Heptophylla gongshana Bezděk & Král, n. sp. from China (Yunnan) and a Checklist of the Tribe Heptophyllini (Scarabaeidae: Melolonthinae) ^†

Abstract

Heptophylla gongshana Bezděk & Král, n. sp. from Yunnan, China, is described. The new species, conspicuous in the genus Heptophylla Motschulsky, 1858, by its black body and octomerous antennal club in males, is compared with similar congeners. An iconography including the male genitalia of the new species and the type material of the Heptophyllini taxa described by You-Wei Zhang and found in the Institute of Zoology, the Chinese Academy of Sciences, Beijing, China, is provided. An updated checklist of Heptophyllini (Scarabaeidae: Melolonthinae) is presented, and the validity of the tribe is discussed.

1. Introduction

The tribe Heptophyllini is one of the insufficiently known and also poorly defined groups of the Old World Melolonthinae (Coleoptera: Scarabaeidae). This tribe was proposed by Medvedev [1] to accommodate the following four genera: Heptophylla Motschulsky, 1858 (=Hypochrus Fairmaire, 1891), Hexataenius Fairmaire, 1891, Hilyotrogus Fairmaire, 1886, and Toxospathius Deyrolle & Fairmaire, 1878. Medvedev [1] pointed out that Heptophyllini occupies a certain intermediate position between the tribes Melolonthini and Rhizotrogini. However, this opinion was weakly supported by morphological data (e.g., general habitus and wing venation, as in Rhizotrogini, and a multimerous antennal club, as in Melolonthini). Kalinina [2] examined the larval stages of Hilyotrogus bicoloreus (Heyden, 1887) in detail and found no differences from the larvae of Rhizotrogini, but retained Heptophyllini as a valid tribe. Although weakly supported, the tribe Heptophyllini has been considered valid in the last 50 years. Two additional genera, Laotrichia Keith, 2007, and Taiwanotrichia Kobayashi, 1990, were described, and the total number of taxa assigned to Heptophyllini exceeded 50 (see the checklist below).

The genus Heptophylla was established by Motschulsky [3] for a single species, H. picea, which was described in the same work from Japan. The genus currently comprises eight valid taxa distributed in East and Northeast Asia: China, the Russian Far East, the Korean Peninsula, and Japan. With the exception of the quite common and widespread H. picea and its subspecies, the other Heptophylla species are only known from their primary description or from a very small number of localities.

During our (A.B. and D.K.) visit to several large Chinese institutional collections in 2024, we had the opportunity to study part of the type material of Heptophyllini described by You-Wei Zhang and deposited in the Institute of Zoology, the Chinese Academy of Sciences (IZAS), as well as interesting specimens from the collection of Hebei University Museum (HBUM), including the new Heptophylla species described below. We have also compiled an up-to-date checklist of taxa currently assigned to the Heptophyllini.

2. Materials and Methods

Photographs of the studied material were taken with a Canon MP-E 65 mm f/2.8 1–5× macro lens mounted on a Canon EOS 5DS R digital camera. Multiple partially focused images of each specimen were combined using Zerene Stacker (Zerene Systems LLC, Richland, WA, USA). All images were digitally processed and enhanced in Adobe Photoshop CC.

Specimens of the newly described species are provided with one red printed label “Heptophylla gongshana sp. nov.|HOLOTYPE, male [or] PARATYPE, male [or] PARATYPE, female|det. A. Bezděk & D. Král 2024”.

The exact label data are cited for the type material examined. Separate labels are indicated by a double vertical bar “||”, and lines within each label are separated by a single vertical bar “|”. Information in quotation marks indicates the original spelling. Our remarks and additional comments are placed in brackets: [p]—preceding data (within quotation marks) are printed; [hw]—the same but handwritten; HT—holotype; and PT—paratype.

The following codes identify the collections housing the material examined:

HBUM—Hebei University Museum, Baoding, Hebei, China (Xinglong Bai, Guodong Ren);

IZAS—the Institute of Zoology, the Chinese Academy of Sciences, Beijing, China (Ming Bai, Kuiyan Zhang);

NMPC—the National Museum, Prague, Czech Republic (Lukáš Sekerka, Jiří Hájek).

The updated checklist is based on the evaluation of data from the literature, including verification of all primary descriptions.

3. Results

3.1. Description of New Species

Heptophylla gongshana Bezděk & Král, n. sp. [贡山七鳃金龟].

Figure 1A,B, Figure 2A–F, and Figure 6A.

Zoobank:urn:lsid:zoobank.org:act:6AF08941-6804-4DB7-819D-D0E3C8352842.

Type locality. Yunnan, Gongshan, Dulongjiang township, N 27°50.422′ E 98°19.319′, 1445 m a.s.l.

