA;

CHS Collection Carolus Holzschuh, Villach, Austria;

HNHM Hungarian Natural History Museum, Budapest, Hungary;

IZCAS Insect collection of the Institute of Zoology, Chinese Academy of Sciences, Beijing, China;

MNHN Muséum National d’Histoire Naturelle, Paris, France.

The copyrights of other photographs were added to the legends of the corresponding figures. All photographs and figures herein were produced using Photoshop CS5 software (Adobe company, San Jose, CA, USA).

3. Results

Pararondibilis Breuning, 1961

Pararondibilis Breuning, 1961: 547 [4]; Breuning 1963: 523 (catalogue) [9]; Löbl and Smetana 2010: 210 (catalogue) [10]; Lin 2015: 158 (catalogue) [11]; Kariyanna et al., 2017: 92 (catalogue) [12]; Danilevsky 2020: 296 (catalogue) [13]. Type species: Pararondibilis sikkimensis Breuning, 1961.

Comparative analyses of the holotypes of Graphidessa variegata Hayashi, 1974, Tuberenes minuta (Pic, 1925), and Tuberenes sikkimensis Breuning, 1978 reveal that these species do not belong to Graphidessa or Tuberenes because they lack several diagnostic characteristics of these genera: the post-basal bump on each elytron is not covered with a small fascicle of setae; each elytron is not covered with short white or pale yellow pubescence forming several longitudinal lines at the apical half; and the femora is not covered with erect pubescence.

However, comparative analyses of the holotypes of Pararondibilis sikkimensis (Figure 2H,I), Eryssamena minuta (now Tuberenes minuta) (Figure 2A), Tuberenes sikkimensis (Figure 2C–F), and Graphidessa variegata (Figure 2K,L) reveal that the latter three species belong to the genus Pararondibilis based on the shared commonality of morphological characteristics with P. sikkimensis: the scape is short and quite thick, antennomere III is longer than the scape, antennomere III is shorter than antennomere IV, the pronotum has a tubercle at the middle sides, each elytron has a post-basal bump, and the mesocoxal cavities open externally to the mesepimera.

To date, Pararondibilis consists of eight species.

Distribution: China, India, Nepal, Vietnam.

Pararondibilis acrosa Holzschuh, 2003

Pararondibilis acrosa Holzschuh, 2003: 313 (type locality: “Nawakot, Trisuli Khola, Dhunche-Syabru Bensi, Nepal”), pl. VI, fig. 11 (holotype, male) [5]; Weigel 2006: 506 (catalogue) [14]; Löbl and Smetana 2010: 210 (catalogue) [10]; Lin 2015: 158, figs. page 158 bas and 159 (paratypes) [11]; Danilevsky 2020: 296 (catalogue) [13].

Type material examined: One paratype (CHS), C-NEPAL, NAWAKOT, Trisuli Khola, 2200–1600 m Dhunche-Syabru Bensi, 28.IX.1982, leg. C. Holzschuh (printed with black ink on a rectangular white label)/GESCHLÜPFT AUS DÜRREM ÄSTEN: 2.7.1983 (“GESCHLÜPFT AUS DÜRREM ÄSTEN:” printed and “2.7.1983” handwritten with black ink on a rectangular white label)/PARATYPUS Pararondibilis acrosa n. sp. det. C. Holzschuh 2003 (printed with black ink on a rectangular red label), examined from two photographs (Figure 1B,C); one paratype, ♂ (IZCAS), NEPAL oc. 2500 m ü.NN 25 km N Jumla, Pina W Jhyari Kh. 29°29′47N″ 82°07′51″E, 23.VI.1999 leg. A. Weigel KL (printed with black ink on a rectangular white label)/PARATYPUS Pararondibilis acrosa n.sp. det.C. Holzschuh 2003 (printed with black ink on a rectangular red label)/IOZ(E)1859491 (printed with black ink on a rectangular yellow label), examined from two photographs (Figure 1D,E); one paratype, ♂ (IZCAS), NEPAL oc. 2500 m ü.NN 25km N Jumla, Pina W Jhyari Kh. 29°29′47N″ 82°07′51″E, 23.VI.1999 leg. A. Weigel KL (printed with black ink on a rectangular white label)/von Juglans regia geldopft (printed with black ink on a rectangular white label)/PARATYPUS Pararondibilis acrosa n.sp. det. C.Holzschuh 2003 (printed with black ink on a rectangular red label)/IOZ(E)1859490 (printed with black ink on a rectangular yellow label), examined from three photographs (Figure 1F–H).

