Research Article |
Corresponding author: Chi-Feng Lee ( chifeng@tari.gov.tw ) Academic editor: Ron Beenen
© 2021 Chi-Feng Lee, Jan Bezděk.
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, Bezděk J (2021) Revision of the genera Xanthogaleruca Laboissière, 1932 and Pyrrhalta Joannis, 1865 (Coleoptera, Chrysomelidae, Galerucinae) of Taiwan, with type designation of Galerucella lineatipes Takei. ZooKeys 1039: 1-108. https://doi.org/10.3897/zookeys.1039.64740
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The taxonomic status of Pyrrhalta Joannis, 1865 and allied genera Tricholochmaea Laboissière, 1932 and Xanthogaleruca Laboissière, 1934 is discussed based on the study of Taiwanese species. Tentatively, Xanthogaleruca and Pyrrhalta are regarded as valid genera while Tricholochmaea is a synonym of Pyrrhalta. Fourteen species are recognized and redescribed, including P. gressitti Kimoto, 1969; P. taiwana Kimoto, 1969; P. viridipennis Kimoto, 1981; P. igai Kimoto, 1981; P. meifena Kimoto, 1976; P. maculata Gressitt & Kimoto, 1963; P. tsoui Bezděk & Lee, 2019; P. semifulva (Jacoby, 1885); P. discalis Gressitt & Kimoto, 1963; P. ishiharai Kimoto, 1994; P. shirozui Kimoto, 1969; P. kobayashii Kimoto, 1974; P. ohbayashii Kimoto, 1984; and P. takizawai Kimoto, 1996. Taiwanese populations identified as Xanthogaleruca aenescens (Fairmaire) were misidentified and those are described as a new species, X. yuae sp. nov. Xanthogaleruca aenescens is redescribed for comparison. Eight additional new species of Pyrrhalta are described: P. alishanensis sp. nov., P. houjayi sp. nov., P. formosanensis sp. nov., P. jungchani sp. nov., P. lui sp. nov., P. meihuai sp. nov., P. tahsiangi sp. nov., and P. wulaiensis sp. nov. Type specimens of Galerucella lineatipes Takei, 1916 were rediscovered and are designated as lectotype and paralectotype. Galerucella lineatipes is removed from synonymy with G. calmariensis (Linnaeus, 1767) and regarded as a senior synonym of P. humeralis (Chen, 1942), syn. nov. Most Pyrrhalta species can be classified into four species groups based on their morphological and genitalic similarity. host plants and other biological information are provided for almost all species.
host plant, leaf beetles, new species, new synonym, nomenclature, taxonomy, Tricholochmaea
The genus Pyrrhalta Joannis, 1865 is one of the most speciose genera of Galerucinae.
In Taiwan,
New species | Authority (reference) | New records or nomenclatural acts |
---|---|---|
P. gressitti, P. shirozui, P. taiwana | Kimoto, 1969 | P. aenescens (Fairmaire), P. humeralis (Chen), P. maculata Gressitt & Kimoto |
P. kobayashii | Kimoto, 1974 | P. semifulva Jacoby, P. discalis Gressitt & Kimoto |
P. meifena | Kimoto, 1976 | P. aurata (Maulik) |
P. igai, P. viridipennis | Kimoto, 1981 | |
P. ohbayashii | Kimoto, 1984 | |
P. ishiharai | Kimoto, 1994 | P. aurata (Maulik): misidentification |
P. takizawai | Kimoto, 1996 |
Taxonomic status of the genera Pyrrhalta and its allied genera is controversial. Tricholochmaea Laboissière and Xanthogaleruca Laboissière are regarded as distinct genera by some European and American taxonomists (e.g.,
The Taiwan Chrysomelid Research Team (TCRT) was founded in 2005 and is composed of ten members. All of them are amateurs interested in producing a complete inventory of chrysomelid species in Taiwan. Members of the genus Pyrrhalta have been collected and studied, and host plants recorded. Life histories for almost all species were documented by laboratory rearing. The results of these efforts are the subject of the current paper.
For rearing studies, larvae were placed in small glass containers (diameter 142 mm × height 50 mm) with cuttings from their host plants. When mature larvae began searching for pupation sites, they were transferred to smaller plastic containers (diameter 90 mm × height 57 mm) filled with moist soil (~ 80% of container volume).
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 examinations, a Nikon ECLIPSE 50i microscope was used.
At least three 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 studied herein are deposited at the following institutes and collections:
JBCB Jan Bezděk collection, Brno, Czech Republic;
HSC Haruki Suenaga collection, Okayama, Japan;
Exact 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, [g] – green label, [b] – blue label, and [r] – red label.
Galerucella (Xanthogaleruca)
Laboissière, 1934: 67 (type species: Chrysomela luteola Müller, 1766, by original designation);
Pyrrhalta (Xanthogaleruca): Wilcox, 1965: 36.
Xanthogaleruca: Silfverberg, 1974: 7;
Xanthogaleruca aenescens (Fairmaire, 1878), X. yuae sp. nov., and the additional ca. ten Palaearctic species (
Large sized species (7.9–9.5 mm). Antenna slender, antennomeres III–VII long (2.5–3.1 × longer than wide), VIII–X shorter. Body flattened (Fig.
Larvae and adults feed on leaves of Ulmus species and Zelkova serrata (Thunb.) Makino (Ulmaceae).
Tentatively we accept Xanthogaleruca as valid genus. Internal sclerite of aedeagus of Xanthogaleruca is characteristic, comb-like, and presumed to be an apomorphy (
Galeruca aenescens Fairmaire, 1878: 140 (China).
