Research Article |
Corresponding author: Chi-Feng Lee ( chifeng@tari.gov.tw ) Academic editor: Astrid Eben
© 2022 Chi-Feng Lee.
This is an open access article distributed under the terms of the Creative Commons Attribution License (CC BY 4.0), which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited.
Citation:
Lee C-F (2022) Revision of the genus Arthrotus Motschulsky, 1858 (Coleoptera, Chrysomelidae, Galerucinae) of Taiwan, with notes on color polymorphism. ZooKeys 1091: 161-208. https://doi.org/10.3897/zookeys.1091.79486
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Seven species of Arthrotus are recognized and redescribed: A. abdominalis (Chûjô, 1962), A. gressitti Kimoto, 1969, A. hirashimai Kimoto, 1969, A. fulvus Chûjô, 1938, A. saigusai Kimoto, 1969, A. tricolor (Chûjô, 1965), and A. testaceus Gressitt & Kimoto, 1963. Also, two new species are described: A. yuae sp. nov. and A. yangi sp. nov. Three new synonyms are proposed: Proegmena taiwana Takizawa, 1978 syn. nov., Dercetina nakanei Kimoto, 1969 syn. nov. and A. shibatai Kimoto, 1984 syn. nov. Lectotypes are designated for A. fulvus Chûjô, 1938 and Dercestra abdominalis Chûjô, 1962. Color polymorphism of each species is delimited base on more than 1800 specimens.
Dercetina, food plant, leaf beetles, new species, new synonym, nomenclature, taxonomy
The genus Arthrotus Motschulsky, 1858 includes 48 species from the Palaearctic and Oriental regions (
Reference | New species, new records, or nomenclatural acts |
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A. fulvus |
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Dercestra abdominalis |
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Dercetis tricolor |
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A. abdominalis (Chûjô, 1962) comb. nov. (transferred from Dercestra) |
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A. gressitti, A. hirashimai, A. saigusai, Dercetina nakanei; A. testaceus Gressitti & Kimoto, 1963 (new record); A. tricolor (Chûjô, 1965) comb. nov. (transferred from Dercetis) |
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Proegmena taiwana |
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A. shibatai |
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A. taiwanus (Takizawa, 1978) comb. nov. (transferred from Proegmena) |
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A. nakanei (Kimoto, 1969) comb. nov. (transferred from Dercetina) |
Arthrotus Motschulsky is similar to Dercetina Gressitt & Kimoto, 1963, with the following combination of shared characters: pronotum usually with one pair of lateral depressions, basal margin entirely marginate; closed anterior coxal cavity; elytra without setae; tibia of hind leg without one apical spine; first tarsomere of hind leg usually shorter than or subequal to combination of the rest; tarsal claws appendiculate. These genera differ from each other only by the structure of the male antennae: antennomere III is approximately twice as long as antennomere II in Dercetina, while antennomeres II and III are subequal in length in Arthrotus.
Most members of Arthrotus have similar shapes of male aedeagi and great color variation. Species boundaries are hard to determine without sufficient material. Fortunately, adults are collected easily by sweeping. More than 1800 specimens are available for study thanks to collecting efforts by members of the Taiwan Chrysomelid Research Team (TCRT), and borrowed material from several museums (see below).
For taxonomic study, the abdomens of adults were separated from the forebodies and boiled in 10% KOH solution, followed by washing in distilled water to prepare genitalia for illustrations. The genitalia were then dissected from the abdomens, mounted on slides in glycerin, and studied and drawn using a Leica M165 stereomicroscope. For detailed examination, a Nikon ECLIPSE 50i microscope was used.
At least three sex pairs from each species were examined to delimit variability of diagnostic characters. For species collected from more than one locality or with color variations, at least one sex pair of specimens from each locality and color morph was examined. Length was measured from the anterior margin of the eye to the elytral apex, and width at the greatest width of the elytra.
Specimens studied herein are deposited at the following institutes and collections:
HTC Haruo Takizawa private collection;
NMNS National Museum of Natural Science, Taichung, Taiwan [Jing-Fu Tsai];
Precise label data are cited for all type specimens of described species; a double slash (//) divides the data on different labels and a single slash (/) divides the data in different rows. Other comments and remarks are in square brackets: [p] – preceding data are printed, [h] – preceding data are handwritten, [w] – white label, [y] – yellow label, [b] – blue label, and [r] – red label.
