Research Article |
Corresponding author: Ming Bai ( baim@ioz.ac.cn ) Academic editor: Frank Krell
© 2016 Li Sha, Enrico Ricchiardi, Ming Bai, Xing-Ke Yang.
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:
Li S, Ricchiardi E, Bai M, Yang X (2016) A taxonomy review of Oreoderus Burmeister, 1842 from China with a geometric morphometric evaluation (Coleoptera, Scarabaeidae, Valgini). ZooKeys 552: 67-89. https://doi.org/10.3897/zookeys.552.6096
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The species of the genus Oreoderus are morphologically similar, and can be challenging to distinguish without dissecting the male genitalia. In this study, the Oreoderus species from China are reviewed. Three new species of Oreoderus are described: O. dasystibialis Li & Yang, sp. n., O. brevitarsus Li & Yang, sp. n. and O. oblongus Li & Yang, sp. n. A key of the male Oreoderus and a distribution map are provided. Oreoderus coomani Paulian, 1961 was found as a new record in China. The first description of the female of O. arrowi Ricchiardi, 2001 is provided. Oreoderus humeralis Gestro, 1891, O. quadricarinatus Arrow, 1944, O. crassipes Arrow, 1944, and O. momeitensis Arrow, 1910 are excluded from the Chinese fauna. Furthermore, we utilize geometric morphometric approaches (GM) to analyze the shape variation of four characters (pronotum, elytra, protibia and aedeagus) in Oreoderus. The morphological variations of Oreoderus and the taxonomic value of each character are discussed. The combined analysis of geometric morphometrics and comparative morphology support recognition of the three new species.
Beetles, Cetoniinae , China, new species, GM
Oreoderus Burmeister, 1842 is a genus of Valgini (Coleoptera: Scarabaeidae) which comprises 29 species, distributed only in the Oriental Region. Adults are flower-visitors, the larvae develop in the rotten wood (with or without termites) (
Geometric morphometrics (GM) is a useful tool for shape analysis in biology. This tool has an important advantage: not only does it offer precise and accurate description, but it also serves the equally important purposes of visualization, interpretation and communication of results (
In this study, all known species and three new species described in this paper (32 species and 82 specimens total) of Oreoderus and 2 species (2 specimens) of the out groups Hybovalgus Kolbe, 1904 and Dasyvalgus Kolbe, 1904 were selected for geometric morphometric analyses (Table
Species | Characters | ||||
---|---|---|---|---|---|
Pronotum | Elytra | Protibia | Aedeagus (♂) | ||
1. | Oreoderus aciculatus Paulian, 1961 | 1 | 1 | 1 | 1 |
2. | Oreoderus ahrensi Ricchiardi, 2001 | 1 | 1 | 1 | 1 |
3. | Oreoderus argillaceus (Hope, 1841) | 1 | 1 | 1 | 1 |
4. | Oreoderus arrowi Ricchiardi, 2001 | 6 | 6 | 6 | 5 |
5. | Oreoderus bengalensis Ricchiardi, 2001 | 1 | 1 | 1 | 1 |
6. | Oreoderus bhutanus Arrow, 1910 | 1 | 1 | 1 | 1 |
7. | Oreoderus bidentatus Ricchiardi, 2001 | 2 | 2 | 2 | 1 |
8. | Oreoderus birmanus Ricchiardi, 2001 | 1 | 1 | 1 | 1 |
9. | Oreoderus brevicarinatus (Pic, 1928) | 2 | 1 | 1 | 1 |
10. | Oreoderus brevipennis Gestro, 1891 | 1 | 1 | 1 | 1 |
11. | Oreoderus brevitarsus sp. n. | 10 | 10 | 7 | 6 |
12. | Oreoderus clypealis Arrow, 1944 | 1 | 1 | 1 | 1 |
13. | Oreoderus coomani Paulian, 1961 | 9 | 9 | 9 | 7 |
14. | Oreoderus crassipes Arrow, 1944 | 1 | 1 | 1 | 1 |
15. | Oreoderus dasystibialis sp. n. | 3 | 3 | 3 | 3 |
16. | Oreoderus gestroi Ricchiardi, 2001 | 1 | 1 | 1 | 1 |
17. | Oreoderus gracilicollis Paulian, 1961 | 1 | 1 | 1 | 1 |
18. | Oreoderus gravis Arrow, 1910 | 1 | 1 | 1 | 1 |