Galerucella aenescens:
Galerucella (Xanthogaleruca) aenescens:
Pyrrhalta aenescens: Gressitt & Kimoto, 1963: 443 (China: Jilin, Rehe, Hebei, Shandong, Jiangsu);
Pyrrhalta (Pyrrhalta) aenescens:
Pyrrhalta (Xanthogaleruca) aenescens:
Xanthogaleruca aenescens:
Presumably deposited at the
China. Beijing: 1♂, 1♀ (TARI), Wofosi (臥佛寺), 27.IV.1961, leg. S.-Y. Wang; Hebei: 8♂, 13♀ (TARI), 保定 (= Baoding), 5.IX.1943, leg. A. Tanaka; Tianjin: 1♂, 2♀ (JBCB), Wuquing Co., Dahuanqpu wetland natural conservation, 15.VII.2010, leg. P. Kment; Manchuria (outdated name, refers to Heilongjiang, Jilin, and Liaoning): 5♂ (TARI), 4♀ (TARI), Tokuniji, 23.VII.1937, leg. M. Hanano; 2♂, 2♀ (TARI), Mt. Riutan, Tolisu, 30.V.1937, leg. M. Hanano; 1♂, 3♀ (TARI), same but with “30.VII.1939”; 1♂ (TARI), Anto, 23.VII.1933, leg. K. Nomura.
Length 8.2–9.5 mm, width 3.9–4.5 mm. Body color (Fig.
Ulmaceae: Ulmus pumila Linnaeus, U. laevis Pallas, and U. davidiana Planch (
adults of X. aenescens (Fairmaire) and X. yuae sp. nov. may be separated from those of other species in the genus by the entirely green elytra, presence of three black spots on the pronotum, elytra with fine and dense punctures. Xanthogaleruca aenescens differs from X. yuae sp. nov. by the wider aedeagus, 5.1 × longer than wide (Fig.
Diagnostic characters of Xanthogaleruca aenescens (Fairmaire) A antenna, male B antenna, female C aedeagus, dorsal view D ditto, lateral view E gonocoxae F abdominal ventrite VIII G abdominal ventrite V, female H abdominal ventrite V, male I apex of tibia of middle leg, male J tarsi of front leg, male K tarsi of front leg, female L spermatheca.
Russia (Far East), Mongolia, North China (Gansu, Hebei, Henan, Hunan, Inner Mongolia, Jiangsu, Jilin, Shandong, Shanxi, Shaanxi;
Pyrrhalta aenescens: Kimoto, 1969: 28 (Taiwan);
Holotype
♂ (TARI), Taiwan. Taoyuan: Paling (巴陵), 27.V.2009 (reared from eggs), leg. M.-H. Tsou. Paratypes. 3♂, 6♀ (TARI), same data as holotype; 1♀ (TARI), same but with “25.V.2009”; 3♀ (TARI), same but with “26.V.2009”; 7♂, 8♀ (TARI), same but with “28.V.2009”; 37♂, 29♀ (TARI), same but with “29.V.2009”; 1♀ (TARI), same locality, 19.IV.2009, leg. S.-F. Yu; 1♀ (TARI), same locality, 19.VI.2010, leg. H.-J. Chen; Chiayi: 3♀ (TARI), Shounouryo (= Channaoliao, 樟腦寮), near Mt. Ari (阿里山), 14.XII.1937, leg. Y. Yano; 1♂ (TARI), Dokuritsuzan (= Tulishan, 獨立山), near Mt. Ari (阿里山), 14.XII.1937, leg. Y. Yano; Nantou: 2♂ (TARI), Lienhuachi (蓮華池), 23–26.V.1980, leg. K. S. Lin & B. H. Chen; 1♂ (
Body flattened. Pronotum with three large black spots, one in middle, two laterally. Elytra metallic green
Length 7.9–8.8 mm, width 3.3–3.8 mm. Body color (Fig.
Diagnostic characters of Xanthogaleruca yuae sp. nov. A antenna, male B antenna, female C aedeagus, dorsal view D ditto, lateral view E gonocoxae F abdominal ventrite VIII G abdominal ventrite V, female H abdominal ventrite V, male I tarsi of front leg, male J tarsi of front leg, female K apex of tibia of middle leg, male L spermatheca.
Adults of X. yuae sp. nov. and X. aenescens may be separated from those of other species in the genus by their entirely green elytra, presence of three black spots on the pronotum, and elytra with fine and dense punctures. Xanthogaleruca yuae sp. nov. differs from X. aenescens in having a narrower aedeagus, 5.7 × longer than wide (Fig.
Larvae and adults feed on leaves of Zelkova serrata (Thunb.) Makino (Ulmaceae) (present study).
Xanthogaleruca yuae sp. nov. populations are presumed to be univoltine. The following life cycle information is based on our (TCRT) observations made by Mr Mei-Hua Tsou (Lee and Cheng 2010). Females began to deposit an average of 10–20 eggs in two rows of a single egg mass on the undersides of leaves (Fig.
Widespread in lowlands of Taiwan.
Dedicated to Mrs Su-Fang Yu who was the first member of TCRT to collect specimens of this new species and rear them successfully from eggs to adults.
Pyrrhalta Joannis, 1865: 82 (type species: Galeruca vibruni Paykull, 1799).
Galerucella (Pyrrhalta): Weise, 1886: 621;
Galeruca (Pyrrhalta): Seidlitz, 1891: 705.
Decoomanius
Laboissière, 1927: 55 (type species: Decoomanius limbatus Laboissière, 1927; by monotypy). Synonymized by
Chapalia
Laboissière, 1929: 269 (type species: Chapalia jeanvoinei Laboissière, 1929; by monotypy). Synonymized by
Lochmaea (Tricholochmaea)
Laboissière, 1932: 963 (type species: Gallerucella semifulva Jacoby, 1885; by original designation). Synonymized by
Tricholochmaea: Chûjô & Kimoto, 1961: 169;
Pyrrhalta (Tricholochmaea): Wilcox, 1965: 37;
Pyrrhalta (Pyrrhalta): Wilcox, 1971: 84.