Type material. China, Yunnan, HT (HBUM), male, labeled as follows: “2017-VII-26 | 云南贡山独龙江乡 | 白兴龙 魏中华 邵宪磊 | 河北大学博物馆 [p] [=2017-VII-26|Yunnan, Gongshan, Dulongjiang township | Xinglong Bai, Zhonghua Wei, Xianlei Shao lgt. | Hebei University Museum] || N 27°50.422′ | E 98°19.319′ | Alt. 1445 | 河北大学博物馆 [p] [=N 27°50.422′ | E 98°19.319′ | Alt. 1445 | Hebei University Museum]”. PT, three males and six females, with the same data; one male and four females in HBUM; one male and one female in NMPC; and one male and one female in IZAS.

Diagnosis. Body length: 13.6–14.7 mm. Elongated, expanded posteriad, and plump abdomen; almost all black-colored, shiny surface. Outline of clypeus is broadly trapezoidal, with considerably upturned margin, and anterior margin shallowly emarginate. Punctation of clypeus is remarkably coarse and dense, almost regularly distributed. Antennae are decamerous. Antennomere 2 is small, conical, and longer than wide; antennal club in males is octomerous, with basal mere shortened to half the length of next mere; club is almost straight in basal two-thirds, and apical third is moderately outcurved; and club is distinctly longer than antennal shaft. Antennal club in females is considerably smaller, consisting of six meres, gradually increasing in length toward apex. Pronotum is moderately convex, transversal, and broadest approximately at the base, except for broad basal interruption bordered all around; anterior bead is narrow; lateral margin is glabrous and bare; basal margin appears with a row of densely, almost regularly distributed punctures, bare; surface is bare, coarsely, densely, and almost regularly punctate, with punctures separated by their 1–2 diameters. Elytra is bare, absent from macrosetation. Macropterous. Protibia is sharply tridentate, basal tooth is smaller, and terminal calcar is slender, sharply acute apically, and inserted against emargination between basal and medial teeth. Calcars of metatibia in males are both simply shaped.

Description of the holotype, male. Body length: 14.4 mm. Body is elongated, with plump abdomen; all black except for considerably chestnut-colored penultimate and last abdominal segments (propygidium and pygidium); whole surface is strongly shiny; and considerably sparse and short macrosetation is yellowish–whitish (Figure 1A).

Head (Figure 1A). Small labrum, bilobed; lobes are rectangular apically; impunctate. Outline of clypeus is broadly trapezoidal, with considerably upturned margin, anterior margin shallowly emarginate, anterior angles rounded, and sides broadly arcuate. Genae are narrow, long, and rounded apically; partially dividing eyes appear dorsally. Frontoclypeal suture is almost straight, finely impressed. Eyes are relatively large; distance between eyes in ventral aspect remarkably exceeds diameter of eye. Punctation of clypeus is remarkably coarse and dense, almost regularly distributed, with punctures separated by approximately their diameter or less laterally and are rather confluent medially. Vertex is rather rugo-punctate, with punctures separated by less than their diameter to confluent.

Head appendages. Antennae are decamerous. Antennomere 2 is small, conical, and longer than wide; antennal club is octomerous, with basal mere shortened to half the length of next mere. Club is almost straight in basal two-thirds, and apical third is moderately outcurved; club is distinctly longer than antennal shaft (antennomeres 1–2 combined). Antennomeres 1–2 have sparse, long, whitish macrosetae; club is sparsely, shortly macrosetaceous, mainly in apical third (Figure 2A). Terminal maxillary palpomere is elongate, almost truncate apically, without dorsal depression, approximately as long as palpomeres 1 and 2 combined.

Pronotum is moderately convex, transversal, and broadest approximately at the base, except for broad basal interruption bordered all around; anterior bead is narrow; lateral margin is glabrous and bare; and basal margin has a row of densely, almost regularly distributed, punctures, bare. Anterior angles are prominent, projecting over anterior margin, acute-angled, with a rounded apex, bearing several long macrosetae; sides are shallowly bisinuate, divergent posteriad to very broadly obtuse posterior angles; and posterior margin is broadly rounded. Surface is bare, coarsely, densely, and almost regularly punctate, with punctures separated by their 1–2 diameters (Figure 1A).

Scutellar plate is approximately broader than long, triangulate, with broadly arcuate sides and acute apex; punctation is somewhat sparser than in pronotum (Figure 1A).

Elytra is convex, bare, and slightly dilated posteriad, with obtuse-angled sutural corner; striae, except for sutural stria, are missing or only very feebly indicated; punctation is coarse, dense, and almost regular, and punctures are separated by their 1–2 diameters, with moderately convex, impunctate sutural interval; sutural stria has a row of irregularly distributed punctures; and lateral margin is distinctly bordered, bare (Figure 1A).

Macropterous.

Legs are shiny, and surface is almost bare. Femora are very sparsely irregularly punctate, with punctures on posterior margin bearing long, erect macrosetae. Protibia is sharply tridentate, with smaller basal tooth and slender terminal calcar, sharply acute apically, inserted against emargination between basal and medial teeth. Meso- and metatibia only slightly expanded apically, with one setiferous transversal carina. Mesotibial terminal calcars are unequal in length, and acute apically. Metatibial terminal calcars are unequal in length, considerably flattened, and acute apically. Bifid claws (Figure 2C). Ventral surface of thorax has sparse, long, and recumbent macrosetation.