Distribution: Nepal (Dhunche-Syabru Bensi).

Remarks: Holzschuh described the body color of this species as variable from reddish-brown to blackish, with elytra either without or with two dark transverse bands (a narrow one extending between the humeri and covering the post-basal bumps, another one ascending somewhat obliquely to the lateral margin in the center, usually not touching either the lateral margin or the suture) [5]. However, we found the holotype (Figure 1A) and two paratypes (Figure 1D,G,H) have very different colors of their antennae, pronotum, elytra, and leg, except for the characteristics mentioned above. Therefore, we suggest that the two paratypes may not be Pararondibilis acrosa. Andreas Weigel told the senior author that Carolus Holzschuh also thought there may be two species in the type series of Pararondibilis acrosa.

Pararondibilis eluta Holzschuh, 2003

Pararondibilis eluta Holzschuh, 2003: 314 (type locality: “Dhawalagiri, Kali-Gandaki-Khola, Kalopani, Nepal”) [5]; Weigel 2006: 506 (catalogue) [14]; Löbl and Smetana 2010: 210 (catalogue) [10]; Danilevsky 2020: 296 (catalogue) [13].

Distribution: Nepal (Kalopani).

Pararondibilis macularia Holzschuh, 2003

Pararondibilis macularia Holzschuh, 2003: 314 [type locality: “Jammu, Yourdu, Jammu and Kashmir (Kishtwar District), India”] [5]; Löbl and Smetana 2010: 210 (catalogue) [10]; Kariyanna et al., 2017: 92 (catalogue) [12]; Danilevsky 2020: 296 (catalogue) [13].

Distribution: India (Kishtwar).

Pararondibilis minuta (Pic, 1925) comb. nov.

Eryssamena minuta Pic, 1925: 31 (type locality: “Tonkin, Vietnam”) [15]; Breuning 1963: 523 (catalogue) [9].

Tuberenes minuta: Breuning 1978: 21 (redescription) [8].

Type material examined: Holotype (MNHN): Lac Thô Tonkin R.P. a de Cooman (handwritten with black ink on a rectangular white label)/Eryssamena minuta n sp (handwritten with black ink on a rectangular white label)/Museum Pairs Coll. M. Pic (printed with black ink on a rectangular white label with black borders)/Type (handwritten with black ink on a rectangular white label)/TYPE (printed with black ink on a red rectangular label), examined from two photographs (Figure 2A,B).

Distribution: Vietnam (Tonkin).

Pararondibilis pinicola Holzschuh, 2018

Pararondibilis pinicola Holzschuh, 2018: 504 (type locality: “Kathmandu, Nepal”), figs. 2 (holotype male), 2 (paratype female) [6]; Danilevsky 2020: 296 (catalogue) [13].

Distribution: India (Kumaon Himalaya), Nepal (Kathmandu).

Pararondibilis rubina nom. nov.

Tuberenes sikkimensis Breuning, 1978: 21 (type locality: “Lachen-Lachung, Sikkim, India”) [8]; Breuning 1982: 23 (redescription) [16]; Löbl and Smetana 2010: 213 (catalogue) [10]; Danilevsky 2020: 299 (catalogue) [13].