Pyrrhalta gressitti Kimoto, 1969; P. houjayi sp. nov.; P. tahsiangi sp. nov.; P. taiwana Kimoto, 1969; and P. viridipennis Kimoto, 1981.
Small to median sized species (3.5–7.8 mm). Antenna extremely slender, antennomeres III–VI long (3.1–4.5 × longer than wide), VII–X shorter. Body convex. Elytra relatively narrow, 1.6–1.8 × longer than wide. Aedeagus asymmetric, ostium covered by a membrane; endophallic sclerites composed of two slender sclerites (Figs
Larvae and adults feed on leaves of Rhododendron species or Vaccinium randaiense Hayata (Ericaceae).
Pyrrhalta gressitti
Kimoto, 1969: 25;
Pyrrhalta (Pyrrhalta) gressitti: Wilcox, 1971: 86.
Holotype
♀ (
Taiwan. Chiayi: 12♂, 3♀ (TARI), Alishan (阿里山), 5–9.VIII.1981, leg. L. Y. Chou & S. C. Lin; 2♀ (TARI), same locality, 17–20.VIII.1982, leg. K. C. Chou & C. C. Pan; 2♀ (
Length 3.9–5.4 mm, width 1.7–2.4 mm. Body color (Fig.
Habitus of Pyrrhalta gressitti Kimoto, P. houjayi sp. nov., and P. tahsiangi sp. nov. A P. gressitti, male, dorsal view B ditto, ventral view C ditto, lateral view D P. houjayi sp. nov., male, dorsal view E ditto, ventral view F ditto, lateral view G P. tahsiangi sp. nov., male, dorsal view H ditto, ventral view I ditto, lateral view.
adults of P. gressitti Kimoto and P. viridipennis Kimoto are characterized by their partly green elytra, which possess longitudinal ridges. However, P. gressitti can be separated from P. viridipennis by its smaller body sizes, 3.9–5.4 mm long (5.3–7.8 mm long in P. viridipennis), smooth and shining elytra, with coarse punctures (rough elytra with fine punctures in P. viridipennis); recurved apex of aedeagus and broadly rounded apex of primary endophallic sclerite lacking teeth (Fig.
Larvae and adults feed on leaves of Rhododendron rubropilosum var. rubropilosum Hayata (Ericaceae) (Fig.
The species is widespread at mid-altitudes (1,500–2,500 m) in southern Taiwan.
Holotype
1♂ (TARI), Taiwan. Pingtung: Lilungshan (里龍山), 30.VI.2016, leg. J.-C. Chen. Paratypes. 2♂♂, 3♀♀ (TARI), same data as holotype; Hsinchu: 1♂ (TARI), Talu trail (大鹿林道), 1.VIII.2015, leg. Y.-L. Lin; Kaohsiung: 1♂, 1♀ (TARI), Chungchihkuan (中之關), 1.VII.2009, leg. S.-F. Yu; 1♂, 2♀♀ (TARI), same locality, 3.VII.2009, leg. M.-H. Tsou; 1♂ (TARI), Shihshan logging trail (石山林道), 19.VIII.2008, leg. C.-T. Yao; 1♀ (
Elytra smooth, lacking longitudinal ridges; green with wide reddish brown band along suture.
Length 4.4–7.5 mm, width 2.5–3.1 mm. Body color (Fig.
Specimens from southern Taiwan possess a broader aedeagus and the broader endophallic sclerite near apex that is almost straight in lateral view.
Adults of P. houjayi sp. nov. and P. taiwana Kimoto are characterized by their partly green elytra lacking longitudinal ridges. Pyrrhalta houjayi sp. nov. can be distinguished from P. taiwana by presence of the wide brown band along the suture of the elytra, and more slender elytra (Figs
Larvae and adults feed on leaves of Rhododendron leptosanthum Hayata (Ericaceae)
The first author, Mrs Hsueh Lee, and Mr Hou-Jay Chen found young larvae feeding on tender shoots (Fig.
Field photographs of Pyrrhalta houjayi sp. nov. on host plant A young larvae feeding on tender shoots B host plant blooming and sprouting at the same time in Tahsuehshan (大雪山) C flower buds with holes caused by larvae D Larva found inside the flower buds E one larva feeding on pedicels F adults.
This new species is widespread at mid-altitudes (1,500–2,500 m) in Taiwan.
Dedicated to Mrs Su-Fang Yu who was the first member of TCRT to collect specimens of this new species and rear them successfully from eggs to adults.
Holotype ♂ (TARI), Taiwan. Ilan: Tsuifenghu (翠峰湖), 4.VII.2010, leg. M.-H. Tsou. Paratypes. 3♂, 8♀ (TARI), same data as holotype; Ilan: 6♀ (TARI), Yuanyanghu (鴛鴦湖), 23.VIII.2011, leg. M.-H. Tsou; 1♂, 5♀ (TARI), same but with “leg. H. Lee”; 7♀ (TARI), Taipingshan (太平山), 25.V.2009 (reared from larvae), leg. C.-F. Lee.
Elytra smooth, lacking longitudinal ridges; yellowish brown, with brown longitudinal stripes.
Length 4.8–5.6 mm, width 2.1–2.4 mm. Body color (Fig.
Diagnostic characters of Pyrrhalta tahsiangi sp. nov. A antenna, male B antenna, female C aedeagus, dorsal view D ditto, lateral view E abdominal ventrite VIII F apex of tibia of middle leg, male G spermatheca H tarsi of middle leg, male I abdominal ventrite V, female J abdominal ventrite V, male K gonocoxae.