19. | Oreoderus humeralis Gestro, 1891 | 1 | 1 | 1 | 1 |
20. | Oreoderus insularis Ricchiardi, 2001 | 1 | 1 | 1 | 1 |
21. | Oreoderus longicarinatus Ricchiardi, 2001 | 1 | 1 | 1 | 1 |
22. | Oreoderus maculipennis Gestro, 1891 | 15 | 15 | 11 | 4 |
23. | Oreoderus meridionalis Paulian, 1961 | 1 | 1 | 1 | 1 |
24. | Oreoderus momeitensis Arrow, 1910 | 1 | 1 | 1 | 1 |
25. | Oreoderus oblongus sp. n. | 10 | 10 | 7 | 6 |
26. | Oreoderus pseudohumeralis Ricchiardi, 2001 | 1 | 1 | 1 | 1 |
27. | Oreoderus quadricarinatus Arrow, 1944 | 1 | 1 | 2 | 1 |
28. | Oreoderus quadrimaculatus Miyake, Yamaguchi & Aoki, 2004 | - | - | 1 | 1 |
29. | Oreoderus rufulus Gestro, 1891 | 1 | 1 | 1 | 1 |
30. | Oreoderus siamensis Ricchiardi, 2001 | 1 | 1 | 1 | 1 |
31. | Oreoderus sikkimensis Ricchiardi, 2001 | 1 | 1 | 1 | 1 |
32. | Oreoderus waterhousei Gestro, 1891 | 1 | 1 | 1 | 1 |
33. | Dasyvalgus ichangcius Moser, 1915 | 1 | 1 | 1 | 1 |
34. | Hybovalgus yunnanus Moser, 1906 | 1 | 1 | 1 | 1 |
The specimens are deposited in the following collections:
ERC Enrico Ricchiardi Collection, Turin, Italy;
IEZU Institute of Applied Entomology,
IZAS
NHML The
PCRD Private Collection of Ran DAI, Kunming, Yunnan, China.
The description of morphological characters follows the terminology of
Four characters (pronotum, protibia, elytra, and aedeagus) were examined and analyzed. There was negligible difference in the shape of pronotum and elytra for the male and female. However, sexual dimorphism often occurs in the protibia of cetoniines and other scarabs (
The morphology of the four characters (pronotum, protibia, elytra, and aedeagus) was represented by curves. Each curve was based on homologous or corresponding criteria. The pronotum was represented by 5 curves. Curve 1 represented the outline of the pronotum, which resampled into 50 semi-landmarks (SLM). Curve 2 and Curve 3 represented the outline of the carinae, which resampled into 15 SLM. Curve 4 and Curve 5 represented the outline of lateral carinae, which resampled into 10 SLM. The pronotum, elytra and aedeagus were each represented by a single curve, which resampled into 50 SLM (Fig.
Curve selection of four characters. A the red curve (Curve 1) is the outline of pronotum, which resampled into 50 semi-landmarks (SLM); the two green curves (Curve 2, 3) are outline of the carinae, which resampled into 15 SLM; two blue curves (Curve 4, 5) are the outline of lateral carinae, which resampled into 10 SLMB the curve is the outline of elytra, which resampled into 50 SLMC the curve is the outline of protibia, which resampled into 50 SLMD the curve is the outline of the left paramere, which resampled into 50 SLM.
These curves were digitized with tps-DIG 2.05 (
The Procrustes distances and Mahalanobis distances computed from canonical variate analysis (CVA) can be used to explain the differences and modes of evolution. Procrustes distance is a measure of the absolute magnitude of the shape deviation and indicates how big the differences are between the average group shape. Mahalanobis distance provides an indication of how different an individual is relative to the others in the sample, and how distinctly groups are separated from one another.
Oreoderus Burmeister, 1842: 726.
Valgus argillaceus Hope, 1841, by monotypy.
Oreoderus can be distinguished from all other genera within the tribe Valgini by the following characters: a) protibia with only two or three external teeth; b) the first joint of the hind tarsus shorter than the second one; c) pronotum elongated with four carinae; d) visible sternite V twice longer than sternite IV.