Dercetis metallica:
Dercestra abdominalis
Chûjô, 1962: 166;
Arthrotus abdominalis: Kimoto, 1965 (transferred from Decesta);
Lectotype
♂ (
A total of 316 specimens was examined (Suppl. material
Adults of Arthrotus abdominalis (Chûjô) (Fig.
Color metallic blue, antennae and legs black, abdomen yellow (Fig.
Male. Length 6.2–7.6 mm, width 3.2–4.5 mm. Antennae filiform (Fig.
Female. Length 7.8–9.2 mm, width 3.9–5.1 mm. Antennae similar to those of males, but antennomere III slightly longer than II in females (Fig.
Adults feed on leaves of Quercus glauca var. glauca Thunb. (Fagaceae), Hydrangea chinensis Maxim. (Hydrangeaceae), Persicaria chinensis (L.) H. Gross (Polygonaceae), Prunus campanulata Maxim. (Rosaceae), Zelkova serrata (Thunb.) Makino (Ulmaceae), and Debregeasia orientalis C.J. Chen (Urticaceae).
Arthrotus gressitti Kimoto, 1969: 61.
Paratypes. 2♀ (
(n = 21). 1♂, 1♀ (
Adults of Arthrotus gressitti Kimoto (Fig.
Color blackish brown, elytra metallic blue, abdomen yellow (Fig.
Male. Length 5.9–6.6 mm, width 2.8–3.0 mm. Antennae filiform (Fig.
Female. Length 7.4–7.9 mm, width 3.6 mm. Antennae (Fig.
Unknown.
Adults have been collected from several localities of central Taiwan, including Pasienshan (八仙山) in Taichung county; Peitungyanshan (北東眼山), Lushan (廬山), and Nanshanchi (南山溪) in north Nantou county (Fig.
Arthrotus hirashimai Kimoto, 1969: 60.
Proegmena taiwana Takizawa, 1978: 125. syn. nov.
Arthrotus taiwana:
Arthrotus hirashimai. Holotype ♂ (
Proegmena taiwana. The male holotype and one male paratype should be deposited at the Hokkaido University but were not found (Takemoto pers. comm., 23 Sept 2021). Two paratypes were deposited at the Takizawa’s private collection. 1♀ (HTC): “Chitou (溪頭) Chu- / shan (竹山) Taiwan / 6–7.VII.1975 / H. Takizawa [p, w] // Paratype [h, red letter] // Proegmena / taiwana n. sp. [h] / para-T [h, red letters] 197[p]7.2[h] / Det. H. Takizawa [p, w] (on the back of the same card)”; 1♂ (HTC): “(male aedeagus glued on the card) // Chitou (溪頭) Chu- / shan (竹山) Taiwan / 6–7.VII.1975 / H. Takizawa [p, w] // PARATYPE (?) / Proegmena / taiwana [h, w] // 2021.X.15 / H. Takizawa / det. [p, w] (on the back of the same card). However, both paratypes should be females based on the original description (
(n = 40). Taiwan. Chiayi: 3♂, 1♀ (
Adults of Arthrotus hirashimai Kimoto (Fig.
Body color yellowish brown, elytra metallic blue, antennae black, vertex darkened in most of individuals (Fig.
Male. Length 5.6–6.4 mm, width 2.6–2.7 mm. Antennae filiform (Fig.
Female. Length 6.1–7.2 mm, width 2.8–3.4 mm. Antennae (Fig.
Unknown.
Adults are restricted to several localities at mid-elevations of central Taiwan, including Hsitou (溪頭), Tungfu (同富), and Tungpu (東埔) in south Nantou county; Alishan (阿里山), Fenchihu (奮起湖), and Tutzuhu trail (杜仔湖步道) in Chiayi county (Fig.
(n = 160). Holotype ♂ (
Adults of Arthrotus yuae sp. nov. (Fig.
Color (Fig.
Male. Length 5.7–7.0 mm, width 2.8–3.4 mm. Antennae filiform (Fig.
Female. Length 6.0–7.7 mm, width 3.3–3.9 mm. Antennae much shorter than in males, antennomere III a little longer than II (Fig.
Leaves of Achyranthes bidentata Blume (Amaranthaceae), and Prunus phaeosticta var. phaeosticta (Hance) Maxim. (Rosaceae).