Type material examined: Holotype (MNHN): BRITISH INDIA SIKKIM Lachen-Lachung (printed with black ink on a rectangular white label)/Tuberenes sikkimensis mihi Typ Breuning dét. (“Tuberenes sikkimensis mihi Typ” handwritten with blue ink and “Breuning dét.” printed with black ink on a rectangular white label)/HOLOTYPE (printed with black ink on a rectangular red label)/HOLOTYPE Tuberenes sikkimensis Breuning, 1978 (printed with black ink on a rectangular white label)/MNHN, Paris EC47847 plus a QR code (printed with black ink on a rectangular white label), examined from five photographs (Figure 2C–G).

Etymology: The new specific epithet of this species is derived from the Latin word “rubina”, referring to its dark red body.

Distribution: India (Sikkim).

Remarks: Breuning first described Tuberenes sikkimensis [8]; then, Breuning described Tuberenes sikkimensis again as a new species [16]. Therefore, the available published date of Tuberenes sikkimensis is 1978, following Article 23.1 of ICZN [17].

Since T. sikkimensis belongs to Pararondibilis, it becomes a secondary homonym of Pararondibilis sikkimensis Breuning, 1961. According to ICZN Art. 57.3.1, Art. 59.1, and Art. 60.3 [17], we hereby propose the replacement name Pararondibilis rubina nom. nov. for Pararondibilis sikkimensis (Breuning, 1978).

Pararondibilis sikkimensis Breuning, 1961

Pararondibilis sikkimensis Breuning, 1961: 547 (type locality: “Pedong, Sikkim, India”) [4]; Breuning, 1963: 523 (catalogue) [9]; Breuning, 1977: 137 (redescription), pl. II, fig. 5 [18]; Löbl and Smetana 2010: 210 (catalogue) [10]; Kariyanna et al., 2017: 92 (catalogue) [12]; Danilevsky 2020: 296 (catalogue) [13].

Type material examined: Holotype (MNHN): Inde Anglaise Pedong Région de Darjecling. Chasseurs indigènes 1934 (printed with black ink on a rectangular white label with black borders)/MUSÉUM PARIS 1952 COLL R OBERTHUR (printed with black ink on a rectangular white label with black borders)/Pararondibilis sikkimensis mihi Typ Breuning dét (“Pararondibilis sikkimensis mihi Typ” handwritten and “Breuning dét” printed with black ink on a rectangular white label)/TYPE (printed with black ink on a rectangular red label), examined from three photographs (Figure 2H–J).

Distribution: India (Sikkim).

Pararondibilis variegata(Hayashi, 1974) comb. nov.

Graphidessa variegata Hayashi, 1974: 50 (type locality: “Lishan in Tachiachi, Taichung Hsien, Taiwan, China”) [19]; Hua 1982: 88 (catalogue) [20]; Nakamura et al., 1992: 92 (catalogue) [21]; Hua 2002: 211 (catalogue) [22]; Chou 2004: 321, fig. male (misidentified) [23]; Mizuno and Shiyake 2004: 52 (catalogue), pl. 21, fig. 476 (holotype) [24]; Hua et al., 2009: 84, pl. LXXXIV, fig. 972 (male), 219 (redescription in Chinese), 361 (redescription in English) [25]; Löbl and Smetana 2010: 223 (catalogue) [10]; Nakamura et al., 2014: 155 (catalogue) [26]; Lin and Yang 2019: 260 (catalogue) [27]; Danilevsky 2020: 314 (catalogue) [13]; Liu et al., 2023: 17, fig. 1 (distribution map), 22, fig. 7C (elytra of holotype, male), 23 (key) [2].

Hayashi compared Graphidessa variegata with two known species (Graphidessa venata Bates, 1884 and Graphidessa obliquefasciata Komiya and Kusama, 1974) of Graphidessa when he described it [19]. However, comparative analyses of the holotypes of Graphidessa venata venata Bates, 1884 (Figure 3A–C, type species of Graphidessa) and G. variegata (Figure 2K,L) reveal that G. variegata does not belong to Graphidessa and should be transferred to Pararondibilis Breuning, 1961, as mentioned above.

