Review Article |
Corresponding author: Chi-Feng Lee ( chifeng@tari.gov.tw ) Academic editor: Alexander Konstantinov
© 2020 Chi-Feng Lee, Ron Beenen.
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, Beenen R (2020) Taiwanoshaira Lee & Beenen, a new genus and first record of moss-inhabiting Galerucinae sensu stricto (Coleoptera, Chrysomelidae) from Taiwan. ZooKeys 944: 129-146. https://doi.org/10.3897/zookeys.944.53099
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Taiwanoshaira Lee & Beenen gen. nov. is described. It represents the first genus of Galerucinae (s. str.) in Taiwan documented to inhabit moss cushions. Shaira chujoi Kimoto, 1982 is transferred to Taiwanoshaira, as follows: T. chujoi (Kimoto), comb. nov. Two new species, T. taipingshanensis sp. nov. and T. tsoui sp. nov., are described. Adults of T. taipingshanensis sp. nov. were observed feeding on the moss species Plagiomnium vesicatum (Besch.) T.J. Kop. (Mniaceae). Microhabitats and distribution of Taiwanoshaira species are discussed.
cloud forest, leaf beetle, Malaise trap, moss, nocturnal, taxonomy
Moss cushions constitute a special environment inhabited by a limited diversity of leaf beetles. Members of more than 50,000 known species of leaf beetles live mainly on the leaf surface of various flowering plants on which they feed.
The Taiwan Chrysomelid Research Team (TCRT) started their inventory of all species of Chrysomelidae during 2005. We found only adults of Ivalia Jacoby inhabited moss cushions early in the project. A TCRT colleague, Sin-Syue Li, found several galerucines (sensu stricto) inhabiting moss cushions at Yuanyang Lake (= Yuanyanghu, 鴛鴦湖) (Fig.
Habitat photographs. A microhabitat for Taiwanoshaira taipingshanensis sp. nov. and T. tsoui sp. nov. at Yuanyang Lake (鴛鴦湖) B active adults of T. chujoi comb. nov. inside moss cushions at Pilu (畢祿) C adult of T. taipingshanensis sp. nov. feeding on leaves of Plagiomnium vesicatum at Yuanyang Lake (鴛鴦湖) D adults of T. taipingshanensis sp. nov. mating at Yuanyang Lake (鴛鴦湖) E adult of T. tsoui sp. nov. at Hsiaofengkou (小風口) F adult of T. tsoui sp. nov. at Meifeng (梅峰).
These same moss-inhabiting galerucines were collected using Malaise traps by colleagues at the National Museum of Natural Science at Meifeng (梅峰), Yuanfeng (鳶峰), Hsiaofengkou (小風口), and Bilu Divine Tree (碧綠神木). Malaise traps are mostly used to collect flying insects and the base of the canvas (tent cloth) does normally not touch the ground. When the canvas touches the ground, then beetles that lack flying capacities can crawl upward. Adults of Taiwanese Paraplotes species are good examples of flightless insects collected with Malaise traps (
These galerucines were initially identified as Shaira chujoi Kimoto, 1982 and allied undescribed species. However, they were clearly different in diagnostic characters from the type species of the genus, S. maculata Maulik, 1936. Thus, generic placement of these species was re-evaluated, species diversity was analyzed, and the results are presented here.
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. For detailed examination a Nikon ECLIPSE 50i microscope was used.
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.
Specimens were available for study and deposited in the following institutions:
NMNS National Museum of Natural Science, Taichung, Taiwan [Jing-Fu Tsai];
RBCN Ron Beenen collection, Nieuwegein, The Netherlands;
TARI Applied Zoology Division, Taiwan Agricultural Research Institute, Taichung, Taiwan [Chi-Feng Lee].
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.
Taiwanoshaira tsoui Lee & Beenen, sp. nov.
Coloration (Figs
Head. Labrum trapezoidal, transverse, with about ten pairs of pores in a transverse row bearing pale, short or long setae, anterior margin medially depressed. Anterior part of head short, almost impunctate and glabrous, lined with setae along anterofrontal ridge. Compound eyes small, interantennal space 3.1–3.6× as wide as diameter of antennal insertion. Frontal tubercles transverse, subtriangular, slightly elevated, glabrous. Vertex smooth and glabrous. Antennae filiform, covered with dense setae, antennomere II subequal or a slightly shorter than III; similar in both sexes.
Pronotum 1.61–1.68 times as broad as long, lateral margins slightly rounded, basally narrowed. Disc smooth, with dense, fine punctures bearing tiny setae in T. taipingshanensis sp. nov. and T. chujoi comb. nov.; setae reduced in T. tsoui sp. nov. Anterior, lateral and posterior margins with marginal bead, without setae along margin. Anterior and posterior angles moderately swollen, rectangular; all angles with setigerous pores bearing long pale setae. Two pairs of longitudinal furrows starting from base, one pair deeper and shorter near middle, the other pair longer but shallow near sides. Scutellum subtriangular, impunctate, glabrous, with rounded apex.