Adults are restricted to several localities at lowlands of southern Taiwan including Chungchihkuan (中之關), Shinanshan (溪南山), Tona trail (多納林道), and Wutai (霧台) in Kaohsiung county; Tahanshan (大漢山) in Pingtung county; Lichia trail (利嘉林道), Liyuan (栗園), Tulanshan (都蘭山), and Yanping trail (延平林道) in Taitung county (Fig.
Dedicated to Mrs Su-Fang Yu (余素芳) who was the first member of TCRT to collect specimens of this new species.
Arthrotus fulvus
Chûjô, 1938: 139;
Arthrotus testaceus:
Dercetina nakanei Kimoto, 1969: 65. syn. nov.
Arthrotus nakanei:
Arthrotus fulvus. Lectotype ♀ (
Dercetina nakanei. Types were studied by
A total of 603 specimens was examined (Suppl. material
Adults of Arthrotus fulvus Chûjô are similar to those of A. tricolor (Chûjô) in possessing rounded lateral margins of their pronota (straight lateral margins in A. abdominalis (Chûjô), A. gressitti Kimoto, A. hirashimai Kimoto, and A. yuae sp. nov.), the less transverse pronotum and elytra, 1.7–2.0 × wider and long in pronotum and 1.5–1.6 × longer than wide in elytra (more transverse pronotum and elytra, 2.1–2.2 × wider and long and 1.4 × longer than wide in elytra of A. testaceus Gressitt & Kimoto and A. yangi sp. nov.), the more transverse antennomere III in males, 0.7–0.8 × longer than wide (Fig.
Habitus of Arthrotus fulvus Chûjô A collected from Kuanwu (觀霧), male, dorsal view B collected from Kuanwu (觀霧), male, ventral view C collected from Tzuchung (自忠), male, dorsal view D collected from Tungfu (同富), male, dorsal view E collected from Motien (摩天), female, dorsal view F collected from Motien (摩天), male, dorsal view.
Some color patterns characteristic and restricted to particular areas. Form A (described as A. nakanei) (Fig.
Habitus of Arthrotus fulvus Chûjô A collected from Liyuan (栗園), male, dorsal view B collected from Liyuan (栗園), male, ventral view C collected from Tengchih (藤枝), male, dorsal view D collected from Tahanshan (大漢山), male, dorsal view E collected from Tahanshan (大漢山), male, dorsal view F collected from Erhchituan (二集團), male, dorsal view.
Habitus of Arthrotus fulvus Chûjô A collected from Motien (摩天), male, dorsal view B collected from Chenghsipao (鎮西堡), male, dorsal view C collected from Pilu (碧綠), female, dorsal view D collected from Taipingshan (太平山), female, dorsal view E collected from Wufeng (五峰), female, dorsal view F collected from Chungchihkuan (中之關), male, dorsal view G collected from Tsuifeng (翠峰), male, dorsal view H collected from Tsuifeng (翠峰), male, venral view I collected from Meifeng (梅峰), male, dorsal view.
Male. Length 5.1–5.4 mm, width 2.5–2.8 mm. Antennae filiform (Fig.
Female. Length 5.4–6.3 mm, width 2.6–3.3 mm. Antennae (Fig.
One specimen collected from Penpuchi (本部溪) was misidentified as A. testaceus by
Leaves of Acer insulare var. caudatifolium (Hayata) S.Y. Lu & Y.P. Yang (Sapindaceae), Alnus formosana (Burkill) Makino (Betulaceae), Stachyurus himalaicus Hook. f. & Thomson (Stachyuraceae), and Persicaria chinensis (L.) H. Gross (Polygonaceae).
Arthrotus fulvus is widespread from lowlands to mid-elevations of Taiwan. Adult color forms A, B, and C are allopatric. Members of form A were collected from central Taiwan, including Hsinchu, Ilan, Hualien, north Nantou, and Taichung counties; color form B from southwest Taiwan, including south Nantou, Chiayi, and Kaoshiung counties; color form C from southeast Taiwan only, including Taitung county. Most adults with yellowish or reddish brown elytra were collected from lowlands, while most adults with maculate or metallic blue elytra were form mid-elevations (Fig.