Pygidium is shiny, bordered all around, and sparsely, finely, and almost regularly punctate. Abdominal ventrites are almost bare, and finely and irregularly punctate.

Male genitalia. Aedeagus is symmetrical, parameres are slender and relatively long, distinctly longer than phallobasis, and distal part is not dilated in dorsal aspect, with rounded apex (Figure 2D–F).

Variability in males. Paratypes are somewhat variable in body length (13.6–14.7 mm), and slightly variable in punctation density of dorsal surface, length, and distribution of macrosetae. One paratype is dark brown with soft cuticle, evidently a teneral specimen.

Female (Figure 1B). Body length: 14.3–16.7 mm. Differs from males as follows: antennal club is considerably smaller, consisting of six meres, gradually increasing in length toward apex (Figure 2B).

Etymology. Species name refers to the type locality: Gongshan County in northwestern Yunnan.

Distribution. China: northwestern Yunnan.

Differential diagnosis. Heptophylla gongshana n. sp. differs from other known species of this genus in the octomerous antennal club in males and the unusual black color of the body. There is only one additional black-colored Heptophylla species, H. picea Motschulsky, 1858, known in Japan, the Russian Far East, and the Korean Peninsula (e.g., [4]). The nominotypical subspecies is usually brown with rare black forms, and H. p. iriei Kusui, 1971 (an endemic subspecies of a small Japanese island, Shimokoshiki-shima), is always entirely black-colored. These two species differ well in the shape of their bodies. Heptophylla picea is slender, nearly parallel-sided, with a heptamerous antennal club in males, while the body of H. gongshana n. sp. is stouter, distinctly expanded posteriad, with an octomerous antennal club in males. Because the generic classification of Heptophyllini is unstable, we compared the new species also with Hilyotrogus vietnamicus Keith, 2012, an entirely black-colored species described in the type locality “Hà Giang” in northern Vietnam, very close to the Vietnam–Yunnan border. Hilyotrogus vietnamicus shares with H. gongshana a rather stout shape of the body, but clearly differs in the pentamerous antennal club in males and also in the well-visible dense, pale setation on the pronotum, elytra, and abdominal ventrites (while in the new species, the pronotum and elytra are completely bare).

A personal study of the type material of Heptophyllini described by You-Wei Zhang and found in IZAS enabled us to take photographs of the relevant type specimens for the first time (see Figure 3A,B, Figure 4A–G, Figure 5A–E, and Figure 6B–D) and to tabulate differential characters from H. gongshana n. sp. (see

Table 1. Comparison of Heptophylla gongshana n. sp. with type material of Heptophyllini species found in IZAS.

Characters	Heptophylla gongshana n. sp.	Heptophylla calcarata	Heptophylla laticollis	Hilyotrogus flavescens
Color	Black, two apical abdominal segments chestnut brown, shiny (Figure 1A,B)	Pale, yellow elytra, head, pronotum, and scutellum pale brownish, matt (Figure 3A,B)	Brown, matt (Figure 5D)	Yellow, head brown, pronotum with two brown longitudinal spots, shiny (Figure 5A)
Shape of body	Elongate, convex, distinctly expanded posteriad (Figure 1A,B)	Elongate, convex, moderately expanded posteriad (Figure 3A,B)	Elongate, convex, parallel-sided (Figure 5D)	Elongate, flattened, parallel-sided (Figure 5A)
Setation of elytra	Absent (Figure 1A,B)	Absent (Figure 3A,B)	Absent (Figure 5D)	Dense, short, erect setae throughout (Figure 5A)
Clypeus	Transversal, slightly emarginate in middle, anterior angles broadly rounded, densely coarsely punctate (Figure 1A, B)	Transversal, slightly emarginate in middle, sides regularly rounded, densely coarsely punctate (Figure 3A,B)	Transversal, emarginate in middle, sides regularly rounded, densely coarsely punctate (Figure 5E)	Transversal, emarginate in middle, anterior angles broadly rounded, sparsely coarsely punctate (Figure 5A)
Male antennal club	Almost straight in basal two-thirds, apical third moderately outcurved, octomerous, basal mere distinctly shortened (Figure 2A)	Strongly curved outwardly, heptamerous, basal mere distinctly shortened (Figure 4A)	Moderately curved outwardly, heptamerous, basal mere only slightly shortened (Figure 5E)	Strongly curved outwardly, heptamerous, basal mere not shortened (Figure 5B)
Female antennal club	Straight, hexamerous, meres gradually increasing in length toward apex (Figure 2B)	Straight, hexamerous, meres gradually increasing in length toward apex (Figure 4B)	Female, unknown	Female, unknown
Pronotum	Transversal, anterior angles prominent, acute-angled (Figure 1A,B)	Transversal, anterior angles weak, acute-angled (Figure 3A,B)	Transversal, anterior angles prominent, rectangular (Figure 5E)	Transversal, anterior angles obtuse-angled (Figure 5A)
Male calcars of metatibia	Both shaped as usual	Lower calcar modified, broadened, and hooked apically (Figure 4C)	Both shaped as usual	Both shaped as usual

).