Elytra ca 1.10–1.26 times as long as wide, almost glabrous, lateral margins rounded, apically tapering in males of T. chujoi comb. nov. and T. tsoui sp. nov., or both sexes of T. taipingshanensis sp. nov. Humeral calli reduced. Epipleura broad at base (Fig.
Ventral surface glabrous except abdomen, which is covered with pale setae. Anterior coxal cavities widely open (Fig.
Legs slender. Tibiae lacking apical spines. Protarsomeres I slightly swollen in males of T. taipingshanensis sp. nov., but unmodified in either sex of T. chujoi sp. nov. and T. tsoui sp. nov. Metatarsomeres I a little longer than pro- and mesotarsomeres I, subequal to II and III combined. Claws appendiculate.
Penis (Figs
Gonocoxae (Figs
Adults of Taiwanoshaira gen. nov. lack metathoracic wings, but elytra present, which completely cover the abdomen. Furthermore, the morphology of the elytra differs from that of Shaira. In Shaira the elytra possess an elongate ridge from the humeral area to the apex, dividing the elytra into a horizontal part and a lateral inclined part. The narrow epipleura are situated below this vertically inclined part of the elytra. In Taiwanoshaira gen. nov. the elytra possess a margin that separates the horizontal elytral surface from the epipleura, which are inclined.
The genus Shaira was proposed by
Taiwanoshaira gen. nov. is likely not closely related to Shaira, although also included in Luperini. It is unique and, although apterous, likely to be more closely related to the genus Sikkimia Duvivier, 1891 with all bordered pronotal margins, two pairs of longitudinal furrows starting from base, one pair deeper and shorter near middle, the other pair longer but shallow near sides. This new genus is easily separated from Sikkimia with the open procoxal cavities and uniform antennae in both sexes (the closed procoxal cavities and modified antennomeres X and XI in males of Sikkimia).
The new genus name combines “Taiwan” and “Shaira” to indicate that this is a new genus endemic to Taiwan that is similar to Shaira. The gender is feminine.
Taiwanoshaira chujoi (Kimoto), comb. nov., T. taipingshanensis sp. nov., and T. tsoui sp. nov.
Shaira chujoi Kimoto, 1982
Holotype
♂ (
(N = 68). Hualien: 1♀ (TARI), Bilu Divine Tree (碧綠神木), 10–11.X.2013, leg. K. Takahashi; 5♂♂, 4♀♀ (NMNS), same locality, 1.VI.-28.VII.2011; 2♂♂ (TARI), same locality (= Pilu), 13.VI.2014, leg. C.-F. Lee; 1♀ (TARI), same but with “leg T.-H. Lee”; 6♂♂, 12♀♀ (TARI), same locality, 30.VII.2014, leg. C.-F. Lee; 1♂ (TARI), same but with “leg. T.-H. Lee”; 8♂♂, 9♀♀ (TARI), same locality, 7.VII.2015, leg. C.-F. Lee; 4♂♂, 3♀♀ (TARI), Chian (吉安), 18.VI.2015, leg. T.-H. Lee; 1♂ (TARI), Kalapao (卡拉寶), 15–17.VII.2019, leg. B.-H. Ho; 4♂♂, 3♀♀ (3♂♂, 1♀TARI; 1♂, 2♀♀: RBCN), Kuanyuan (關原), 25.VIII.2014, leg. F.-S. Huang; 2♂♂, 1♀ (1♂, 1♀: TARI; 1♂: RBCN), Pilu (畢祿), 8.VIII.2014, leg. M.-H. Tsou; Nantou: 1♂ (NMNS), Meifeng (梅峰), 8.VII.-5.VIII.2003, leg. C. S. Lin & W. T. Yang; 1♂ (NMNS), Yuanfeng (鳶峰), 13.VIII.-10.IX.2002, leg. C. S. Lin & W. T. Yang; 1♂ Taitung: 1♂ (TARI), Liyuan (栗園), 5.X.2010, leg. T.-H. Lee.
Length 4.1–5.0 mm, width 2.5–3.3 mm. General color dark brown or blackish-brown (Fig.
Adults of Taiwanoshaira chujoi (Kimoto) comb. nov. are similar to those of T. tsoui sp. nov. based on the following shared characters: elytra smooth and lacking longitudinal ridges (Figs
Probably some species of moss, currently unknown (Fig.
(N = 77). Holotype ♂ (TARI): Taiwan. Ilan: Taipingshan (太平山), 5.VIII.2015, leg. Y.-T. Chung. Paratypes. 5♂♂, 15♀♀ (TARI), same as holotype; 10♂♂, 16♀♀ (TARI), Yuanyanghu (鴛鴦湖), 19.VIII.2010, leg. S.-S. Li; 4♂♂, 5♀♀ (TARI), same locality, 22.VIII.2011, leg. C.-F. Lee; 2♂♂, 8♀♀ (2♀♀: TARI; 2♂♂, 2♀♀: RBCN), same but with “leg. M.-H. Tsou”; 5♂♂, 6♀♀ (TARI), same but with “leg. H. Lee”.
Length 4.0–5.7 mm, width 2.6–3.4 mm. General color dark brown or blackish-brown (Fig.