1 | Pronotal carinae forms two noticeable hooked tubercles at the anterior margin | 2 |
– | Pronotal carinae do not forms any tubercles at the anterior margin | 3 |
2 | Pronotal scale tufts not present; propygidial spiracles completely obsolete; propygidium without any scale tufts at the hind margin | O. argillaceus (Hope, 1841) |
– | Pronotum with scale tufts on the small lateral carinae; propygidial spiracles moderately elevated; propygidium with two scale tufts at the hind margin | O. insularis Ricchiardi, 2001 |
3 | Pronotal carinae very long, reaching over 3/4 of the pronotum length | 4 |
– | Pronotal carinae not reaching over 2/3 of the pronotum length | 5 |
4 | Pronotal carinae obsolete | O. gravis Arrow, 1910 |
– | Pronotal carinae interrupted twice, sharp | O. longicarinatus Ricchiardi, 2001 |
5 | Pronotal carinae long, reaching about 2/3 of the pronotum length | 6 |
– | Pronotal carinae very short, not reaching 1/2 of the pronotum length | 27 |
6 | Pronotal carinae interrupted once, sharp | 7 |
– | Pronotal carinae never interrupted | 9 |
7 | Pronotal scale tufts on the small lateral carinae | 8 |
– | Pronotal scale tufts close to the scutellum | O. brevipennis Gestro, 1891 |
8 | Propygidial spiracles moderately elevated; propygidium without any scale tufts at the hind margin | O. meridionalis Paulian, 1961 |
– | Propygidial spiracles completely obsolete; propygidium with two scale tufts at the hind margin | O. siamensis Ricchiardi, 2001 |
9 | Pronotal carinae obsolete | 10 |
– | Pronotal carinae sharp | 11 |
10 | Elytra with four patches of dark-colored scales | O. quadrimaculatus Miyake, Yamaguchi & Aoki, 2004 |
– | Elytra without patches of scales | O. waterhousei Gestro, 1891 |
11 | Pronotal lateral carinae join the lateral margin or arrive very close | 12 |
– | Pronotal lateral carinae stop well before the lateral margin | 14 |
12 | Propygidial spiracles completely obsolete | 13 |
– | Propygidial spiracles sharply elevated | O. momeitensis Arrow, 1910 |
13 | Propygidium without any scale tufts at the hind margin | O. quadricarinatus Arrow, 1944 and O. rufulus Gestro, 1891* |
– | Propygidium with two scale tufts at the hind margin | O. arrowi Ricchiardi, 2001 |
14 | Third teeth of protibia not present | 15 |
– | Third teeth of protibia present | 18 |
15 | Mesotibia without bush of thick scales | 16 |
– | Mesotibia with bush of thick scales | 17 |
16 | Meso- and metatibia without a spine at the middle of posterior margin | O. brevitarsus Li & Yang, sp. n. |
– | Meso- and metatibia with a spine at the middle of posterior margin | O. ahrensi Ricchiardi, 2001 |
17 | Protibia sharp, cariane reaching over the middle of the pronotum | O. bidentatus Ricchiardi, 2001 |
– | Protibia blunt, cariane not reaching the middle of the pronotum | O. dasystibialis Li & Yang, sp. n. |
18 | Third tooth of protibia much smaller than first and second | 19 |
– | Third tooth of protibia similar in size as the second | O. gracilicollis Paulian, 1961 |
19 | Anterior margin of clypeus sinuated, with a bifid processes | O. gestroi Ricchiardi, 2001 |
– | Anterior margin of clypeus pointed, simply sinuated or rounded but without a bifid process | 20 |
20 | Hind margin of propygidium centrally projected toward the back, not pointed | O. humeralis Gestro, 1891 |
– | Hind margin of propygidium centrally rounded or smoothly curved inward | 21 |
21 | First joint of hind tarsi shorter than the second | 22 |
– | First joint of hind tarsi as long as the second | 25 |
22 | Propygidium covered by simple, not coffee grain shaped scales | 23 |
– | Propygidium centrally or near the scale tufts with areas of raised c-shaped scales | O. pseudohumeralis Ricchiardi, 2001 |
23 | Propygidial spiracles moderately elevated | O. aciculatus Paulian, 1961 |
– | Propygidial spiracles completely obsolete | 24 |
24 | Meso- and metatibia with a spine at the middle of posterior margin | O. maculipennis Gestro, 1891 |
– | Meso- and metatibia without a spine at the middle of posterior margin | O. oblongus Li & Yang, sp. n. |
25 | Pronotal carinae continue after the middle of the length with a triangular area made by C- shaped, black, raised scales that reaches the hind margin; propygidium centrally or near the scale tufts with areas made by raised c-shaped scales; propygidium with two scale tufts at the hind margin | O. bengalensis Ricchiardi, 2001 |
– | No triangular black scales area is present at the end of the pronotal carinae; propygidium covered by simple, not coffee grain shaped, scales; propygidium without any scale tufts at the hind margin | 26 |
26 | Metatibial intrusion interposed between the two mobile spurs present; pronotal carinae not parallel, but arched or sinuated | O. birmanus Ricchiardi, 2001 |
– | Metatibial intrusion interposed between the two mobile spurs not present; pronotal carinae almost parallel and the included area is narrow | O. coomani Paulian, 1961 |
27 | Pronotal carinae obsolete | 28 |
– | Pronotal carinae sharp | 30 |
28 | First joint of hind tarsi shorter than the second | 29 |
– | First joint of hind tarsi as long as the second | O. crassipes Arrow, 1944 |
29 | Propygidium without any scale tufts at the hind margin | O. sikkimensis Ricchiardi, 2001 |
– | Propygidium with two scale tufts at the hind margin | O. bhutanus Arrow, 1910 |
30 | Third tooth of protibia much smaller than first and second; meso- and metatibia without any scales brush covering the median posterior tooth; first hind tarsi joint shorter than the second | O. clypealis Arrow, 1944 |
– | Third and second teeth of protibia similar in size; meso- and metatibia with a well noticeable brush made by ochraceous scales covering the medial posterior tooth; first hind tarsi joint as long as the second | O. brevicarinatus (Pic, 1928) |
*To separate them see the shape of the parameres.