Figure 1. Dorsal habitus of Heptophylla gongshana n. sp. (A) Holotype, male; length: 14.4 mm. (B) Paratype, female; length: 14.9 mm. Not to scale. Photographs taken by Y.-P.L.

Figure 1. Dorsal habitus of Heptophylla gongshana n. sp. (A) Holotype, male; length: 14.4 mm. (B) Paratype, female; length: 14.9 mm. Not to scale. Photographs taken by Y.-P.L.

Figure 2. Heptophylla gongshana n. sp. (A,C–F) Holotype, male. (B) Paratype, female. (A) Left antenna, dorsal view. (B) Left antenna, dorsal view. (C) Claw of left hind leg. (D) Parameres, dorsal view. (E) Parameres, left lateral view. (F) Parameres, ventral view. Not to scale. Photographs taken by Y.-P.L.

Figure 2. Heptophylla gongshana n. sp. (A,C–F) Holotype, male. (B) Paratype, female. (A) Left antenna, dorsal view. (B) Left antenna, dorsal view. (C) Claw of left hind leg. (D) Parameres, dorsal view. (E) Parameres, left lateral view. (F) Parameres, ventral view. Not to scale. Photographs taken by Y.-P.L.

Figure 3. Dorsal habitus of Heptophylla calcarata Y.-W. Zhang, 1981. (A) Paratype, male; length: 14.2 mm. (B) Paratype, female; length: 14.9 mm. Not to scale. Photographs taken by Y.-P.L.

Figure 3. Dorsal habitus of Heptophylla calcarata Y.-W. Zhang, 1981. (A) Paratype, male; length: 14.2 mm. (B) Paratype, female; length: 14.9 mm. Not to scale. Photographs taken by Y.-P.L.

Figure 4. Heptophylla calcarata Y.-W. Zhang, 1981. (A,C–G) Paratype, male. (B) Paratype, female. (A) Left antenna, dorsal view. (B) Left antenna, dorsal view. (C) Detail with modified calcar of left metatibia. (D) Claw of left middle leg. (E) Parameres, dorsal view. (F) Parameres, left lateral view. (G) Parameres, ventral view. Not to scale. Photographs taken by Y.-P.L.

Figure 4. Heptophylla calcarata Y.-W. Zhang, 1981. (A,C–G) Paratype, male. (B) Paratype, female. (A) Left antenna, dorsal view. (B) Left antenna, dorsal view. (C) Detail with modified calcar of left metatibia. (D) Claw of left middle leg. (E) Parameres, dorsal view. (F) Parameres, left lateral view. (G) Parameres, ventral view. Not to scale. Photographs taken by Y.-P.L.

Figure 5. Hilyotrogus flavescens (Y.-W. Zhang, 1997), holotype, male; length: 13.8 mm (A–C). Heptophylla laticollis Y.-W. Zhang, 1995, holotype, male; length: 13.5 mm (D–F). (A,D) Dorsal habitus. (B) Right antenna, dorsal view. (E) Part of head with right antenna, dorsal view. (C,F) Claw of right middle leg. Not to scale. Photographs taken by Y.-P.L.

Figure 5. Hilyotrogus flavescens (Y.-W. Zhang, 1997), holotype, male; length: 13.8 mm (A–C). Heptophylla laticollis Y.-W. Zhang, 1995, holotype, male; length: 13.5 mm (D–F). (A,D) Dorsal habitus. (B) Right antenna, dorsal view. (E) Part of head with right antenna, dorsal view. (C,F) Claw of right middle leg. Not to scale. Photographs taken by Y.-P.L.

Figure 6. Associated labels. (A) Heptophylla gongshana n. sp., holotype. (B) H. calcarata Y.-W. Zhang, 1981, paratype. (C) Hilyotrogus flavescens (Y.-W. Zhang, 1997), holotype. (D) Heptophylla laticollis Y.-W. Zhang, 1995, holotype. Photographs taken by Y.-P.L.

Figure 6. Associated labels. (A) Heptophylla gongshana n. sp., holotype. (B) H. calcarata Y.-W. Zhang, 1981, paratype. (C) Hilyotrogus flavescens (Y.-W. Zhang, 1997), holotype. (D) Heptophylla laticollis Y.-W. Zhang, 1995, holotype. Photographs taken by Y.-P.L.

3.2. Type Material of Heptophyllini Found in IZAS and Used for Comparison (See Table 1)

Heptophylla calcarata Y.-W. Zhang, 1981 [5] [距七鳃金龟].

Figure 3A,B, Figure 4A–G, and Figure 6B.