Figure 2. (A,B) Eryssamena minuta, holotype: (A) dorsal habitus; (B) labels (photographs (A,B) were taken by Xavier Gouverneur). (C–G) Tuberenes sikkimensis, holotype, male: (C) dorsal habitus; (D) lateral habitus; (E) ventral habitus; (F) frontal habitus; (G) labels (photographs (C–G) were taken by Christophe Rivier). (H–J) Pararondibilis sikkimensis, holotype: (H) dorsal habitus; (I) lateral habitus; J labels (photographs (H–J) were taken by Xavier Gouverneur). (K,L) Graphidessa variegata, holotype, male: (K) dorsal habitus (photograph (K) was reproduced from Mizuno and Shiyake 2004 [24]); (L) elytra (photograph (L) was reproduced from Liu et al., 2023 [2]).

Figure 2. (A,B) Eryssamena minuta, holotype: (A) dorsal habitus; (B) labels (photographs (A,B) were taken by Xavier Gouverneur). (C–G) Tuberenes sikkimensis, holotype, male: (C) dorsal habitus; (D) lateral habitus; (E) ventral habitus; (F) frontal habitus; (G) labels (photographs (C–G) were taken by Christophe Rivier). (H–J) Pararondibilis sikkimensis, holotype: (H) dorsal habitus; (I) lateral habitus; J labels (photographs (H–J) were taken by Xavier Gouverneur). (K,L) Graphidessa variegata, holotype, male: (K) dorsal habitus (photograph (K) was reproduced from Mizuno and Shiyake 2004 [24]); (L) elytra (photograph (L) was reproduced from Liu et al., 2023 [2]).

Distribution: China (Taiwan).

Key to species of Pararondibilis

• 1. Elytral apices subtruncate..................................................................................................................................................................................................................P. pinicola
• - Elytral apices sub-rounded or acute....................................................................................................................................................................................................................2
• 2. Elytral apices sub-rounded.................................................................................................................................................................................................................................3
• - Elytral apices acute................................................................................................................................................................................................................................................4
• 3. Each elytron with a median grayish-yellow transverse band running up towards the lateral edge...........................................................................................P. minuta
• - Each elytron without a median grayish-yellow transverse band running up towards the lateral edge........................................................................................P. rubina
• 4. Elytron without a band basally...........................................................................................................................................................................................................................5
• - Elytron with a band basally..................................................................................................................................................................................................................................6
• 5. Elytra densely covered with short yellowish-gray pubescence..................................................................................................................................................P. macularia
• - Elytra densely covered with short gray pubescence............................................................................................................................................................................P. acrosa
• 6. Elytra with a pair of oblique black bands in middle..........................................................................................................................................................................P. acrosa
• - Elytra without a pair of oblique black bands in middle....................................................................................................................................................................................7
• 7. Post-basal bump on elytron angularly processed in profile.........................................................................................................................................................P. variegata
• - Post-basal bump on elytron flat in profile...........................................................................................................................................................................................................8
• 8. Elytra not covered with protruding pubescence........................................................................................................................................................................P. sikkimensis
• - Elytra covered with protruding pubescence............................................................................................................................................................................................P. eluta

Graphidessa Bates, 1884

Graphidessa Bates, 1884: 248 [1]; Aurivillius 1922: 231 [28]; Breuning 1963: 492 (catalogue) [9]; Breuning 1975: 7 (key) [29]; Breuning 1976: 75 (redescription) [30]; Nakamura et al., 1992: 92 (catalogue) [21]; Ohbayashi and Niisato 2007: 618 (catalogue) [31]; Löbl and Smetana 2010: 223 (catalogue) [10]; Nakamura et al., 2014: 155 (catalogue) [26]; Lin and Yang 2019: 260 (catalogue) [27]; Danilevsky 2020: 314 (catalogue) [13]. Type species: Graphidessa venata Bates, 1884.