The genus Oreoderus was recently revised by
Seven species are now known from China, including the three new species (O. brevitasus Li & Yang, sp. n., O. dasystibialis Li & Yang, sp. n., and O. oblongus Li & Yang, sp. n.), plus a range extension (O. coomani). The three species previously known from China and confirmed in our study are O. arrowi, O. bidentatus, and O. maculipennis.
Holotype ♂, P.R. CHINA, Zhejiang, Mt. Gutianshan, G45–G15, 560m, broad-leaved mixed forest, 2009.VIII.2–5, leg. Liu Chongling. Holotype deposited in IZAS. Paratypes: P.R. CHINA, 1♂, Zhejiang, Mt. Gutianshan, G24ha-140, 446–715m, Broad-leaved mixed forest, 2009.VII. 5–8, leg. Liu Chongling, (IZAS); 1♀, Zhejiang, Mt. Gutianshan, G24ha-157, 446–715m, broad-leaved mixed forest, 2009.IX. 24–27, leg. Liu Chongling, (IZAS); 1♀, Zhejiang, Mt. Gutianshan, G24ha-83, 446–715m, broad-leaved mixed forest, 2009.VII. 26–29, leg. Liu Chongling, (IZAS); 1♀, Zhejiang, Mt. Gutianshan, G24ha-111, 446–715m, Broad-leaved mixed forest, 2009.VII. 5–8, leg. Liu Chongling, (IZAS); 1♂, Zhejiang, Mt. Gutianshan, 1992.VII.27, leg. Wu Hong, IOZ(E)902215, (IZAS); 1♂, Fujian, Zhang???hu (unrecognized name), 1981.VI.19, IOZ(E)902216, (IZAS); 1♂, Yunnan, Qiubei, Shupi 1278, Quercus, 1500m, leg. Kui Meihua, 1979.VII.6, IOZ(E)902199, (IZAS); 1♀, Yunnan, Xishuangbanna, Mengzhe, 1200m, 1958.VIII.23, leg. Pu Fuji, IOZ(E)902201, (IZAS); 1♂, Yunnan, Xishuangbanna, Xiaomengyang, 850m, 1958.IX.4, leg. MengXuwu, IOZ(E)90221, (IZAS); 1♂, Yunnan, Yiwu, Banna, Menglun, 650m, 1959.VIII.3, leg. Pu Fuji, IOZ(E)902211, (ERC).
1♀, P.R. CHINA, Zhejiang, Thousand island Lake (IEZU).
Based on the morphological comparison and PCA analysis of four characters (see below), this new species is close to O. dasystibialis and O. bidentatus, but differs from O. bidentatus by the shape of the carinae of the pronotum and can be separated from O. dasystibialis by the sharp teeth on the protibia and the absence of a thick brush on meso- and metatibia in the male. Finally, the aedeagi are very distinctive in the three species. The parameres are much slender than the other two species and the outer margin is sinuate. The female O. brevitarsus can be distinguished from O. bidentatus by the short stylus and the presence of a brush on the inner margin of the protibia.
Length 8.2 mm; width 3.9 mm. Color: light brown to brown. Head: clypeus anteriorly rounded, lateral margin extended, with erected setae. Frons densely covered with testaceous scales. Ocular canthus short and broad, covered with same scales as clypeus. Antenna with 10 segments, clubs much longer than antennomeres 2–7. Pronotum: widest at the base, lateral margins sinuate. Surface covered with oblong lied-down scales. Carinae sharp, highly prominent, ending over 1/2 of the pronotum length; lateral small carinae short, sharp, not reaching the lateral margin of the pronotum. Scutellum: triangular, rounded at the apex, covered with shorter oval scales than pronotum. Elytra: with rows of punctate lines, covered with similar scales as scutellum. Propygidium: anteriorly glabrous and slightly punctate, posterior densely punctate and covered with scales. Spiracles moderately elevated. Pygidium: with thick, lied-down scales and one central scale tuft at the hind margin. Venter: coarsely and densely punctate. Visible sternite V smooth and bald in the middle. Visible sternite V twice longer than sternite IV. Legs: slender, femora and tibia covered with testaceous scales except for protibia. Outer margin of mesometatibia covered with more dense scales. Protibia short and bidentate, external teeth sharp. Tarsomeres with short setae. Parameres: short, lateral margin sinuate, the apex covered with yellow setae.