Type material examined (20 paratypes). China, Xizang: PT, male, labeled as follows: “西藏 [p] 吉隆 [h] | 中国科学院 | [p] [=Xizang, Jilong | Chinese Academy of Sciences] || 1978. VI. 14. | 324 [h] | 采集者: [p] 农科所 [=日喀则农科所] [h] [=collector: Institute of Agricultural Sciences [=Shigatse Institute of Agricultural Sciences]] || 32004 | Inst. Zool. | Acad. Sin. [p] || IOZ (E) 218297 [p]”. PT, four males and three females, with the same collection data; PT, nine males and three females, with the same collection data, but with the following modification: “1978. VI. 9. | 2750 m [h]”.

 

Heptophylla laticollis Y.-W. Zhang, 1995 [6] [阔背七鳃金龟].

Figure 5D–F and Figure 6D.

Type material examined (holotype and three paratypes). China, Zhejiang: HT, male, labeled as follows: “浙江庆元百山祖 [p] [=Zhejiang, Qingyuan, Baishanzu] | 1993—[p] VIII. 19. [h] | 吴鸿 [p] [=Hong Wu lgt.] 1600 m [h]”. PT, two males, with the same collection data, but with the following modification: “1993—[p] VIII. 22. [h] | 吴鸿 [p] [=Hong Wu lgt.] 1500 m [h]”; PT, male, with the same collection data, but with the following modification: “1993—[p] VIII. 22. [h] | 林敏 [p] [=Lin Min lgt.] 1500 m [h]”.

 

Hilyotrogus flavescens (Y.-W. Zhang in Y.-W. Zhang & Y. Li, 1997) [7] [淡黄希鳃金龟].

Figure 5A–C and Figure 6C.

Type material examined (holotype). China, Sichuan, HT, male, labeled as follows: “四川巫山梨子坪 [=Sichuan, Wushan, Liziping] | 1850 m | 中国科学院 [p] [=Chinese Academy of Sciences] || 1994. V. 19 | 采集人李文柱 [p] [=Wenzhu Li lgt.]”.

3.3. Checklist of Genera and Species Currently Classified in Heptophyllini—Distribution Is Derived from the Primary Descriptions and Bezděk [4], Unless Otherwise Stated

Genus Heptophylla Motschulsky, 1858 [3] [七鳃金龟属].

Heptophylla Motschulsky, 1858: 32.

Hypochrus Fairmaire, 1891: cci [8]. Synonymized by Medvedev (1951: 202) [1].

 

Heptophylla brevicollis (Fairmaire, 1891) [8] [短七鳃金龟].

Hypochrus brevicollis Fairmaire, 1891: cci.

Heptophylla brevicollis: Medvedev 1951: 204 [1] (new combination).

Type locality. “Tchang-Yang”.

Distribution. China (Hubei).

 

Heptophylla calcarata Y.-W. Zhang, 1981 [5] [距七鳃金龟].

Heptophylla calcarata Y.-W. Zhang, 1981: 349.

Type locality. “Xizang: Gyirong, 2800 m”.

Distribution. China (Xizang).

 

Heptophylla dimidiata Arrow, 1934 [9] [二色七鳃金龟].

Heptophylla dimidiata Arrow, 1934: 9.

Type locality. “Kiangsu”.

Distribution. China (Jiangxi).

 

Heptophylla laticollis Y.-W. Zhang, 1995 [6] [阔背七鳃金龟].

Heptophylla laticollis Y.-W. Zhang, 1995: 243.

Type locality. “Mt. Baishanzu, 1600 m, Zhejiang Prov.”.

Distribution. China (Zhejiang).

 

Heptophylla longilamella Y.-W. Zhang, 1981 [5] [长角七鳃金龟].

Heptophylla longilamella Y.-W. Zhang, 1981: 350.

Type locality. “Xizang: Cona”.

Distribution. China (Xizang).

 

Heptophylla picea iriei Kusui, 1971 [10].

Heptophylla picea iriei Kusui, 1971: 117.

Type locality. “Aose, Shimokoshiki Is.”.

Distribution. Japan (Shimo-koshiki-shima).

 

Heptophylla picea maenamii Nomura, 1969 [11].

Heptophylla picea maenamii Nomura, 1969: 73.

Type locality. “Toshima, Myiakejima, Mikurajima”.

Distribution. Japan (Izu-shotō).

 

Heptophylla picea picea Motschulsky, 1858 [3].

Heptophylla picea picea Motschulsky, 1858: 33.

Heptophylla tosana Miyatake, 1963: 23 [12]. Type locality. “Kochi pref. (Tosa province), Shikoku, Japan”. Synonymized by Fujioka (2001: 89) [13].

Holotrichia transversa Motschulsky, 1861: 15 [14]. Type locality. “Japon”. Synonymized by Lewis (1895: 399) [15].

Type locality. “Japon”.

Distribution. Japan, Russia (Far East), and South Korea.

 

Genus Hexataenius Fairmaire, 1891 [16] [六鳃金龟属].

Hexataenius Fairmaire, 1891: ix.

 

Hexataenius ezakii Niijima & Kinoshita, 1923 [17] [江崎六鳃金龟].

Gymnogaster Ezakii Niijima & Kinoshita, 1923: 70.

