Research Article |
Corresponding author: Chi-Feng Lee ( chifeng@tari.gov.tw ) Academic editor: Alexander Konstantinov
© 2020 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 (2020) Revision of Taiwanese species of Atrachya Chevrolat, 1836 (Coleoptera, Chrysomelidae, Galerucinae): descriptions of three new genera, two new species, and designations of three new synonyms. ZooKeys 940: 117-159. https://doi.org/10.3897/zookeys.940.51800
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The genus Atrachya Chevrolat is redefined based on study of the type species A. menetriesii (Faldermann, 1835). All Taiwanese species of Atrachya are transferred to three new genera: A. hirashimai Kimoto, 1976 and A. nitidissima (Chûjô, 1935) are transferred to Neochya gen. nov.; A. mediofasciata Kimoto, 1976 is transferred to Tsouchya gen. nov.; A. unifasciata Takizawa, 1978 is transferred to Chinochya gen. nov. Two species are described: N. chengi sp. nov. and N. tsoui sp. nov. Atrachya bicoloripennis (Chûjô, 1938) and A. saramao (Chûjô, 1962) are regarded as synonyms of N. nitidissima (Chûjô, 1935) comb. nov., and Monolepta tsoui Lee, 2009 is synonymized with T. mediofasciata (Kimoto, 1976) comb. nov. Monolepta sublata Gressitt & Kimoto, 1963 is redescribed and transferred to Chinochya gen. nov. Taiwanese records of Monolepta sublata are based on misidentifications and represent specimens of C. unifasciata. Variablity of adult color patterns is discussed.
Chinochya, leaf beetles, Neochya, new genus, taxonomic revision, Tsouchya
The genus Atrachya Chevrolat, 1836 is attributed to “Monoleptites” sensu
Diagnostic characters of the genus Atrachya were discussed (Wagner and Bieneck 2017) based on Cnecodes bisignatus Motschulsky, 1858. However, one important character, the sexually dimorphic elytral impression, does not occur in any Taiwanese species. This paper redefines the genus Atrachya based on the type species, A. menetriesii (Faldermann, 1835). The taxonomic status of the Taiwanese species of Atrachya is reevaluated, and their identities reviewed. In addition,
The Taiwan Chrysomelid Research Team (TCRT) has been inventorying chrysomelid fauna since 2005. Adults of the taxa covered in this paper were collected by sweeping host plants, species of Celastrus and Euonymus (Celastraceae) (Fig.
The abdomens of adults were separated from the bodies and boiled in 10% KOH solution, followed by washing in distilled water to clear and soften genitalia. The genitalia were then dissected from the abdomen, mounted on slides in glycerin, and studied and drawn using a Leica M165 stereomicroscope. A Nikon ECLIPSE 50i microscope was used for detailed examination.
At least two pairs from each species were examined to delimit variability of diagnostic characters. For species collected from more than one locality, at least one pair from each locality 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. Descriptions are all based on adult specimens.
Specimens were available for study and deposited in the following institutions:
NMNS National Museum of Natural Science, Taichung, Taiwan [Jing-Fu Tsai];
Exact label data are cited for all type specimens of previously 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, [r] – red label, [b] – blue label.
Habitat photographs A adult of Neochya chengi sp. nov. feeding on leaves of Celastrus hindsii B adult of N. hirashimai C Celastrus kusanoi, food plant for Neochya species D adult of N. nitidissima feeding on leaves of Euonymus spraguei E adult of N. tsoui resting on underside of leaves of Euonymus spraguei F Euonymus spraguei, food plant for Neochya species.
Atrachya Chevrolat, 1836: 401 (type species: Galleruca menetriesii Faldermann, 1835, by monotypy).
Iphidea
Baly, 1865: 127 (type species: Iphidea discrepens Baly, 1865, by original designation) (= Galleruca menetriesii Faldermann, 1835. Synonymized by
Cnecodes
Motschulsky, 1858: 99 (type species: Cnecodes bisignatus Motschulsky, 1858, by
(Faldermann, 1835). Japan. Hiroshima: 1♂ (
Atrachya is a distinct genus similar to Paleosepharia Laboissière (redefined based on type species by
Excluding Taiwanese species, 21 species in the African, Palaearctic, and Oriental regions (
Atrachya hirashimai Kimoto, 1969
Coloration : extremely variable but without metallic color. Body length 4.6–7.0 mm.