Adults of this new species are easily separated from other members of the species group by their yellowish brown elytra bearing longitudinal brown stripes and modified tarsi of the middle leg. In addition, some putative autapomorphies are found in genitalic characters, including the characteristic shape of the aedeagus and single endophallic sclerite bearing teeth near the apex (Fig.
adults feed on leaves of Rhododendron chilanshanense Kurashige (Fig.
The first author and Mr Ta-Hsiang Lee collected young larvae on tender leaves of Rhododendron mariesii May 1, 2009 in Taipingshan, northeastern Taiwan, and reared them in the laboratory. Newly eclosed adults emerged from soil May 25. Mr. Mei-Hua Tsou collected a number of adults July 5, 2010 at the same locality (= Tsuifenghu) (Fig.
This new species is restricted to mid-altitudes (1,000–2,000 m) in northeastern Taiwan.
Dedicated to Mr. Ta-Hsiang Lee. He and the first author were the first to find larvae of this new species and rear them successfully to adults.
Pyrrhalta taiwana
Kimoto, 1969: 27 (Taiwan);
Pyrrhalta (Pyrrhalta) taiwana: Wilcox, 1971: 90 (catalogue).
Holotype
♂ (
Taiwan. Chiayi: 11♂, 7♀ (TARI), Tzuchung (自忠), 5.VI.2011 (reared from larvae), leg. C.-F. Lee; Kaohsiung: 2♂, 2♀ (TARI), Chungchihkuan (中之關), 3.VII.2009, leg. S.-F. Yu; 1♂, 2♀ (TARI), same locality, 1.VII.2019, leg. M.-H. Tsou; 3♂ (TARI), Tengchih (藤枝), 31.VII.2008, leg. C.-T. Yao; Taichung: 1♂ (TARI), Tahsuehshan (大雪山), 23.VII.2011, leg. J.-C. Chen; Taitung: 6♂, 7♀ (TARI), Lichia trail (利嘉林道), 15.VII.2014, leg. B.-X. Guo.
Length 5.6–7.0 mm, width 2.6–3.0 mm. Body color (Fig.
Adults of P. taiwana Kimoto and P. houjayi sp. nov. are characterized by their partly green elytra without longitudinal ridges. Pyrrhalta taiwana can be distinguished from P. houjayi sp. nov. by the entirely green elytra, except lateral margins, and wider elytra (Figs
Larvae and adults feed on leaves of Vaccinium randaiense Hayata (Fig.
Mrs Su-Fang Yu found adults feeding on leaves of Rhododendron leptosanthum July 3, 2009 in Chungchihkung, southern Taiwan. The first author and Mr. Mei-Hua Tsou found a number of larvae feeding tender leaves of Vaccinium randaiense May 9, 2011 in Tzuchung, southern Taiwan. These were reared in the laboratory. They began burrowing into soil May 12 and built underground chambers for pupation. The newly eclosed adults emerged from soil May 25.
This species is widespread at mid-altitudes (1,500–2,500 m) in southern Taiwan.
Pyrrhalta viridipennis
Kimoto, 1981: 2;
Holotype
♂ (
Taiwan. Chiayi: 1♂, 1♀ (TARI), Alishan (阿里山), 5–9.VIII.1981, leg. L. Y. Chou & S. C. Lin; Kaohsiung: 1♂ (TARI), Kuanshanyakou (關山啞口), 30.VII.2015, leg. C.-F. Lee; 1♂, 1♀ (TARI), Tengchih (藤枝), 7.IX.2012, leg. W.-C. Liao; 2♂, 5♀ (TARI), same but with “10.VIII.2013”; 1♀ (TARI), same but with “27.IX.2013”; 2♂, 4♀ (TARI), same but with “5.X.2013”; 4♂, 1♀ (TARI), same locality, 4.VIII.2012, leg. J.-C. Chen; 2♂, 2♀ (TARI), same locality, 30.VIII.2014, leg. B.-X. Guo; 2♀ (TARI), Tsuyunshan (出雲山), 25.IV.1990, leg. C. C. Chiang; Nantou: 1♀ (
Length 5.3–7.8 mm, width 2.3–3.5 mm. Body color (Fig.
Diagnostic characters of Pyrrhalta viridipennis Kimoto A antenna, male B antenna, female C aedeagus, from Alishan (阿里山), dorsal view D ditto, lateral view E apex of aedeagus, from Shihfen (十分), dorsal view F ditto, lateral view G apex of aedeagus, from Hsiangyang, dorsal view H ditto, lateral view I abdominal ventrite VIII J gonocoxae K abdominal ventrite V, female L abdominal ventrite V, male M apex of tibia of middle leg N spermatheca.
The apex of the aedeagus is variable among populations; more slender in north and central Taiwan (Fig.
Larvae and adults feed on leaves of Rhododendron rubropilosum var. rubropilosum Hayata (Ericaceae) (Fig.
The first author and Mr. Ta-Hsiang Lee collected one larva feeding on leaves April 20, 2010 in Tahsuehshan, central Taiwan (Fig.
Adults of P. viridipennis Kimoto and P. gressitti Kimoto are both characterized by the green elytra with longitudinal ridges. However, P. viridipennis differs from P. gressitti by the larger body sizes, 5.3–7.8 mm long (3.9–5.4 mm long in P. gressitti), rough elytra with fine punctures (smooth and shining elytra with coarse punctures in P. gressitti); curved apex of aedeagus and narrowly rounded apex of primary endophallic slerite with teeth (Fig.
The species is widespread at mid-altitudes (1,500–2,500 m) in central and southern Taiwan.
Pyrrhalta alishanensis sp. nov.; P. igai Kimoto, 1981; P. meifena Kimoto, 1976; and P. meihuai sp. nov.