Hexataenius esakii: Nomura 1970: 69 [18] (incorrect subsequent spelling, new combination).

Type locality. “Formosa: Urai”.

Distribution. China (Taiwan).

 

Hexataenius protensus Fairmaire, 1891 [16] [展六鳃金龟].

Hexataenius protensus Fairmaire, 1891: ix.

Type locality. “Tchang-Yang”.

Distribution. China (Chongqing, Hubei, Sichuan) and Japan.

 

genus Hilyotrogus Fairmaire, 1886 [19] [希鳃金龟属].

Hilyotrogus Fairmaire, 1886: 325.

Melichrus Brenske, 1892: 156 [20]. Synonymized by Arrow (1944: 633) [21].

 

Hilyotrogus assamensis Moser, 1913 [22].

Hilyotrogus assamensis Moser, 1913: 278.

Type locality. “Assam”.

Distribution. India (Assam, Sikkim).

 

Hilyotrogus aurosericeus (Brenske, 1892) [23].

Holotrichia aurosericea Brenske, 1892: 172.

Hilyotrogus aurosericeus: Moser 1913: 277 [22] (new combination).

Type locality. “Tenasserim”.

Distribution. Myanmar.

 

Hilyotrogus bicoloreus (Heyden, 1887) [24] [二色希鳃金龟].

Lachnosterna (Holotrichia) bicolorea Heyden, 1887: 265.

Hilyotrogus bicolor: Brenske 1892: 154 [20] (incorrect subsequent spelling, new combination).

Type locality. “Korea”.

Distribution. Widely distributed in northern and northeastern China, the Korean Peninsula, and Russian Far East ([1,25,26]).

 

Hilyotrogus birmanicus Arrow, 1946 [27].

Hilyotrogus birmanicus Arrow, 1946: 30.

Type locality. “Kambaiti, 7000 ft.”.

Distribution. Myanmar.

 

Hilyotrogus calcaratus (Frey, 1971) [28].

Melichrus calcaratus Frey, 1971: 132.

Hilyotrogus calcaratus: Frey 1975: 236 [29] (new combination).

Type locality. “Nepal, East Junbesi, 2780 m und Nepal Dudh Kosi”.

Distribution. Nepal.

 

Hilyotrogus changbaishanensis J.-K. Li, 1992 [30] [长白希鳃金龟].

Hilyotrogus changbaishanensis J.-K. Li, 1992: 77.

Type locality. “Jilin, Changbai Shan, 700 m”.

Distribution. China (Jilin).

 

Hilyotrogus cribratulus Fairmaire, 1893 [31].

Hilyotrogus cribratulus Fairmaire, 1893: 308.

Type locality. “Haut Tonkin, 6 kilomètres autour du poste de Ha-lâng, situé dan sun région montagneuse, versant occidental de la chaîne du Yunnan”.

Distribution. Vietnam.

 

Hilyotrogus flavescens (Y.-W. Zhang in Y.-W. Zhang & Y. Li, 1997) [7] [淡黄希鳃金龟].

Melichrus flavescens Y.-W. Zhang in Y.-W. Zhang & Y. Li, 1997: 770.

Hilyotrogus flavescens: Bezděk 2006: 183 [32] (new combination).

Type locality. “Liziping, Wushan County, Sichuan province, 1850 m”.

Distribution. China (Chongqing).

 

Hilyotrogus formosanus (Niijima & Kinoshita, 1927) [33] [台湾希鳃金龟].

Heptophylla formosana Niijima & Kinoshita, 1927: 223.

Hilyotrogus formosanus: Arrow 1944: 633 [21] (new combination).

Type locality. “Formosa, Urai”.

Distribution. China (Taiwan).

 

Hilyotrogus hirsutissimus Keith, 2004 [34].

Hilyotrogus hirsutissimus Keith, 2004: 727.

Type locality. “Myanmar, 65 km NW de Putao, Zi Yar Dam, 1250 m”.

Distribution. Myanmar.

 

Hilyotrogus holosericeus (Kollar & Redtenbacher, 1844) [35].

Ancylonycha holosericea Kollar & Redtenbacher, 1844: 524.

Hilyotrogus holosericeus: Brenske 1892: 155 [20] (new combination).

Hilyotrogus rufoflavus Moser, 1921: 65 [36]. Type locality. “Sikkim”. Synonymized by Arrow (1944: 633) [21].

Lachnosterna stridulans Sharp, 1878: 172 [37]. Type locality. “Murree”. Synonymized by Arrow (1944: 633) [21].

Type locality. “Himalaya, Massuri”.

Distribution. India (Himachal Pradesh, Sikkim), Nepal, and Pakistan ([38]).

 

Hilyotrogus iridipennis Fairmaire, 1886 [19] [红翅希鳃金龟].

Hilyotrogus iridipennis Fairmaire, 1886: 326.

Type locality. “Yunnan”.

Distribution. China (Yunnan).

 

Hilyotrogus kolbei (Brenske, 1892) [20].

Melichrus Kolbei Brenske, 1892: 156.