As mentioned above, Graphidessa variegata Hayashi, 1974 was transferred to Pararondibilis. Graphidessa now consists of three species and one subspecies.

Distribution: China, Japan.

Graphidessa jinfoensis Liu et al., 2023

Graphidessa jinfoensis Liu et al., 2023: 17 (type locality: “Jinfoshan National Nature Reserve, Chongqing, China; Dongfeng Lake National Wetland Park, Xishui County, Zunyi City, Guizhou Province, China”), fig. 1 (distribution map), fig. 2A (holotype, male), 2B (paratype, female), 3A (holotype, male), 3B (paratype, female), 4A (details of pronotum of holotype, male), 4B (details of pronotum of paratype, female), 4C (details of bump near base of elytra of holotype, male), 4D (details of bump near base of elytra of paratype, female), 5A–F (holotype, male genitalia), 6A–C (paratype, female ovipositor), 7D (elytra of holotype, male), 23 (key) [2].

Distribution: China (Chongqing, Guizhou).

Graphidessa obliquefasciata Komiya and Kusama, 1974

Graphidessa obliquefasciata Komiya and Kusama, 1974: 138 (type locality: “Sungkang, Nantou Hsien, Taiwan, China”), pl. I [32]; Hua 1982: 88 (catalogue) [20]; Nakamura et al., 1992: 92 (catalogue) [21]; Hua 2002: 211 (catalogue) [22]; Chou 2004: 321, figs. male and female [23]; Löbl and Smetana 2010: 223 (catalogue) [10]; Nakamura et al., 2014: 155 (catalogue) [26]; Lin and Yang 2019: 260 (catalogue) [27]; Danilevsky 2020: 314 (catalogue) [13]; Liu et al., 2023: 23 (key) [2].

Distribution: China (Taiwan).

Graphidessa venata venataBates, 1884

Graphidessa venata Bates, 1884: 248 (type locality: “Higo, Kyushsu, Japan”) [1]; Aurivillius 1922: 321 (catalogue) [28]; Breuning 1963: 492 (catalogue) [9]; Breuning 1976: 75 (redescription) [30].

Graphidessa venata venata Ohbayashi and Niisato 2007: 618, pl. 67, fig. 6 (male) [31]; Löbl and Smetana 2010: 223 (catalogue) [10]; Danilevsky 2020: 314 (catalogue) [13]; Liu et al., 2023: 17, fig. 1 (distribution map), 22, fig. 7A (elytra of holotype, male), 23 (key) [2].

Type material examined: Holotype (MNHN): Graphidessa venata Bates (handwritten with black ink on a rectangular white label)/Higo (handwritten with black pencil on a rectangular white label)/venata Bates (handwritten with black ink on a rectangular white label with black borders and a transversal black line in middle)/MUSEUM PARIS COLL. H. W. BATES 1952 (printed with black ink on a rectangular white label with black borders)/TYPE (printed with black ink on a rectangular red label), examined from three photographs (Figure 3A–C).

Distribution: Japan (Kyushsu).

Graphidessa venata takakuwai Fujita, 1980

Graphidessa venata takakuwai Fujita, 1980: 15 (type locality: “Mikurajima Island, Izu Islands, Japan”), figs. 36 and 37 [33]; Ohbayashi and Niisato 2007: 618, pl. 67, fig. 7 (male) [31]; Löbl and Smetana 2010: 223 (catalogue) [10]; Danilevsky 2020: 314 (catalogue) [13]; Liu et al., 2023: 17, fig. 1 (distribution map), 22, fig. 7B (elytra of holotype, male), 23 (key) [2].

Distribution: Japan (Izu Islands).