Head. Labrum trapezoidal, transverse, with six pores in transverse row bearing pale setae, anterior margin medially depressed. Anterior part of head very short, almost impunctate and glabrous, several setae on anterior margin of clypeus and anterofrontal ridge. Interantennal space broad, 1.5–2.6× as wide as diameter of antennal insertion. Frontal tubercles transverse, subtriangular, slightly elevated, glabrous. Vertex smooth and glabrous. Antennae slender, covered with dense setae, antennomere II as long as antennomere III; similar in both sexes.
Pronotum 1.75–2.00 times as broad as long, lateral margins slightly rounded, basally narrowed. Disc covered with dense, coarse punctures, moderately or strongly convex, with lateral depressions, except N. chengi sp. nov. and N. nitidissima. Anterior margin lacking marginal bead, lateral and posterior margins with marginal bead. Anterior and posterior margins without setae, lateral margins with two pairs of setae near base and apex respectively. Anterior angles moderately swollen, rectangular, posterior angles obtuse angulate, all angles with setigerous pore bearing long pale seta.
Scutellum subtriangular, impunctate, glabrous, with rounded apex.
Elytra
ca 1.35–1.68 times as long as wide, almost glabrous (with indistinct, sparse, short, pale setae on humeri, lateral margins and apical slopes), parallel sided, except N. nitidissima (Fig.
Ventral
surface sparsely covered with fine punctures and pale setae. Anterior coxal cavities widely open (Fig.
Legs slender. All tibiae with one apical spine, the longest spine on metatibia. Protarsomeres I not modified in males. Metatarsomeres I much longer than pro- and mesotarsomeres I, much longer than II and III combined. Claws appendiculate.
Penis
broad, with one pair of small lateral processes near apex (Figs
Gonocoxae
(Figs
Neochya gen. nov. differs from Atrachya Chevrolat and Monolepta Chevrolat in the following combination of characters: antennomere II subequal to III in length (antennomere II much shorter than III in Atrachya (Fig.
Composed from new and Atrachya to indicate that this is a new genus similar to Atrachya.
Neochya chengi sp. nov., N. hirashimai (Kimoto), comb. nov., N. nitidissima (Chûjô) comb. nov., and N. tsoui sp. nov.
(N = 49). Holotype ♂ (
Length
4.2–5.2 mm, width 2.3–3.0 mm. General color reddish brown (Fig.
Most of specimens from Tahanshan have a distinct color pattern on elytra (Fig.
Neochya chengi sp. nov. is similar to N. nitidissima (Chûjô) in having wide elytra, truncate elytral apices and reduced lateral depressions on the pronotum (Figs
Celastraceae: Celastrus hindsii Benth (Fig.
It is named after Mr. Hsing-Tzung Cheng who was a member of the TCRT and an editor for a series of the books entitled “The Chrysomelidae of Taiwan”. The gender is feminine.
Atrachya hirashimai
Kimoto, 1969: 54;
Holotype
♂ (
(N = 81). Chiayi: 4♂♂, 4♀♀ (
Length
5.1–6.2 mm, width 2.4–3.1 mm. General color yellowish brown (Fig.
A distinct color pattern occurs in beetles from the east part of South Cross-Island Highway (南橫公路): general color black; but apical 3/4 of elytra and abdomen yellowish brown (Fig.
Neochya hirashimai (Kimoto) is similar to N. tsoui sp. nov. in having slender elytra and lateral depressions on the pronotum (Figs
Celastraceae: Celastrus kusanoi Hayata (Fig.
Restricted to several places in central Taiwan. Two color patterns are separated in the eastern and western parts of the range (Fig.
Luperodes nitidissimus
Chûjô, 1935: 161;
Atrachya nitidissima: Kimoto, 1966: 30 (additional records);
Luperodes bicoloripennis
Chûjô, 1938: 138;
Atrachya bicoloripennis: Kimoto, 1969: 55: (additional records);
Atrachya bicolor [sic!]: Kimoto, 1989: 257 (additional records).
Luperodes saramao Chûjô, 1962: 230. syn. nov.
Atrachya saramao: Kimoto, 1989: 257 (additional records).