Length 7.5–10.4 mm; width 3.5–4.7. Pronotum broader than the male one; carinae much shorter, reaching about 1/3 of the disc. Propygidium much longer, and pygidium with a sharp stylus. Visible sternite V much broader. Protibia short and with thick brush in the inner margin; tooth slightly blunt, mesometatibia with same brush on the inner and outer margin.
Male paratypes: length 9.0–9.5 mm; width 4.7–5.1 mm, and very similar to the holotype.
The new species is named for the short tarsi of protibia.
Holotype, ♂, P.R. CHINA, Hainan, Mt. Jiangfengling, Heiling, 1982.VII.10, leg. Hua Lizhong, Sun Yat-sen University/100/IOZ(E)1658787. Holotype deposited in IZAS. Paratypes: P.R. CHINA: 1♂, Hainan, Mt. Jiangfengling, Wufenqu, 1981.VI.29, leg. Wang (26), Sun Yat-sen University/99/IOZ(E)1658786, (IZAS); 1♂, Hainan, Kwangtung, 1934.IX.1, leg. He Chi, Fan Inst. Biol Peiping/ IOZ(E)902260, (ERC).
based on the morphological comparison and PCA analysis of four characters (see below), this new species is close to O. brevitarsus and O. bidentatus, but differs from O. bidentatus by the smaller lateral carinae of the pronotum and can be separated from O. brevitarsus by the blunt teeth on protibia and the appearance of a thick brush on meso- and metatibia in the male. The aedeagi are very distinctive among these three species. The apex of the parameres in the new species is much wider than in the other two.
length 8.5 mm; Width 4.3 mm. Color: light brown to brown. Head: clypeus short, anteriorly straight, with erected setae. Frons densely punctate, covered with testaceous scales. Ocular canthus short, covered with same scales. Antenna with 10 segments, clubs much longer than antennomeres 2–7. Pronotum: widest at the base, lateral margins sinuate. Surface densely punctate, covered with thick testaceous scales. Carinae sharp, highly prominent, ending at 2/3 of the pronotum length; lateral small carinae long, sharp, reaching the lateral margins of the pronotum. Scutellum: triangular, rounded at the apex, covered with testaceous scales. Elytra: with rows of punctate line, covered with same scales as scutellum. Propygidium: covered with lied down scales. Spiracles moderately elevated. Pygidium: triangular form, coarsely punctate, covered with thick lied down oval scales. Venter: coarsely and densely punctate with each point bearing a testaceous scale. A longitudinal groove is visible in the middle of visible sternites I–IV; visible sternite V longer twice than the sternite IV. Legs: slender, covered with testaceous scales except protibia. Protibia extended, bidentate, teeth blunt. Meso- and metatibia with thick brush on the outer margins. Tarsomeres with short setae. Parameres: relatively longer and much broader, the apex is the widest part.
unknown.
paratypes length 7.5–7.7 mm; width 3.8–4.2 mm, and very similar to the holotype.
the new species is named according to its thick brush on meso- and metatibia.
China: Hainan.
Holotype, ♂, P.R. CHINA, Yunnan, Xishuangbanna, Jinghong, 650m, 1958.VIII.12, leg. Meng Xuwu, IOZ(E)902189. Holotype deposited in IZAS. Paratypes: P.R. CHINA: 1♂, Yunnan, Xishuangbanna, Xiaomengyang, 850m, 1958.VIII.19, leg. Zhang Yiran, IOZ(E)902182, (IZAS); 1♂, Yunnan, Xishuangbanna, Xiaomengyang, 850m, 1958.VIII.20, leg. Pu Fuji, IOZ(E)902183, (IZAS); 1♂, Yunnan, Xishuangbanna, Xiaomengyang, 850m, 1958.IX.2, leg. Zheng Leyi, IOZ(E)902185, (IZAS); 1♂, Yunnan, Xishuangbanna, Jinghong, 650m, 1958.VIII.26, leg. Meng Xuwu, IOZ(E)902186, (IZAS); 1♂, Yunnan, Xishuangbanna, Jinghong, 650m, 1958.VII.15, leg. Meng Xuwu, IOZ(E)902188, (IZAS); 1♂, Yunnan, Xishuangbanna, Menglun, 580m, 1993.IX.10, leg. Xu Huanli, IOZ(E)902204, (IZAS); 1♂, Yunnan, Xishuangbanna, Menglun, 600m, 1993.IX.9, leg. Yang Longlong, IOZ(E)902205, (IZAS); 1♂, Yunnan, Xishuangbanna, Jinghong, 650m, 1958.VII.27, leg. Meng Xuwu, IOZ(E)902208, (ERC); 1♀, Yunnan, Xishuangbanna, Xiaomengyang, 850m, 1957.X.20, leg. Pu Fuji, IOZ(E)902223, (IZAS).