Key to species of Graphidessa

• 1. Body dark reddish-brown or light red....................................................................................................................................................................................................................2
• - Body dark brown.........................................................................................................................................................................................................................................................3
• 2. Elytra sparsely covered with short yellowish-white hairs between middle and apical one-quarter...................................................................................G. venata venata
• - Elytra densely covered with short yellowish-white hairs between middle and apical one-quarter...............................................................................G. venata takakuwai
• 3. Post-basal bumps of elytra covered with long black setae......................................................................................................................................................G. obliquefasciata
• - Post-basal bumps of elytra covered with short black and long brown setae....................................................................................................................................G. jinfoensis

Paratimiola Breuning, 1965

Paratimiola Breuning, 1965: 34 [7]; Rondon and Breuning 1970: 493 (catalogue) [34]; Breuning 1975: 7 (key) [29]; Breuning 1976: 137 (redescription) [30]. Type species: Paratimiola rondoni Breuning, 1965, by original designation.

By examining different types of Tuberenes vietnamensis Breuning, 1972, we found that it does not belong to Tuberenes because it lacks several diagnostic characteristics of this genus: each elytron is widely rounded at the apex, neither covered with a small fascicle of setae behind the base nor covered with white pubescence forming several longitudinal lines at the apical half, and the femora is not covered with erect pubescence.

Comparative analyses of the types of Paratimiola rondoni (Figure 4A) and T. vietnamensis (Figure 4B,C,E,F) reveal that both species obviously belong to same genus based on the shared commonality of morphological characteristics: antennae distinctly longer than the body; thick scape; pedicle longer than width; antennomere III slightly shorter than the scape; antennomere IV distinctly longer than antennomere III; antennomere V distinctly shorter than antennomere IV; length of antennomeres V–XI gradually reduced; pronotum with a pair of triangular tubercles at the middle sides; scutellum linguiform; elytra wider than the pronotum at the base; sides of elytra sub-paralleled at the basal third, slightly expanded in the middle, gradually constricted from the apical third to the apex and widely rounded at the apices; disc sparsely covered with erected long black setae, densely covered with short white pubescence at the basal third, forming irregular spots, and densely covered with short white pubescence in the middle, forming an X-shaped band, with coarse and moderately dense punctuations from the base to the apical third (the punctations are sparse from the apical third to the apex); the mesocoxal cavities open externally to the mesepimera; the femora of the legs are strongly clavate; and the mesotibiae have an outer groove in front of the apex.

Paratimiola currently consists of two species.

Distribution: Laos, Vietnam.

Paratimiola rondoni Breuning, 1965

Paratimiola rondoni Breuning, 1965: 35 (type locality: “Phou Kow Khouei, près de Vientiane, Laos”) [7]; Rondon and Breuning 1970: 493 (catalogue), fig. 38 i [34]; Breuning 1976: 137 (redescription) [30].

Type material examined: Holotype (BPBM): Phou Kow Khouei region, a high-altitude station near Vientiane, 15 April 1965 (collecting data cited from Breuning 1965 [7]), examined from one photograph (Figure 4A).

Diagnosis: Paratimiola rondoni is very similar to P. vietnamensis (Breuning, 1972), as mentioned above, but can be clearly distinguished from P. vietnamensis by two diagnostic characteristics that are lacking in this species: the elytral base is not covered with short white pubescence, forming a V-shaped band surrounding the scutellum, and the apical third of each elytron is not covered with short white pubescence, forming a U-shaped band along the margin.

Distribution: Laos (Vientiane).

Paratimiola vietnamensis (Breuning, 1972) comb. nov.

Tuberoenes vietnamensis Breuning, 1972: 237 (type locality: “Xuan dinh, NW of Hanoi, Vietnam”) [35].

Tuberenes vietnamensis: Breuning 1978: 21 (redescription) [8].