Hilyotrogus kolbei: Frey 1975: 236 [29] (new combination).

Type locality. “Darjeeling”.

Distribution. Bhutan and India (West Bengal).

 

Hilyotrogus longiclavis Bates, 1891 [39] [长角希鳃金龟].

Hilyotrogus longiclavis Bates, 1891: 74.

Type locality. “Sze-chuen”.

Distribution. China (Gansu, Sichuan, Xizang) and Myanmar ([27]).

 

Hilyotrogus luteosericeus (Brenske, 1897) [40].

Holotrichia luteosericea Brenske, 1897: 347.

Hilyotrogus luteosericeus: Moser 1913: 277 [22] (new combination).

Type locality. “Kurseong”.

Distribution. India (West Bengal).

 

Hilyotrogus mangkamensis Y.-W. Zhang, 1981 [5] [芒康希鳃金龟].

Hilyotrogus mangkamensis Y.-W. Zhang. 1981: 351.

Type locality. “Xizang: Mangkam; 2600 m”.

Distribution. China (Xizang).

 

Hilyotrogus nitens Frey, 1975 [29].

Hilyotrogus nitens Frey, 1975: 235.

Type locality. “Gogona; 3000 m; Bhutan”.

Distribution. Bhutan.

 

Hilyotrogus ochraceosericeus Moser, 1913 [22].

Hilyotrogus ochraceosericeus Moser, 1913: 279.

Type locality. “Assam (Shillong)”.

Distribution. India (Assam).

 

Hilyotrogus piceosericeus Moser, 1913 [22] [褐丝希鳃金龟].

Hilyotrogus piceosericeus Moser, 1913: 281.

Type locality. “Yunnan”.

Distribution. China (Yunnan).

 

Hilyotrogus pilicollis Moser, 1913 [22] [毛希鳃金龟].

Hilyotrogus pilicollis Moser, 1913: 280.

Type locality. “Yunnan (Tali)”.

Distribution. China (Yunnan).

 

Hilyotrogus pilifer Moser, 1921 [36] [胶州希鳃金龟].

Hilyotrogus pilifer Moser, 1921: 66.

Type locality. “China (Kiautschau)”.

Distribution. China (Shandong).

 

Hilyotrogus rufosericeus Moser, 1913 [22].

Hilyotrogus rufosericeus Moser, 1913: 279.

Type locality. “Assam (Shillong)”.

Distribution. India (Assam).

 

Hilyotrogus sericeus Moser, 1915 [41] [丝光希鳃金龟].

Hilyotrogus sericeus Moser, 1915: 144.

Type locality. “China (Szetshwan)”.

Distribution. China (Sichuan).

 

Hilyotrogus setiger Moser, 1915 [41] [刚毛希鳃金龟].

Hilyotrogus setiger Moser, 1915: 145.

Type locality. “China (Kiukiang; Kiautschau)”.

Distribution. China (Shandong).

 

Hilyotrogus setipennis Moser, 1913 [22] [鬃翅希鳃金龟].

Hilyotrogus setipennis Moser, 1913: 281.

Type locality. “China (Tchao Pin-Jo)”.

Distribution. China (“Tchao Pin-Jo”).

 

Hilyotrogus sikkimensis Moser, 1913 [22].

Hilyotrogus sikkimensis Moser, 1913: 277.

Type locality. “Sikkim”.

Distribution. India (Sikkim).

 

Hilyotrogus stolidus Fairmaire, 1886 [19] [云南希鳃金龟].

Hilyotrogus stolidus Fairmaire, 1886: 326.

Type locality. “Yunnan”.

Distribution. China (Yunnan).

 

Hilyotrogus subsericeus (Moser, 1908) [42].

Holotrichia subsericea Moser, 1908: 341.

Hilyotrogus subsericeus: Moser, 1913: 277 [22] (new combination).

Type locality. “Montes Mauson”.

Distribution. Vietnam.

 

Hilyotrogus unguicularis Fairmaire, 1886 [19] [爪希鳃金龟].

Hilyotrogus unguicularis Fairmaire, 1886: 325.

Type locality. “Yunnan”.

Distribution. China (Yunnan).

 

Hilyotrogus vietnamicus Keith, 2012 [43].

Hilyotrogus vietnamicus Keith, 2012: 333.

Type locality. “Vietnam; Ha Giang”.

Distribution. Vietnam.

 

Hilyotrogus wittmeri Frey, 1975 [29].

Hilyotrogus wittmeri Frey, 1975: 233.

Type locality. “Bhutan; Nobding”.

Distribution. Bhutan.

 

Hilyotrogus yasuii (Nomura, 1970) [18].

Brahmina yasuii Nomura, 1970: 68.

Hilyotrogus yasuii: Matsumoto 2007: 34 [44] (new combination).

Type locality. “Mt. Hateruma; Iriomote Is.”.

Distribution. Japan (Iriomote-shima).

 

Genus Laotrichia Keith, 2007 [45] [老鳃金龟属].