Female genitalic characters are variable among different localities. The apices of the gonocoxae are widely rounded in specimens from Taipingshan (太平山) (Fig.
Diagnostic characters of Taiwanoshaira taipingshanensis sp. nov. A antenna, male B antenna, female C penis, apex, dorsal view D penis, base, dorsal view E penis, lateral view F endophallic sclerites, dorsal view G ditto, lateral view H gonocoxae, from Yuanyanghu (鴛鴦湖) I same, from Taipingshan (太平山) J abdominal ventrite VIII, from Yuanyanghu (鴛鴦湖) K same, from Taipingshan (太平山) L spermatheca.
Taiwanoshaira taipingshanensis sp. nov. is easily separated from other congeners by the presence of longitudinal ridges on the elytra (Fig.
Mniaceae: Plagiomnium vesicatum (Besch.) T.J. Kop. We observed that adults fed on leaves of host plants (Fig.
This new species is named for its type locality.
Known from two localities in northern Taiwan (Fig.
(N = 54). Holotype ♂ (TARI): Taiwan. Nantou: Hsiaofengkou (小風口), 9.VIII.2012, leg. C.-F. Lee. Paratypes. 14♂♂, 21♀♀ (12♂♂, 19♀♀TARI; 2♂♂, 2♀♀: RBCN), same data as holotype; 7♂♂, 6♀♀ (TARI), same but with “leg. T.-H. Lee”; 2♂♂, 2♀♀ (TARI), same locality, 29.VII.2014, leg. C.-F. Lee; 1♀ (NMNS), same locality, 23.VI.–24.VIII.2009, leg. W. T. Yang & K. W. Huang; 1♂, 3♀♀ (NMNS), same locality, 24.VIII.–24.IX.2009, leg. W. T. Yang & K. W. Huang; Ilan: 2♂♂, 1♀ (TARI), Taipingshan (太平山), 5.VIII.2015, leg. Y.-T. Chung; 1♀ (TARI), Yuanyanghu (鴛鴦湖), 19.VIII.2010, leg. S.-S. Li; Nantou: 1♀ (TARI), Meifeng (梅峰), 11.VI.2014, leg. C.-F. Lee; 1♂ (TARI), same locality, 29.VII.2014, leg. C.-F. Lee; 9♂♂, 2♀♀ (TARI), Peitungyanshan (北東眼山), 3.VII.2014, leg. C.-F. Lee; Taichung: 8♂♂, 4♀♀ (TARI), Tahsuehshan (大雪山), 2.VIII.2019, leg. B.-X. Guo.
Length 4.1–4.8 mm, width 2.5–2.9 mm. General color dark brown or blackish-brown (Fig.
Diagnostic characters of Taiwanoshaira tsoui sp. nov. A antenna, male B antenna, female C penis, dorsal view D penis, lateral view E endophallic sclerites, from Meifeng (梅峰), dorsal view F ditto, lateral view G same, from Peitungyanshan (北東眼山), dorsal H ditto, lateral view I samel, from Tahsuehshan (大雪山), J ditto, laeral view K same, form Taipingshan (太平山), dorsal L ditto, lateral view M gonocoxae, from Hsiaofengkou (小風口) N same, from Taipingshan (太平山) O abdominal ventrite VIII, from Peitungyanshan (北東眼山) P same, from Taipingshan (太平山) Q spermatheca.
Specimens from Hsiaofengkou (小風口) have paler bodies and shorter antenna than others. The endophallic spiculae complexes are variable among localities: subbases of endophallic spiculae are shorter and wider in specimens from Meifeng (梅峰) (Fig.
Adults of T. tsoui sp. nov. are similar to those of T. chujoi (Kimoto) comb. nov. in sharing the following characters: elytra smooth and lacking longitudinal ridges (Figs
Probably some species of moss, currently unknown (Fig.
This new species is dedicated to Mei-Hua Tsou, a member of TCRT and the first to collect this new species.
Northern and central Taiwan (Fig.
1 | Elytra with longitudinal ridges, apically narrowed in both sexes (Fig. |
T. taipingshanensis sp. nov. |
– | Elytra smooth, without longitudinal ridges, apically narrowed in males but widely rounded in females (Figs |
2 |
2 | Elytra entirely black (Fig. |
T. chujoi (Kimoto) comb. nov. |
– | Elytra with yellowish-brown sides and suture (Fig. |
T. tsoui sp. nov. |
Mosses are common all over Taiwan since it is a country with high humidity. They are most dominant in cloud forests. The montane cloud forest of Taiwan was mapped using 12-year MODIS-derived ground fog frequency data (
We would like to thank all curators listed above for giving us the opportunity to study the specimens from their collections. We thank the Taiwan Chrysomelid Research Team for assistance in collecting material, including Yi-Ting Chung, Bo-Xin Guo, Hsueh Lee, Ta-Hsiang Lee, and Mei-Hua Tsou. We especially thank Chi-Lung Lee, Hsueh, Lee and Mei-Hua Tsou for taking photographs of specimens and live individuals in the field. We thank 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.