Luperodes nitidissimus. Holotype ♀ (
Luperodes bicoloripennis. Lectotype ♂ (
Luperodes saramao. Holotype ♂ (
Form A (N = 84). Chiayi: 1♂, 1♀ (
Form B (N = 255). Hsinchu: 1♀ (
Form C (N = 85). Hsinchu: 1♂ (
Form D (N = 44). Chiayi: 4♂♂, 1♀ (
Form E (N = 1). Pingtung: 1♂ (
Length
5.6–7.0 mm, width 2.9–3.8 mm. General color extremely variable, with five distinct color patterns (see variation). Antennae (Fig.
Color pattern divided into four forms. Form A (Fig.
Habitus of Neochya nitidissima A form A, female, from Alishan (阿里山), dorsal view B same, ventral view C form B, male, from Kuanwu (觀霧), dorsal view D same, ventral view E form C, female, from Pilu (碧綠), dorsal view F same, ventral view G form D, female, from Taikang trial (泰崗林道), dorsal view H same, ventral view I form E, male, from Peitawushan (北大武山), dorsal view.
Neochya nitidissima (Chûjô) is similar to N. chengi sp. nov. in having wide elytra, truncate elytral apices and reduced lateral depressions on the pronotum (Figs
The holotype of Luperodes nitidissimus was described as a male (
Celastraceae: Celastrus kusanoi Hayata, C. hindsii Benth., Euonymus spraguei Hayata (Fig.
Widespread in Taiwan. Most individuals with different color patterns can be separated based on distributions (Fig.
(N = 49). Holotype ♂ (
Length
4.6–5.5 mm, width 2.1–3.0 mm. General color reddish brown or yellowish brown (Fig.
Neochya tsoui sp. nov. is similar to N. hirashimai (Kimoto) in having slender elytra and lateral depressions on the pronotum (Figs
Celastraceae: Euonymus japonicus Thunb., E. spraguei Hayata (Fig.
This new species is dedicated to Mei-Hua Tsou, a member of TCRT and the first to collect this new species.
Widespread but scattered in Taiwan (Fig.
Atrachya mediofasciata Kimoto, 1976.
Color extremely variable (Fig.
Head. Labrum trapezoidal, transverse, with six pores in transverse row bearing pale seta, anterior margin truncate. Anterior part of head short, almost impunctate and glabrous, four setae on anterior margin of clypeus and several setae along anterior margin of anterofrontal ridge. Interantennal space narrow, 1.0–1.2× as wide as diameter of antennal insertion. Frontal tubercles transverse, slightly reduced, glabrous. Vertex smooth and glabrous. Antennae slender, covered with dense setae, antennomere II much shorter than antennomere III (0.61–0.68× as long as III); similar in both sexes (Fig.
Habitus of Tsouchya mediofasciata A female, from Shihshan trail (石山林道), dorsal view B male, from Lilungshan (里龍山), dorsal view C same, ventral view D female, from Lilungshan (里龍山), dorsal view view E male, from Shihshan trail (石山林道), dorsal view F female, from Shihshan trail (石山林道), dorsal view G same, dorsal view H male, from Talu trail (大鹿林道), dorsal view I female, from Meifeng (梅峰), dorsal view.
Pronotum 1.52–1.56 times as broad as long, lateral margins straight, basally narrowed. Disc covered with dense coarse punctures, moderately convex. Posterior half of disc with wide shallow transverse impression. Anterior margin lacking marginal bead, lateral and posterior margins with marginal bead. Anterior and posterior margins without setae, lateral margins with two pairs of setae near base and apex, respectively. Anterior angles moderately swollen, rectangular, posterior angles obtuse angulate, all angles with setigerous pores bearing long pale setae.
Scutellum subtriangular, impunctate, glabrous, with rounded apex.
Elytra 1.55–1.61 times as long as wide, almost glabrous (with indistinct, sparse, short, pale setae on humeri, lateral margins and apical slopes), parallel-sided, densely covered with coarse confused punctures. Humeral calli well developed. Epipleura broad at base, strongly narrowed at basal 1/3, abbreviated from apical 1/3 to apex. Macropterous.
Ventral
surface sparsely covered with fine punctures and pale setae. Anterior coxal cavities closed (Fig.