Based on the morphological comparison and PCA analysis of four characters (see below), this new species is close to O. maculipennis, but lacks a spine on the outer margin of meso- and metatibia and elytra without white patch scales in the new species. The parameres are very short in O. oblongus.
Length 8.4 mm; width 4.0 mm. Color: light brown to brown. Head: clypeus anteriorly rounded, with erect setae on the anterior margin. Frons covered with testaceous scales. Ocular canthus short, covered with same scales. Antenna with 10 segments, clubs not much longer than antennomeres 2–7. Pronotum: widest at base, lateral margins sinuate. Surface densely punctate, covered with lied down scales. Carinae sharp, highly prominent, ending around 2/3 of the pronotum length; Lateral small carinae short, sharp, not joining the lateral margin of the pronotum. Scutellum: triangular, rounded at the apex, covered with testaceous scales. Elytra: with rows of punctate line, densely covered with oval scales. Propygidium: broad, punctate, covered with lied down scales. Spiracles moderately elevated. Pygidium: triangular, punctuation rounded, with thick lied down scales. Venter: coarsely and densely punctate with testaceous scales. Visible sternite V twice longer than sternite IV. Legs: slender, covered with oval scales except for protibia. Protibia tridentate; the third tooth small, far from the first two teeth. Meso- and metatibia covered with only sparse scales. Tarsomeres with short setae. Parameres: perpendicular to phallobase, the apex is sharp.
Length 9.7 mm; width 4.5 mm. Pronotum a little broader than the male; carinae slightly shorter. Propygidium much longer, and pygidium more highly prominent. Visible sternite V much broader. External tooth of protibia apparently blunt. Tarsomeres more robust.
paratypes length 7.0–9.2 mm; width 3.7–4.3 mm, and very similar to the holotype.
the new species is named for the oblong shape of the body.
Oreoderus coomani Paulian, 1961: 31.
Lectotype (designated by Ricchiardi, 2001), ♂, Tonkin, Hoa-Binh, A. de Cooman, (
P.R. CHINA: 1♂, Yunnan, Xihuangbanna, Xiaomengyang, 850m, 1957.X.20, leg. ZangLingchao, IOZ(E)902180; 1♂, Yunnan, Xihuangbanna, Menglun, 600m, 1993.IX.11, Mt. Shihuishan, leg. Yang Longlong, IOZ(E)902190; 1♂, Yunnan, Xihuangbanna, Menghai, 1100m, 1957.VIII.15, leg. Wang Shuyong, IOZ(E)902191; 1♂, Yunnan, Xihuangbanna, Menga, 1050–1080m, 1958.VIII.12, leg. Wang Shuyong, IOZ(E)902192; 1♂, Yunnan, Xihuangbanna, Menga, 1050–1080m, 1958.VIII.7, leg. Pu Fuji, IOZ(E)902193; 1♂, Yunnan, Xihuangbanna, Menga, 1050–1080m, 1958.VIII.20, leg. Pu Fuji, IOZ(E)902194; 1♂, Yunnan, Xihuangbanna, Menga, 1050–1080m, 1958.VIII.19, leg. Wang Shuyong, IOZ(E)902195; 1♂, Yunnan, Xihuangbanna, Menga, 1050–1080m, 1958.VIII.10, leg. Wang Shuyong, IOZ(E)902196; 1♂, Yunnan, Xihuangbanna, Xiaomengyang, 850m, 1957.X.26, leg. Wang Shuyong, IOZ(E)902197; 1♂, Yunnan, Malipo, 1958.VII.21, (IZAS).
China: Yunnan; Vietnam and Laos.
This species was previously known from Vietnam and Laos. This is the first record for Yunnan, China.
Oreoderus arrowi Ricchiardi, 2001: 521.
Holotype, ♂, S. China, 10–14.VII.1990, Jinghong, Prov., Yunnan, leg. S. Bečvář, (
P.R. CHINA: 1♂, Yunnan, Xihuangbanna, Jinghong, 650m, 1958.VII.7, leg. MengXuwu, IOZ(E)902175; 1♂, Yunnan, Xihuangbanna, Damenglong, 650m, 1958.VII.11, leg. ZhengLeyi, IOZ(E)902176; 1♂, Yunnan, Xihuangbanna, Damenglong, 650m, 1958.VII.11, leg. ZhengLeyi, IOZ(E)902177; 1♂, Yunnan, Xihuangbanna, Mengzhe, 870m, 1958.IX.7, leg. Wang Shuyong, IOZ(E)902178; 1♂, Yunnan, Xihuangbanna, Xiaomengyang, 850m, 1958.IX.2, leg. MengXuwu, IOZ(E)902179; 1♂, Yunnan, Xihuangbanna, Xiaomengyang, 1400m, 1957.X.4, leg. Wang Shuyong, IOZ(E)902187; 1♀, Yunnan, Naban River Nature Reserve, Mengsong, Danuoyou, 2007.XII.14, 770m, Danuoyou IV A, 14.XII.2007, leg. A. Weigel, 22.20699°N, 100.63761°E (trap), leg. A. Weigel, IOZ(E)1945434, (IZAS).