Arthrotus saigusai
Kimoto, 1969: 62;
Arthrotus fulvus:
Holotype
♂ (
Yellowish brown form (n = 114): Taiwan. Chiayi: 1♀ (HTC), 阿里山 (Alishan), 23–24.IV.1928, leg. Matsumura; 1♀ (
(n = 17): Taiwan. Hualien: 2♀ (
Adults of Arthrotus saigusai Kimoto are similar to those of A. tricolor (Chûjô) and A. fulvus Chûjô in possessing rounded lateral margins of pronota (straight lateral margins of pronotum in A. abdominalis (Chûjô), A. gressitti Kimoto, A. hirashimai Kimoto, and A. yuae sp. nov.), the less transverse pronotum and elytra, 1.7–2.0 × wider and long in pronotum and 1.5–1.6 × longer than wide in elytra (the more transverse pronotum and elytra, 2.1–2.2 × wider and long and 1.4 × longer than wide in elytra of A. testaceus Gressitt & Kimoto and A. yangi sp. nov.). Adults of A. saigusai are different from those of A. fulvus and A. tricolor by the less transverse antennomere III in male, 1.1 × longer than wide (Fig.
Pale individuals have yellowish brown bodies with black antennae except antennomeres I, tibiae, and tarsi (Fig.
Habitus of Arthrotus saigusai Kimoto A collected from Kuanshan Wind Gap (關山啞口), male, dorsal view B collected from Kuanshan Wind Gap (關山啞口), male, ventral view C collected from Kuanshan Wind Gap (關山啞口), male, dorsal view D collected from Kuanshan Wind Gap (關山啞口), male, ventral view E collected from Tienchih (天池), female, dorsal view F collected from Huakang (華岡), female, dorsal view.
Habitus of Arthrotus saigusai Kimoto A collected from Hsiaofengko (小風口), male, dorsal view B collected from Hsiaofengko (小風口), male, ventral view C collected from Huakang (華岡), female, dorsal view D collected from Huakang (華岡), female, ventral view E collected from Hsiangyangshan (向陽山), male, dorsal view F collected from Hsiangyangshan (向陽山), male, ventral view.
Male. Length 5.0–5.2 mm, width 2.4–2.5 mm. Antennae filiform (Fig.
Female. Length 5.9–6.5 mm, width 3.0–3.3 mm. Antennae (Fig.
Leaves of Reynoutria japonica Houtt. (Polygonaceae). Adults were found hiding inside the curled tender leaves of the food plants (Fig.
One specimen collected from Ho Huan Shan (合歡山) by K. Baba was misidentified as Arthrotus fulvus by
Arthrotus fulvus Chûjô, 1938: 139 (part).
Dercetis tricolor Chûjô, 1965: 95.
Dercetina tricolor:
Arthrotus tricolor:
Holotype
♀ (
A total of 299 specimens was examined (Suppl. material
Adults of Arthrotus tricolor (Chûjô) are similar to those of A. fulvus Chûjô in possessing rounded lateral margins of pronota (straight lateral margins of pronotum in A. abdominalis (Chûjô), A. gressitti Kimoto, A. hirashimai Kimoto, and A. yuae sp. nov.), the less transverse pronotum and elytra, 1.7–2.0 × wider and long in pronotum and 1.5–1.6 × longer than wide in elytra (more transverse pronotum and elytra, 2.1–2.2 × wider and long and 1.4 × longer than wide in elytra of A. testaceus Gressitt & Kimoto and A. yangi sp. nov.), a more transverse antennomere III in male, 0.7–0.8 × longer than wide (Fig.
Color yellowish brown; head, scutellum, and prothorax reddish brown, but antennae black; elytra with black stripes along basal margin, extending along entire suture, and lateral margins from base to apical 1/3, with two transverse black stripes at basal 1/3 and apical 1/3, legs black (Fig.
Habitus of Arthrotus tricolor (Chûjô) A collected from Chutzuhu (竹子湖), male, dorsal view B collected from Chutzuhu (竹子湖), male, ventral view C collected from Chutzuhu (竹子湖), male, lateral view D collected from Chutzuhu (竹子湖), female, dorsal view E collected from Junghua (榮華), male, dorsal view F collected from Wulai (烏來), male, dorsal view G collected from Lilungshan (里龍山), male, dorsal view H collected from Wushihkeng (烏石坑), male, dorsal view I ditto, ventral view.