Medium to large sized species (5.6–8.7 mm). Antenna stout, antennomeres VII-X shortest (1.5–2.2 × longer than wide), III–VI similar or slender. Body convex. Elytra relatively broad, 1.4–1.6 × longer than wide. Aedeagus apically tapering and symmetric (Figs
Larvae and adults feed on leaves of Acer species (Sapindaceae).
Holotype ♂ (TARI), Taiwan. Chiayi: Alishan (阿里山), 22.IV.2009, leg. M.-H. Tsou. Paratypes. 7♂, 11♀ (TARI), same data as holotype.
Medium-sized species, 7.3–8.7 mm. Body black. Elytra with fine dense punctures.
Length 7.7–8.7 mm, width 3.8–4.6 mm. Body black (Fig.
Adults of P. alishanensis sp. nov. are easily separated from other members of the species group by their black bodies (Fig.
Larvae and adults feed on leaves of Acer rubescens Hayata (Sapindaceae) (Fig.
Field photographs of Pyrrhalta alishanensis sp. nov., P. igai Kimoto, and P. meihuai sp. nov. on host plant A mature larva of P. alishanensis sp. nov. B adult of P. alishanensis sp. nov. C mature larva of P. igai D adult of P. igai E mature larva of P. meihuai sp. nov. F adult of P. meihuai sp. nov.
The first author and Mr Mei-Hua Tsou found more than 20 mature larvae (Fig.
Only known from the type locality.
Dedicated to the type locality, Alishan.
Pyrrhalta igai
Kimoto, 1981: 1;
Holotype
♂ (
Taiwan. Hsinchu: 1♂ (TARI), Talu trail (大鹿林道), 12.V.2018, leg. Y.-L. Lin; Kaohsiung: 3♂ (TARI), Neiyingshan (內英山), 5.V.2016, leg. B.-X. Guo; 1♀ (
Length 8.1–8.5 mm, width 4.0–4.5 mm. Body brown (Fig.
Adults of P. igai Kimoto are similar to those of Pyrrhalta meihuai sp. nov. in body sizes and color patterns (Fig.
Larvae and adults feed on leaves of Acer albopurpurascens Hayata (Sapindaceae).
The first author and Mr Mei-Hua Tsou found larvae feeding on leaves (Fig.
The collecting data of the label on the holotype is inconsistent with
The species is widespread at mid-altitudes (1,500–2,500 m) in Taiwan.
Pyrrhalta meifena
Kimoto, 1976: 4;
Holotype
♂ (
Taiwan. Chiayi: 1♂, 2♀ (TARI), Alishan (阿里山), 22.IX.2009, leg. M.-H. Tsou; 2♂♂ (TARI), same locality, 29.V.2016, leg. Y.-T. Chung; 1♂ (TARI), Shizilu (十字路), 23.VII.2015, leg. U. Ong; Hsinchu: 9♂, 2♀, Litungshan (李棟山), 24–25.III.2009, leg. M.-H. Tsou; 4♂, 3♀ (TARI), same locality, 28.III.2009, leg. S.-F. Yu; 1♀ (TARI), same locality, 16.VI.2010, leg. Y.-L. Lin; Hualien: 1♂, 2♀ (TARI), Kuanyuan (關原), 2.VII.2008, leg. S.-F. Yu and M.-H. Tsou; 1♀ (TARI), Pilu (碧綠), 15.VII.2019, leg. B.-X. Guo; 1♀ (TARI), Tayuling (大禹嶺), 2.VII.2018, leg. J.-C. Chen; Kaohsiung: 1♀ (TARI), Chungchihkung (中之關), 1.VII.2009, leg. S.-F. Yu; 1♀ (TARI), same locality, 2.VII.2009, leg. M.-H. Tsou; 1♀ (TARI), same locality, 17.IV.2012, leg. L.-P. Hsu; 1♂, 1♀ (TARI), Tienchih (天池), 2.VII.2009, leg. M.-H. Tsou; Nantou: 1♂, 2♀ (TARI), Meifeng (梅峰), 24–26.VI.1981, leg. K. S. Lin & W. S. Tang; 1♀ (TARI), same locality, 1–3.VIII.1981, leg. T. Lin & W. S. Tang; 6♂ (TARI), Tatachia (塔塔加), 13.VII.2014, leg. W.-C. Liao; 1♂ (TARI), Tsuifeng (翠峰), 25–27.VI.1981, leg. K. S. Lin & W. S. Tang; 1♀ (TARI), same locality, S. C. Lin & C. N. Lin; Taichung: 5♂, 7♀ (TARI), Anmashan (鞍馬山), 7.VI.2010, leg. C.-F. Lee; Taitung: 1♂ (TARI), Hsiangyang (向陽), 9.V.2013, leg. J.-C. Chen; 1♂ (TARI), same locality, 18.VII.2014, leg. W.-C. Huan; 2♀ (TARI), Liyuan (栗園), 19.VI.2013, leg. C.-F. Lee.
Length 5.6–6.5 mm, width 2.7–3.2 mm. Body yellow (Fig.
adults of P. meifena Kimoto are characterized by their small body sizes, 5.6–6.5 mm long (7.3–8.7 mm long in others), and yellow bodies (Fig.
Larvae and adults feed on leaves of Acer insulare var. caudatifolium (Hayata) and A. rubescens Hayata (Sapindaceae).
Mrs Su-Fang Yu found young larvae (Fig.
The species is widespread at mid-altitudes (1,500–2,500 m) in Taiwan.