Length 8.4 mm; width 2.8mm. Color: light brown to brown. Head: clypeus anteriorly rounded, sharp in the apex, with erected setae. Frons covered with testaceous scales. Ocular canthus short and broad, covered with same scales. Antenna with 10 segments, club much longer than antennomeres 2–7. Pronotum: widest at base, lateral margin sinuate. Surface densely punctate, covered with testaceous scales. Carinae and lateral carinae sharp, highly prominent, ending before middle of pronotum. Scutellum: triangular, rounded at apex, covered with testaceous scales. Elytra: coarsely punctate, covered with testaceous scales. Propygidium: apparently longer than in male, hind margin rounded. Propygidial spiracles moderately elevated. Pygidium: narrower than in male, with thick lied down scales. Venter: coarsely and densely punctate with testaceous scale. Sternite V twice longer than Sternite IV; Sternite VI much narrower than male. Legs: slender, covered with testaceous scales except protibia. Protibia tridentate, tooth blunter than in male; meso- and metatibia with a spine on the outer margin. Tarsomeres much shorter than in male, covered with short setae.
Oreoderus bidentatus Ricchiardi, 2001: 526.
Holotype, ♂, India, Meghalaya, Kashia Hills. Paratype, 1♀, Yunnan, Bao Shan, 1700 m, 1993.V.1–3, (NHML).
1♀, Yunnan, Mt. Gaoligong, (PCRD).
China: Yunnan; North East India.
Oreoderus bidentatus was described from three males from Assam (2 paratypes) and Meghalaya (holotype) and one female from Yunnan (
Oreoderus maculipennis Gestro, 1891: 869.
Holotype, ♂,
1♂, Fujian, Shanghang, 650m, 1988.VII.24, IOZ(E)902212; 1♂, Fujian, Shanghang, 650m, 1988.VII.23, IOZ(E)902213; 1♂, Yunnan, Xishuangbanna, Xiaomengyang, 850m, 1957.X.26, leg. Wang Shuyong, IOZ(E)902203; 1♂, Yunnan, Naban, II/3D, 10.XI.2008, leg. L.Z. Meng, Yunnan, Jinghong, Naban River Nature Reserve Chachang (Forest), 2008.XI.10, 729m, 22.15810°N, 100.66543°E, leg. Meng Lingzeng; 1♀, Yunnan, 991 Fengqing, Fengshan, 1600m, leg. Zhang Fu, 1980.VII.26, IOZ(E)902214; 1♀, Yunnan, Xishuangbanna, Xiaomengyang, 850m, 1957.X.26, leg. Wang Shuyong, IOZ(E)902218; 1♂, Yunnan, Xishuangbanna, Xiaomengyang, 850m, 1957.X.26, leg. Wang Shuyong, IOZ(E)902219; 1♂, Yunnan, Xishuangbanna, Xiaomengyang, 850m, 1957.X.26, leg. Wang Shuyong, IOZ(E)902220; 1♂, Yunnan, Xishuangbanna, Xiaomengyang, 850m, 1957.X.25, leg. Wang Shuyong, IOZ(E)902221; 1♂, Yunnan, Xishuangbanna, Xiaomengyang, 850m, 1957.X.21, leg. Zang Lingchao, IOZ(E)902222; 1♀, Yunnan, Xishuangbanna, Menghai, 1200–1600m, 1958.VII.22, leg. Pu Fuji, IOZ(E)902224; 1♀, Yunnan, Xishuangbanna,, Mengzhe, 1200m, 1958.VIII.28, leg. Wang Shuyong, IOZ(E)902226; 1♂, Naban II/3D, 10.XI.2008, leg. L.Z. Meng, Yunnan, Jinghong, Naban River Nature Reserve Chachang (Forest), 2008.XI.10, 729m, 22.15810°N, 100.66543°E, leg. Meng Lingzeng; 1♂, Naban II/3D, 20.XI.2008, leg. L.Z. Meng, Yunnan, Jinghong, Naban River Nature Reserve Chachang (Forest), 2008.XI.20, 729m, 22.15810°N, 100.66543°E, leg. Meng Lingzeng, (IZAS).
China: Yunnan; Myanmar.