Laotrichia Keith, 2007: 297.

 

Laotrichia cuccodoroi Keith, 2007 [45] [库氏老鳃金龟].

Laotrichia cuccodoroi Keith, 2007: 298.

Type locality. “Laos; Pahia”.

Distribution. China (Yunnan) ([46]), India (Assam), Laos, and Thailand ([45,47]).

 

Genus Taiwanotrichia Kobayashi, 1990 [48] [金背鳃金龟属].

Taiwanotrichia Kobayashi, 1990: 76.

 

Taiwanotrichia dorsopilosa C.-L. Li & Yang, 1991 [49] [细毛金背鳃金龟].

Taiwanotrichia dorsopilosa C.-L. Li & Yang, 1991: 320.

Type locality. “Hsin-Bai-Yang; Hualien Hsien”.

Distribution. China (Taiwan).

 

Taiwanotrichia hainanensis Keith & J.-K. Li, 2009 [50] [海南金背鳃金龟].

Taiwanotrichia hainanensis Keith & J.-K. Li, 2009: 164.

Type locality. “Chine; Hainan; mont Wuzhishan”.

Distribution. China (Hainan).

 

Taiwanotrichia longicornis Kobayashi, 1990 [48] [长角金背鳃金龟].

Taiwanotrichia longicornis Kobayashi, 1990: 76.

Type locality. “Bandar; near Wushe; Nantou Hsien”.

Distribution. China (Taiwan).

 

Taiwanotrichia similis C.-L. Li & Yang, 1991 [49] [鞍马山金背鳃金龟].

Taiwanotrichia similis C.-L. Li & Yang, 1991: 323.

Type locality. “An Ma Mt.; Taichung Hsien”.

Distribution. China (Taiwan).

 

Taiwanotrichia sinocontinentalis fujianensis Keith, 2009 [51] [中华金背鳃金龟福建亚种].

Taiwanotrichia sinocontinentalis fujianensis Keith, 2009: 234.

Tye locality. “Chine, Fujian, 50 km SE Yingtan, Wuyishan”.

Distribution. China (Fujian).

 

Taiwanotrichia sinocontinentalis sinocontinentalis Keith, 2009 [51] [中华金背鳃金龟指名亚种]

Taiwanotrichia sinocontinentalis sinocontinentalis Keith, 2009: 233.

Type locality. “Chine, Guangxi, Gongcheng, Dayaoshan”.

Distribution. China (Guangxi).

 

Genus Toxospathius Deyrolle & Fairmaire, 1878 [52] [弓角鳃金龟属].

Toxospathius Deyrolle & Fairmaire, 1878: 102.

 

Toxospathius auriventris Bates, 1891 [39] [丽腹弓角鳃金龟].

Toxospathius auriventris Bates, 1891: 73.

Type locality: “Ta-tsien-lu”.

Distribution. China (Sichuan, Xizang).

 

Toxospathius brevicollis Arrow, 1934 [9] [短弓角鳃金龟].

Toxospathius brevicollis Arrow, 1934: 8.

Type locality. “S. Kansu”.

Distribution. China (Gansu, Sichuan).

 

Toxospathius inconstans Fairmaire in Deyrolle & Fairmaire, 1878 [52] [变弓角鳃金龟].

Toxospathius inconstans Fairmaire in Deyrolle & Fairmaire, 1878: 103.

Type locality. “Chine centrale”.

Distribution. Northwestern and central China ([1]).

 

Remark on the checklist. Bezděk [4] included in Heptophyllini also Hilyotrogus flavosericeus (Brenske, 1897) from Sikkim, India. This species was originally described in the genus Holotrichia and later transferred to Hilyotrogus by Moser [22]. Nevertheless, based on its trimerous antennal club only, Frey [29] finally transferred it to Brahmina (tribe Rhizotrogini).

4. Discussion

As stated above, the tribe Heptophyllini is only weakly supported by morphological characters ([1,2]). The most commonly used character to distinguish Heptophyllini from Rhizotrogini is a multimerous antennal club with 5–7 meres in Heptohyllini and usually only 3 in Rhizotrogini. Nevertheless, the number of meres of the antennal club is rather inconsistent, because there are a few genera traditionally classified in Rhizotrogini with a multimerous antennal club also (e.g., Megistophylla Burmeister, 1855). Similarly, most genera of Sericini have only trimerous clubs, but with some conspicuous exceptions, like Tetraserica Ahrens, 2004, with a tetramerous club in males, or Microserica Brenske, 1894, with 4–7 meres in the antennal club of males.

None of the species currently classified in Heptophyllini has been included in major phylogenetic studies of Melolonthinae (e.g., [53,54]), and, in general, a comprehensive phylogenetic study of Rhizotrogini and Melolonthini is urgently needed. For this reason, the validity of the tribe Heptophyllini has not yet been clarified.

We hope that the description of the new species of Heptophylla and the checklist of species currently placed in Heptophyllini will draw attention to this group of chafers and encourage specialists in the taxonomy and phylogeny of Melolonthinae to study them.