Holotype ♂ (TARI), Taiwan. Ilan: Mingchi (明池), 2.V.2009, leg. M.-H. Tsou. Paratypes. 3♂, 3♀ (TARI), same data as holotype; 1♂, 3♀ (TARI), same but with “1.V.2009”; Pingtung: 1♂ (TARI), Tahanshan (大漢山), 18.VI.2012, leg. Y.-T. Chung; 1♂ (TARI), same but with “11.VII.2012”; 2♂, 1♀ (TARI), same but with “24.IV.2013”; 3♂, 5♀ (TARI), same but with “15.V.2013”; 2♂ (TARI), same but with “25.V.2013”; 2♀ (TARI), same but with “30.V.2013”; 3♀ (TARI), same but with “17.VI.2013”; 3♂ (TARI), same but with “2.VII.2013”; 1♂ (TARI), same but with “10.VII.2013”; 1♀ (TARI), same but with “30.VII.2013”; 1♀ (TARI), same but with “12.VI.2014”; 1♀ (TARI), same but with “4.VI.2020”; 1♀ (TARI), same locality, 19.VII.2012, leg. C.-F. Lee; 1♂ (TARI), Tahantrail (大漢林道), 20.VIII.2012, leg. J.-C. Chen; 1♀ (TARI), same but with “27.V.2013”; Taipei: 1♀ (TARI), Hsiungkungshan (熊空山), 15.VI.2014, leg. Y.-L. Lin; Taitung: 1♀ (TARI), Lichia (利嘉), 15.VII.2014, leg. Y.-T. Chung; 1♂ (TARI), same but with “16.VII.2014”; Taoyuan: 1♂ (TARI), Hsiaowulai (小烏來), 29.IX.2009, leg. M-H. Tsou; 1♀ (TARI), same locality, 1.VI.2010, leg. S.-F. Yu; 1♀ (TARI), Lalashan (拉拉山), 4.V.2009, leg. H.-J. Chen; 1♂ (TARI), Tungyanshan (東眼山), 12.VII.2015, leg. H. Lee.
Medium-sized species, 7.3–8.7mm. Body brown. Elytra with fine dense punctures. Discs of pronotum and elytra smooth, lacking reticulate microsculpture.
Length 7.3–8.6 mm, width 3.4–4.0 mm. Head and prothorax reddish brown (Fig.
Adults of P. meihuai sp. nov. are similar to those of P. igai Kimoto in body sizes and color patterns (Fig.
Larvae and adults feed on leaves of Acer serrulatum Hayata (Sapindaceae).
Mr Mei-Hua Tsou collected mature larvae (Fig.
The species is widespread at mid-altitudes (1,500–2,500 m) in Taiwan.
Dedicated to Mr. Mei-Hua Tsou. He, the first author, and Mr. Hou-Jay Chen were the first to collect larvae of this new species and rear them successfully to adults.
Pyrrhalta maculata Gressitt & Kimoto, 1963; P. tsoui Bezděk & Lee, 2019; P. formosanensis sp. nov.; P. semifulva (Jacoby, 1885); P. discalis Gressitt & Kimoto, 1963; P. ishiharai Kimoto, 1976; and P. wulaiensis sp. nov.
Small sized species (3.3–5.6 mm). Antenna stout, antennomeres VIII–X stout (1.4–2.0x longer than wide), III-VI slender. Body convex. Elytra relatively wider, 1.4–1.6 × longer than wide. Aedeagus asymmetric; ostium covered by a membrane or lacking cover; endophallic sclerites composed of two slender sclerites, with several teeth on apex of primary sclerite and with one additional tooth near apex of secondary sclerite except P. formosanensis sp. nov. with only primary sclerite (Fig.
Included species can be subdivided into species complexes based on similar color patterns. For example,
Anthophagous species. Larvae and adults feed on flowers of Meliosma rhoifolia (Sabiaceae) or species of Rosaceae.
Pyrrhalta maculata
Gressitt & Kimoto, 1963: 456;
Pyrrhalta (Pyrrhalta) maculata: Wilcox, 1971: 88 (catalogue).
(types examined by
(specimens examined by
Length 4.7–5.2 mm, width 2.3–2.5 mm. Body color (Fig.
Diagnostic characters of Pyrrhalta maculata Gressitt & Kimoto A antenna, male B antenna, female C aedeagus, dorsal view D ditto, lateral view E apex of tibia of middle leg, male F abdominal ventrite VIII G spermatheca H gonocoxae I abdominal ventrite V, female J abdominal ventrite V, male K tarsi of middle leg, male.
Adults of P. maculata Gressitt and Kimoto and P. tsoui Bezděk and Lee may be separated from others within the species group by the five pairs of large black spots on the elytra (Fig.
Possibly adults fed flowers of Lauraceae based on the following events. A specimen was collected by Mr Yi-Ting Chung 10 April 2016 in Sungkang by sweeping flowers of Lauraceae. Two specimens were collected by Mr Fu-Sheng Huang 16 September 2013 in Peitungyanshan by fogging Neolitsea aciculata var. variabillima J.C. Liao (Lauraceae).
China, Taiwan.
Pyrrhalta tsoui Bezděk & Lee, 2019: 531.
(specimens examined by
Length 4.6–5.3 mm, width 2.3–2.8 mm. Body color (Fig.
Adults of P. tsoui Bezděk & Lee and P. maculata Gressitt & Kimoto may be separated from others within the species group by the five pairs of large black spots on the elytra (Fig.
Adults feed on flowers of Meliosma rhoifolia Maxim. (Sabiaceae).
This species is widespread in lowlands of Taiwan.