First record of Oreoderus maculipennis from Yunnan Province which confirmed the prediction of
Oreoderus is the largest genus in the tribe Valgini and easily distinguished by its covering of scales. The clypeus is usually rounded in front except for O. clypealis (the front margin straight and recurved), O. gestroi, O. gravis and O. waterhousei (with a process on the front margin). The pronotum is nearly trapezoidal, longer than wide. There are two pairs of carinae on the pronotum, the length of carinae varies among species. The middle carinae are moderately prominent except O. insularis, O. argillaceus (highly prominent, forming two tubercles) and O. coomani (only slightly prominent). The elytra are short and broad, similar morphologically and sometimes decorated with patches of scales (O. bidentatus, O. maculipennis and O. birmanus etc.). The pygidium is nearly triangular, covered with thick scales. Females of some species have a stylus on the hind margin (O. bidentatus, O. brevitasus).
The morphological variation of four characters (pronotum, protibia, elytra and aedeagus) was investigated based on 34 species (84 specimens) using geometric morphometrics. The shape information was extracted from the landmark data using the Procrustes fit. To see the variations, we used the principal component analysis (PCA). The first two PCs together accounted for 77.11%, 89.14%, 60.50% and 55.96% of the total variance in the analysis of pronotum, elytron, protibia and aedeagus, respectively. The main shape change of the pronotum was observed in the length/width ratio of the pronotum and the carinae (Fig.
Based on the PCA results, the morphological diversity in these four characters suggest they are not equally diagnostic in Oreoderus (Table
To extend the results of the comparative morphological analysis, a canonical variate analysis (CVA) of Chinese Oreoderus including the new species was conducted to quantitatively assess the differences among species, with a particular focus on differentiating the new species. The canonical variates scores of pronotum variables showed the 90% equal frequency ellipse, although there is some overlap between O. maculipennis and O. oblongus (Fig.
Most obtained p-values from permutation tests (10000 permutation rounds) for Procrustes distances based on the morphology of four characters (pronotum, elytra, protibia and aedeagus, respectively) among the seven species were smaller than 0.05 (Suppl. material
In this study, the taxonomy values of four characters (pronotum, elytra, protibia and aedeagus) were evaluated in Oreoderus. According the CVA analyses above, the studied specimens were clustered into seven groups for all four morphological characters. Comparisons among species were significantly different when quantified by Mahalanobis distances. This meant the morphological boundary of species based on the sample specimens from the Oreoderus species were distinctly separated from each other. In other words, the Oreoderus species could be statistically separated and determined based on Mahalanobis distances of the four characters. However, the average shape of these four characters was not always significantly different when measured by Procrustes distance, and may not be a useful metric for taxonomic evaluations of Oreoderus. Our results demonstrate that geometric morphometric analysis of external and internal characters can enhance species diagnosis in cryptic species.
The taxonomical value of the four characters examined was not equivalent. The aedeagus of Oreoderus is very distinctive in Valgini, with an overall longer and more robust shape. Also, the aedeagus contains the most morphological variation in Oreoderus, and provides the best character for taxonomic determination in this genus.
The GM analysis suggests that the pronotum and protibia can also be informative in the taxonomy of Oreoderus. The shape of pronotum in Oreoderus is nearly trapezoidal, longer than it is wide. The apical part is usually narrow and the lateral margin is sinuate. The main shape variations are the outline of pronotum, length and relative position of the carinae on the pronotum according to the result of PCA. The protibia of Oreoderus is usually short and flat, dentate with only one spur. The main variation of the protibia is on the external teeth according to the result of PCA.
The numbers of the external teeth of protibia are already used in the taxonomy of Valgini (
The elytra of Oreoderus are flat and covered with scales. The elytra contained the least morphological variation among four characters in this study. Additionally, the out groups were not separated from Oreoderus. The taxonomic value of the elytral shape is not highly supported.
Traditionally, discrete characters are commonly used in taxonomy. However it is often difficult to find enough discrete characters to resolve confusing taxonomic problem, such as the morphological convergence of Oreoderus. In such a case, geometric morphometric (GM) can been used (
We sincerely thank the museums and institutions that loaned the specimens for our research. Special thanks to Dr Johannes Frisch and Joachim Willers (ZMHB) for loaning specimens; Dr Olaf Jaeger (STMD), Dr Olivier Montreuil and Antoine Mantilleri (
This research was supported by the National Basic Research Program of China (973 Program, 2011CB302102); the National Natural Science Foundation of China (31172143, 51305057); and by a Humboldt Fellowship (M.B.) from the Alexander von Humboldt Foundation.
Figure A.1 Minimum spanning tree mapped onto a PCA plot; Table A.1–4 Difference in shapes of four characters among species.
Data type: calculation results