Male. Length 5.2–5.6 mm, width 2.4–2.8 mm. Antennae filiform (Fig.
Female. Length 5.4–6.4 mm, width 3.1–3.8 mm. Antennae (Fig.
Some specimens collected from Tahsuehshan (大雪山) and nearby Wushihkeng (烏石坑) have a distinctive color form (Fig.
Two types of Arthrotus fulvus [1♀ (
Leaves of Styrax formosanus var. formosanus Matsum. (Styracaceae), Cryptocarya chinensis (Hance) Hemsl. (Lauraceae), and Actinidia rufa (Siebold & Zucc.) Planch. ex Miq. (Actinidiaceae).
Arthrotus testaceus
Gressitt & Kimoto, 1963: 702 (China);
Arthrotus shibatai
Kimoto, 1984: 55 (Taiwan);
Arthrotus testaceus. Paratypes. 1♂ (
Arthrotus shibatai. Holotype ♂ (
A total of 301 specimens was examined (Suppl. material
Adults of Arthrotus testaceus Gressitt & Kimoto (Figs
Adults from China yellowish brown, but extremely variable in Taiwan, some similar to those of China (Fig.
Habitus of Arthrotus testaceus Gressitt & Kimoto A collected from Tengchih (藤枝), male, dorsal view B collected from Peitawushan (北大武山), male, dorsal view C ditto, ventral view D collected from Tahanshan (大漢山), female, dorsal view E collected from Tahanshan (大漢山), male, dorsal view F collected from Tahanshan (大漢山), male, dorsal view G collected from Tahanshan (大漢山), male, dorsal view H collected from Tahanshan (大漢山), male, dorsal view I collected from Tahanshan (大漢山), male, dorsal view.
Habitus of Arthrotus testaceus Gressitt & Kimoto and A. yangi sp. nov. A A. testaceus, collected from Tahanshan (大漢山), male, dorsal view B A. testaceus, collected from Tahanshan (大漢山), male, ventral view C A. testaceus, collected from Tahanshan (大漢山), male, dorsal view D A. yangi sp. nov., collected from Hualuhsi (華綠溪), male, dorsal view E A. yangi sp. nov., collected from Hualuhsi (華綠溪), male, ventral view F A. yangi sp. nov., collected from Hualuhsi (華綠溪), female, dorsal view.
Male. Length 6.5–7.1 mm, width 3.9–4.0 mm. Antennae filiform (Fig.
Female. Length 7.2–7.9 mm, width 4.3–4.4 mm. Antennae (Fig.
Leaves of Sapindus mukorossi Gaertn. (Sapindaceae), Acer insulare var. caudatifolium (Hayata) S.Y. Lu & Y.P. Yang (Sapindaceae), Acer albopurpurascens Hayata (Sapindaceae), Alniphyllum pterospermum Matsum. (Styracaceae), Alnus formosana (Burkill) Makino (Betulaceae), and Lithocarpus hancei (Benth.) Rehder (Fagaceae).
Adults of Arthrotus testaceus Gressitt & Kimoto are widespread in lowlands (below 1500 m) of Taiwan (Fig.
(n = 3). Holotype ♂ (NMNS): Taiwan. Hualien: Hualuhsi (華綠溪), 19.IV.-1.VI.2011, leg. W.-T. Yang & K. W. Huang, with Malaise trap. Paratypes. 1♀ (NMNS), same data as holotype; 1♀ (NMNS): Taiwan. Taitung: Shinkangshan (新港山), 24.III.-19.V.2009, leg. W.-T. Yang & K. W. Huang, with Malaise trap.
Adults of Arthrotus yangi sp. nov. (Fig.
Color (Fig.
Male. Length 5.5 mm, width 3.3 mm. Antennae filiform (Fig.
Female. Length 6.7 mm, width 4.1 mm. Antennae (Fig.
Unknown.
Adults are collected from only two localities in East Taiwan (Fig.
The specific name is dedicated to Mr Wan-Tsun Yang (楊萬琮) who collected type specimens using Malaise traps.