Legs slender. All tibiae with one apical spine, the longest spine on metatibia. Protarsomeres I not modified in males. Metatarsomeres I much longer than pro- and mesotarsomeres I, much longer than II and III combined. Claws appendiculate.
Penis
(Fig.
Gonocoxae
(Fig.
Tsouchya gen. nov. differs from Neochya gen. nov., Monolepta Chevrolat and Atrachya Chevrolat based on the following combination of characters: antennomere II much short than III in length (Fig.
Composed from Tsou and Atrachya to honor Mei-Hua Tsou, who is a member of TCRT (Taiwan Chrysomelid Research Team) and made great contributions to inventorying the chrysomelid fauna in Taiwan. The gender is feminine.
Tsouchya mediofasciata (Kimoto), comb. nov.
Atrachya mediofasciata
Kimoto, 1976: 6;
Monolepta tsoui Lee, 2009: 23. syn. nov.
Monolepta bicavipennis: Kimoto, 1969: 50 (Taiwan). non Monolepta bicavipennis Chen, 1942.
Atrachy mediofasciata. Holotype ♂ (
Kaohsiung: 1♂, 2♀♀ (
This species is described in detail as Monolepta tsoui by
Coloration
(Fig.
Head. Labrum trapezoidal, transverse, with six pores in transverse row bearing pale setae, anterior margin truncate. Anterior part of head short, almost impunctate and glabrous, four setae on anterior margin of clypeus and several setae along anterior margin of anterofrontal ridge. Interantennal space narrow, 0.8–0.9× as wide as diameter of antennal insertion. Frontal tubercles transverse, slightly reduced, glabrous. Vertex smooth and glabrous. Antennae slender, covered with dense setae, antennomere II subequal to III in length; similar in both sexes.
Pronotum 1.62–1.69 times as broad as long, lateral margins rounded, basally narrowed. Disc covered with dense, fine punctures, moderately convex, without transverse impression. Anterior margin lacking marginal bead, lateral and posterior margins with marginal bead. Anterior and posterior margins without setae, lateral margins with two pairs of setae near base and apex, respectively. Anterior angles moderately swollen, rectangular, posterior angles obtuse angulate, all angles with setigerous pores bearing long pale setae.
Scutellum subtriangular, impunctate, glabrous, with rounded apex.
Elytra 1.37–1.61 times as long as wide, almost glabrous (with indistinct, sparse, short, pale setae on humeri, lateral margins and apical slopes), parallel-sided, densely covered with coarse, confused punctures. Humeral calli well developed. Epipleura broad at base, strongly narrowed at basal 1/3, abbreviated from apical 1/3 to apex. Macropterous.
Ventral
surface sparsely covered with fine punctures and pale setae. Anterior coxal cavities almost closed (Fig.
Legs slender. All tibiae with one apical spine, the longest spine on metatibia. Protarsomeres I swollen in males (Figs
Penis
(Figs
Gonocoxae
(Figs
Diagnostic characters of Chinochya sublata A antenna, male B antenna, female C aedeagus, dorsal view D aedeagus, lateral view E aedeagus, ventral view F abdominal ventrite VIII G gonocoxae H dorsal bursal sclerites I ventral bursal sclerites J protarsi, male K protari, female L spermatheca.
Chinochya gen. nov. differs from Tsouchya gen. nov., Neochya gen. nov., Atrachya Chevrolat, and Monolepta Chevrolat based on the following combination of characters: antennomere II subequal to III in length (Figs
Composed from China and Atrachya to indicate the locality of the type species. The gender is feminine.
Chinochya sublata (Gressitt & Kimoto) comb. nov. and C. unifasciata (Takizawa) comb. nov.
Monolepta sublata Gressitt & Kimoto, 1963: 635.
Paratype
: 1♀ (
China. Fujian: 1♀ (
Length
5.3–6.3 mm, width 3.1–3.7 mm. Head, prothorax, and abdomen yellowish brown; antennae black except two basal antennomeres yellowish brown; meso- and metathoracic ventrites black; front legs yellowish brown, but tibia and tarsi darkened; middle and hind legs black; elytra black with one transverse, broad white band at middle (Fig.
Chinochya sublata is similar to C. unifasciata. They cannot be separated based on their external morphology, however, C. sublata (Fig.