Holotype ♂ (TARI), Taiwan. Kaohsiung, Tienchih (天池), 2.VII.2009, leg. M.-H. Tsou. Paratypes. 1♂, 12♀, same data as holotype; Hualien: 1♂ (TARI), Kuanyuan (關原), 2.VII.2008, leg. M.-H. Tsou; 1♂ (TARI), Pilu (碧綠), 6.VII.2018, leg. H.-F. Lu; Ilan: 1♀ (TARI), Chienching trail (見晴步道), 23.IV.2019, leg. M.-D. Chen; 1♀ (TARI), Tsuifenghu (翠峰湖), 15.VIII.2007, leg. S.-S. Li; Kaohsiung: 1♂ (TARI), Chungchihkuan (中之關), 10.VI.2015, leg. C.-F. Lee; Nantou: 1♂ (TARI), Meifeng (梅峰), 24–26.VI.1981, leg. K. S. Lin & W. S. Tang; 1♂ (TARI), Piluhsi (碧綠溪), 8.VII.2015, leg. C.-F. Lee; 1♂ (TARI), Tsuifeng (翠峰), 30.VII.2014, leg. C.-F. Lee.
Legs reddish brown; tibia of middle leg with apical spine; tarsomere I modified. Sides of ventrite V strongly shortened in males.
Length 4.6–5.5 mm, width 2.4–2.9 mm. Body color (Fig.
Diagnostic characters of Pyrrhalta formosanensis sp. nov. A antenna, male B antenna, female C aedeagus, dorsal view D ditto, left-side view E ditto, ventral view F ditto, right-side view G apex of tibia of middle leg, male H abdominal ventrite VIII I spermatheca J gonocoxae K abdominal ventrite V, male L abdominal ventrite V, female M tarsi of middle leg, male.
Adults of P. formosanensis sp. nov. are similar to those in Taiwanese populations of P. semifulva with their reddish bodies, but differ in the reddish brown scutellum, legs, and thoracic ventrites (Fig.
Adults feed on flowers of Prunus campanulata Maxim. (Rosaceae) (Fig.
The species is widespread at mid-altitudes (1,500–2,500 m) in Taiwan.
This species is named after Taiwan, a beautiful island.
Gallerucella semifulva Jacoby, 1885: 745 (Japan: Kiga).
Lochmaea (Tricholochmaea) semifulva:
Tricholochmaea semifulva: Chûjô & Kimoto 1961: 169 (catalogue);
Pyrrhalta semifulva:
Pyrrhalta (Tricholochmaea) semifulva:
Gallerucella modesta
Jacoby, 1885: 745 (Japan: Nikko). Synonymized by
Lochmaea (Tricholochmaea) modesta:
Gallerucella signaticeps
Weise, 1887: 191 (Vladivostok). Synonymized by
Lochmaea japonica
Weise, 1922: 67 (Japan: Honshu). Synonymized by
Gallerucella semifulva. Lectotype ♀ (
Gallerucella modesta. Lectotype (sex undetermined,
Japan. Hokkaido: 1♀ (HSC), Etetsu-shi, Nopporo, 18.VI.2011, leg. H. Suenaga; 1♀ (HSC), Tomakomai-shi, Lake Utonai-ko, 29.VIII.2011, leg. H. Suenaga; 1♂ (HSC), same but with “22.V.2012”; Honshu: 1♂, 1♀ (HSC), Akita Pref., Nikaho-shi, Chôkaisan, Hokodate, 10.VI.2016, leg. S. Sejima; 1♀ (TARI), Aomori Pref., 29.VI. 1934, leg. F. Watanabe; 1♀ (TARI), Aomori Pref., Hatinohe, 1.VI.1933, leg. A. Fukuda; 1♀ (
Length 4.3–5.4 mm, width 2.4–3.0 mm. Body color (Fig.
Diagnostic characters of Pyrrhalta semifulva (Jacoby) A antenna, male B antenna, female C aedeagus, typical form, dorsal view D ditto, lateral view E aedeagus, variation (endophallic sclerites omitted), dorsal view apex of tibia of middle leg, male F ditto, lateral view G abdominal ventrite VIII H ermatheca I abdominal ventrite V, female J abdominal ventrite V, male K gonocoxae.
Aedeagi of many individuals have apically tapering apices and look straight in lateral view (Fig.
Pyrrhalta semifulva (Jacoby) and P. formosanensis sp. nov. may be separated from others within the species group by the reddish brown bodies (Figs
Rosaceae: Prunus jamasakura Sieb., ex Koidz., P. yedoensis Matsum., and Sorbus japonica (Decne.) Hedl.; Hamamelidaceae: Corylopsis gotoana Makino, (
Japan, Russian, Taiwan. The species is widespread at mid-altitudes (1,500–2,500 m) in Taiwan.
Pyrrhalta discalis
Gressitt & Kimoto, 1963: 448 (China: Hubei);
Pyrrhalta (Pyrrhalta) discalis:
Holotype
♂ (
Taiwan. Hsinchu: 1♂ (TARI), Litungshan (李棟山), 15.III.2009, leg. S.-F. Yu; 1♀ (TARI), Lupi (魯壁), 25.II.2010, leg. S.-F. Yu; 1♀ (TARI), Wuchihshan (五指山), 27.III.2008, leg. H. Lee; 1♂ (TARI), same locality, 14.V.2008, leg. S.-F. Yu; Hualien: 1♀ (TARI), Pulowan (布洛灣), 26.III.2016, leg. H.-F. Lu; 1♂, 1♀ (TARI), same but with “31.III.2016”; 10♂, 4♀ (TARI), same but with “30.IV.2016”; 1♀ (TARI), same but with “9.V.2016”; Pingtung: 1♂ (TARI), Tahanshan (大漢山), 29.VI.2018, leg. Y.-T. Chung; Taichung: 1♀ (
Length 4.6–5.6 mm, width 2.3–2.8 mm. Body color (Fig.
Taiwanese populations display great variation of color patterns on the elytra. Some individuals have two additional transverse black stripes (Fig.
adults of P. discalis Gressitt and Kimoto are easily recognized by the yellowish brown bodies. In males of P. discalis, the elongate and apically curved aedeagus is similar to that of P. semifulva (Jacoby), but differs by the relatively shorter secondary endophallic sclerite, 0.6 × as long as primary endophallic sclerite (Fig.