1 | Lateral margins of pronotum straight; elytra metallic blue, with transverse depression at basal 1/ 3 (Figs |
2 |
– | Lateral margins of pronotum rounded; color patterns of elytra variable, without transverse depression (Figs |
5 |
2 | Head, pronotum, underside of thorax, and legs metallic blue (Fig. |
A. abdominalis (Chûjô) |
– | Head, pronotum, underside of thorax, and legs blackish or yellowish brown (Figs |
3 |
3 | Head, pronotum, underside of thorax, and legs blackish brown (Fig. |
A. gressitti Kimoto |
– | Head, pronotum, underside of thorax, and legs yellowish brown (Fig. |
4 |
4 | Tectum of aedeagus covered with short needle-shape setae laterally and stout teeth apically, apex curved in lateral view (Fig. |
A. hirashimai Kimoto |
– | Tectum of aedeagus with scattered, stout setae, apex recurved in lateral view (Fig. |
A. yuae sp. nov. |
5 | Pronotum and elytra more transverse, pronotum 2.1–2.2 × wider than long and elytra 1.4 × longer than wide (Figs |
6 |
– | Pronotum and elytra less transverse, prontum1.7–2.0 × wider than and elytra 1.5–1.6 × longer than wide (Figs |
7 |
6 | Color pattern of elytra characteristic (Fig. |
A. yangi sp. nov. |
– | Color pattern on elytra variable but lacking above color pattern (Figs |
A. testaceus Gressitt & Kimoto |
7 | Color pattern on elytra characteristic (Figs |
A. saigusai Kimoto |
– | Color pattern on elytra variable but lacking above color pattern (Figs |
8 |
8 | Color pattern on elytra characteristic (Fig. |
A. tricolor (Chûjô) |
– | Color pattern on elytra variable but lacking above color pattern (Figs |
A. fulvus Chûjô |
Taxonomic studies on Arthrotus species of Taiwan are difficult due to females being confused with those of Dercetina and the great variation of color patterns of most species. Taiwanese species of Dercetina were revised recently (
Five Arthrotus species are widespread in Taiwan, A. abdominalis, A. testaceus, A. saigusai, A. tricolor, and A. fulvus. Of these, only A. saigusai is alpine. Populations of Arthrotus gressitti, A. hirashimai, A. yuae sp. nov., and A. yangi sp. nov. are localized and have similar color patterns and body shapes but allopatric distributions. In addition, one interesting phenomenon was noticed, the two bicolored patterns (black and white, black and red) of A. fulvus (Fig.
File photographs of Arthrotus fulvus Chûjô and Neochya nitidissima (Chûjô) A color form A of A. fulvus collected from Meifeng (梅峰) B similar color pattern of N. nitidissima collected from Kuanwu (觀霧) C color form B of A. fulvus collected from Tatachia (塔塔加) D similar color pattern of N. nitidissima collecte from Tatachia (塔塔加)
I am grateful to the Taiwan Chrysomelid Research Team (TCRT), including Jung-Chang Chen (陳榮章), Hou-Jay Chen (陳厚潔), Yi-Ting Chung (鍾奕霆), Bo-Xin Guo (郭泊鑫), Hsueh Lee (李雪), Wen-Chuan Liao (廖文泉), His-Feng Lu (陸錫峯), Mei-Hua Tsou (曹美華), and Su-Fang Yu (余素芳) for assistance in collecting material. I especially thank Yi-Chia Chiu (邱奕家), Chi-Lung Lee (李其龍), and Hsing-Tzung Cheng (鄭興宗) for photos of specimens, Ta-Hsiang Lee (李大翔), Chia-Wen Tsai (蔡佳雯), and Mei-Hua Tsou (曹美華), for their field photography, Chih-Kai Yang for identification of host plants. This study was supported by the Ministry of Science and Technology MOST 109-2313-B-055-003. I especially thank Chang Chin Chen for assisting this study in various ways and Chris Carlton for reading the draft and editing for American English style. Finally, I am grateful to Thomas Wagner and Alexey Tischechkin for reviewing the manuscript.
Arthrotus abdominalis (Chûjô, 1962)
Data type: docx file
Explanation note: Arthrotus abdominalis (Chûjô, 1962)
Arthrotus fulvus Chûjô, 1938
Data type: docx file
Explanation note: Arthrotus fulvus Chûjô, 1938
Arthrotus tricolor (Chûjô, 1965)
Data type: docx file
Explanation note: Arthrotus tricolor (Chûjô, 1965)
Arthrotus testaceus Gressitt & Kimoto, 1963
Data type: docx file
Explanation note: Arthrotus testaceus Gressitt & Kimoto, 1963