Males are here described for the first time. Types on which the original description was based are all females (
South China (Fujian, Sichuan, Yunnan).
Atrachya unifasciata Takizawa, 1978: 132.
Monolepta sublata: Kimoto, 1976: 6 (Taiwan). non Monolepta sublata Gressitt & Kimoto, 1963
Holotype
♂ (
Taiwan. Hsinchu: 1♂ (
Length
4.9–5.8 mm, width 2.6–3.0 mm. Head, prothorax, and abdomen yellowish brown; antennae black except two basal antennomeres yellowish brown; meso- and metathoracic ventrites black; front legs yellowish brown, but tibiae and tarsi darker; middle and hind legs black; elytra black with one transverse, broad, white band at middle (Fig.
Diagnostic characters of Chinochya unifasciata A antenna, male B antenna, female C aedeagus, dorsal view D aedeagus, lateral view E aedeagus, ventral view F abdominal ventrite VIII G gonocoxae H dorsal bursal sclerites I ventral bursal sclerites J protarsi, male K protari, female L spermatheca.
Chinochya unifasciata is similar to C. sublata. They cannot be separated based on their external morphology, however, C. unifasciata (Fig.
Widespread but scattered in Taiwan.
1 | Subscutellar groove on the elytra present in males | Paleosepharia Laboissière |
– | Subscutellar groove on the elytra absent in males | 2 |
2 | Tarsomere I of front legs swollen in males (Fig. |
Chinochya gen. nov. |
– | Tarsomere I not modified in males | 3 |
3 | Antennomere III longer than II | Tsouchya gen. nov. |
– | Antennomere III subequal or smaller than II | 4 |
4 | Prothoracic coxal cavities closed (Fig. |
Monolepta Chevrolat |
– | Prothoracic coxal cavities open (Fig. |
Neochya gen. nov. 5 |
5 | Elytra slender (Figs |
6 |
– | Elytra wide (Figs |
7 |
6 | Pronotum and elytra reddish brown (Fig. |
N. tsoui sp. nov. |
– | Pronotum reddish brown and elytra yellowish brown; or pronotum black and elytra with basal half black and apical half yellowish brown (Fig. |
N. hirashimai (Kimoto) |
7 | Elytra parallel sided; punctures on pronotum and elytra coarse | N. chengi sp. nov. |
– | Elytra with lateral margins rounded; punctures on pronotum reduced and punctures on elytra fine | N. nitidissima (Chûjô) |
A number of genera of Monoleptites in Sundaland with elongate metatarsomeres I were reevaluated and redefined recently, together with new genera, including Arcastes Baly, 1865 (
Reliable diagnostic characters for the supraspecific taxonomy of Monoleptites with elongate metatarsomeres I have been limited. Color patterns are useful for most Oriental genera, but not diagnostic for east Palaearctic genera. This character in most species of Neochya and Tsouchya is variable, and in N. nitidissima and T. mediofasciata it is extremely variable. Although color patterns of Chinochya species are similar, some species of Monolepta share these patterns, including M. leechi Jacoby, 1890 and M. maana Gressitt & Kimoto, 1963.
Prothoracic coxal cavities have been used for diagnosis of genera within this group of Monoleptites. This character was evaluated in Atrachya menetriesii, all species of Neochya, Chinochya, Tsouchya, Taiwanese species of Paleosepharia, and Monolepta. It can be separated into three states: widely open for Atrachya (Fig.
Elytral epipleurae of Paleosepharia (Fig.
I would like to thank all curators listed above for giving us the opportunity to study the specimens from their collections, and the Taiwan Chrysomelid Research Team for collecting materials, including Hou-Jay Chen, Jung-Chang Chen, Yi-Ting Chung, Bo-Xin Guo, Hseh Lee, Wen-Chuan Liao, Mei-Hua Tsou, and Su-Fang Yu. I am grateful to Haruki Suenaga and Takuya Takemoto for providing specimens of Atrachya menetriesii, and Chih-Kai Yang for identification of food plants. I especially thank Chang Chin Chen for assisting our study in various ways and Chris Carlton for reading the draft and editing for American English style. This study was supported by the Ministry of Science and Technology (MOST 107-2313-B-055-002).