Type material examined: Holotype (HNHM): VIETNAM: Xuan dinh NW of Hanoi 26-29.IV.1966 Bxp. Gy. TOPÁL (printed with black ink on a rectangular white label)/Tuberenes vietnamensis mihi Typ Breuning dét. (“Tuberenes vietnamensis mihi Typ” handwritten and “Breuning dét.” printed with black ink on a rectangular white label)/TYPE (printed with black ink on a rectangular red label)/Holotypus 1972. Tuberoenes vietnamensis Breuning (“Holotypus” printed with red ink and “1972. Tuberoenes vietnamensis Breuning” handwritten with black ink on a rectangular white label with red borders), examined from three photographs (Figure 4B–D). Paratype (HNHM): VIETNAM: Tanh liet SE of Hanoi 23.IV.1966 Bxp. Gy. TOPÁL (printed with black ink on a rectangular white label)/Nr. 140 beaten from trees (printed with black ink on a rectangular white label)/Tuberenes vietnamensis mihi Paratyp Breuning dét. (“Tuberenes vietnamensis mihi Paratyp” handwritten and “Breuning dét.” printed with black ink on a rectangular white label)/Paratypus 1972. Tuberoenes vietnamensis Breuning (“Paratypus” printed with red ink and “1972. Tuberoemes vietnamensis Breuning” handwritten with black ink on a rectangular white label with red borders), examined from three photographs (Figure 4E–G).

Distribution: Vietnam (Hanoi).

Tuberenes Breuning, 1978

Tuberoenes Breuning, 1972: 237 nomen nudum [35].

Tuberenes Breuning, 1978: 20 [8]; Löbl and Smetana 2010: 213 (catalogue) [10]; Lin and Yang 2019: 224 (catalogue) [27]; Danilevsky 2020: 299 (catalogue) [13]. Type species: Eryssamena robustipes Pic, 1939.

Breuning described Tuberoenes (sic) vietnamensis from Vietnam and compared it with robustipes Pic (this should be Eryssamena robustipes Pic, 1939, which is a type species of Tuberenes) [35], but there is no information on the genus Tuberoenes. Then, Breuning transferred Tuberoenes vietnamensis to Tuberenes when he first established Tuberenes [8]. While the Titan Database considers Tuberoenes a misspelling of Tuberenes [3], it is actually a nomen nudum following the ICZN, Art. 13.1.1 and Art. 13.3 [17].

Distribution: China, India.

Tuberenes robustipes (Pic, 1939)

Eryssamena robustipes Pic, 1939: 15 (type locality: “Kansou, Chine = Gansu, China”) [36]; Gressitt 1951: 523 (catalogue) [37]; Breuning 1963: 524 (catalogue) [9]; Hua 2002: 207 (catalogue) [22].

Tuberenes robustipes: Breuning 1978: 20 (redescription) [8]; Löbl and Smetana 2010: 213 (catalogue) [10]; Mitra et al., 2017: 67 (fauna) [38]; Lin and Yang 2019: 224 (catalogue) [27]; Danilevsky 2020: 299 (catalogue) [13].

Since Tuberenes minuta (Pic, 1925) and T. sikkimensis Breuning, 1978 have been transferred to Pararondibilis and T. vietnamensis Breuning, 1972 has been transferred to Paratimiola, Tuberenes now consists of only one species.

Type material examined: Holotype (MNHN): Chine Kansou (handwritten with black ink on a rectangular white label)/type (handwritten with black ink on a rectangular yellow label)/Eryssamena robustipes n sp (handwritten with black ink on a rectangular white label)/voisin de minuta (i.e., Eryssamena minuta Pic, 1925), scape plus robuste, cuisses plus fortes, pattes post [érieures] plus courtes (the contents mean “similar to minuta, more robust scape, stronger thighs shorter hind legs” and were handwritten with black ink on a rectangular white label)/élytres plus courts à fortes côtes latérales (“elytres” was handwritten with black ink at the end of the previous label; the contents mean “shorter elytra with strong lateral ribs” and were handwritten with black ink on a rectangular white label)/TYPE (printed with black ink on a rectangular red label)/Museum Paris Coll. M. Pic (printed with black ink on a rectangular white label with black borders), examined from three photographs (Figure 5A–C).

Distribution: China (Gansu), India (Sikkim, Darjeeling).