Larvae and adults feed on flowers of Pourthiaea lucida Decne. (Fig.
eggs (Fig.
China, Taiwan. It is widespread at lowlands (0–1,500 m) in Taiwan.
Pyrrhalta aurata: Kimoto, 1976: 4 (Taiwan). Misidentification (after
Pyrrhalta ishiharai
Kimoto, 1994: 191;
Holotype
♀ (
Taiwan. Hsinchu: 1♂, 1♀ (TARI), Chienshih (尖石), 10.VII.2010, leg. M.-H. Tsou; 1♂ (TARI), same locality, 5.VIII.2012, leg. Y.-L. Lin; Nantou: 1♂ (
Length 4.8–5.1 mm, width 2.3–2.5 mm. Body color (Fig.
Diagnostic characters of Pyrrhalta ishiharai Kimoto A antenna, male B antenna, female C aedeagus, dorsal view D ditto, lateral view E apex of tibia of middle leg, male F abdominal ventrite VIII G spermatheca H tarsi of middle leg, male I abdominal ventrite V, female J abdominal ventrite V, male K gonocoxae.
Adults of P. ishiharai Kimoto and P. wulaiensis sp. nov. are easily separated from other species within the species group by the longitudinal ridges on the elytra (Fig.
Adults feed on flowers of Meliosma rhoifolia Maxim. (Sabiaceae) (Fig.
The species is widespread at lowlands (0–1,500 m) in Taiwan.
Holotype
♂ (TARI), Taiwan. Nantou: Peitungyanshan (北東眼山), 3.VII.2014, leg. F.-S. Huang, 變葉新木薑子 (Neolitsea aciculata var. variabillima J.C. Liao) 噴霧 (fogging). Paratypes. 1♂ (TARI), same locality as holotype, 3.VII.2014, leg. C.-F. Lee; Ilan: 1♂ (TARI), Fushan (福山), 5.VII.2013, leg. Y.-T. Wang; Miaoli: 1♂ (TARI), Hsuehchien (雪見), 5.III.2013, leg. W.-B. Yeh; Nantou: 1♀ (TARI), Meifeng (梅峰), 28–29.VIII.1981, leg. L. Y. Chou & S. C. Lin; 1♀ (TARI), same locality, 15.VII.1982, leg. S. C. Lin & C. N. Lin; 1♀ (
Smaller species, 3.3–3.7 mm in length. Elytra relatively broad, 1.5 × longer than wide; unicolorous, without dark spots; with ridges.
Length 3.3–3.7 mm, width 1.6–1.9 mm. Body color (Fig.
Adults of P. wulaiensis sp. nov. and P. ishiharai Kimoto are easily separated from other species within the species group by the longitudinal ridges on the elytra (Fig.
Larvae and adults feed on flowers of Meliosma rhoifolia Maxim. (Sabiaceae).
One female was collected on flowers of the host plant (Fig.
The species is widespread at lowlands (0–1,500 m) in northern Taiwan and mid-altitudes (1,500–2,500 m) in central Taiwan.
The species is named for the locality where specimens were collected and used for laboratory rearing.
Pyrrhalta jungchani sp. nov.; P. lui sp. nov.; and P. shirozui Kimoto, 1969.
adults small to medium sized (3.3–6.8 mm). Antenna slender, antennomere III longest, V–X similar in size. Body convex. Elytra relatively wider for P. shorozui 1.5 × longer than wide (Fig.
Habitus of Pyrrhalta jungchani sp. nov. and P. shirozui Kimoto A P. jungchani sp. nov., male, dorsal view B ditto, ventral view C ditto, lateral view D P. shirozui, female, dorsal view E ditto, ventral view F ditto, lateral view G Same species, color variation H Same species, color variation I Same species, color variation.
Larvae and adults feed on leaves of Viburnum species (Adoxaceae).
Holotype
♂ (TARI), Taiwan. Pingtung, Tahantrail (大漢林道), 30.VII.2012, leg. J.-C. Chen. Paratypes. Chiayi: 1♀ (TARI), Zengwen Reservoir (曾文水庫), 2.IV.2016, leg. U. Ong; Nantou: 2♀ (
Small species, 4.3–5.0 mm in length. Pronotum with three large black spots, one at middle, two laterally. Elytra relatively narrow, 1.7 × longer than wide, disc with dense coarse punctures, with black stripes at humeral calli, with one additional pair of longitudinal dark stripes between humeral calli and suture.
Length 4.3–5.0 mm, width 1.9–2.3 mm. Body yellowish brown (Fig.
Diagnostic characters of Pyrrhalta jungchani sp. nov. A antenna, male B antenna, female C aedeagus, dorsal view D ditto, lateral view E abdominal ventrite VIII F apex of tibia of middle leg, male G gonocoxae H tarsi of middle leg, male I abdominal ventrite V, female J abdominal ventrite V, male K spermatheca.
Adults of P. jungchani sp. nov. (Fig.
adults feed on leaves of Viburnum odoratissimum Ker Gawl. (Adoxaceae) (Fig.
The species is widespread at mid-altitudes (1,500–2,500 m) in central and southern Taiwan.
Dedicated to Mr Jung-Chan Chen who was the first member of TCRT to collect specimens of this new species.
Holotype ♂ (TARI), Taiwan. Hualien: Hahuan Cross-Ridge (合歡越嶺古道), 4.VIII.2018, leg. H.-F. Lu. Paratypes. 16♂, 7♀ (TARI), same data as holotype; Hualien: 3♀ (TARI), Hutoushan (虎頭山), 21.IV.2018, leg. H.-F. Lu; Kaohsiung: 1♀ (TARI), Chungchihkuan (中之關), 17.IV.2012, leg. L.-P. Hsu; 4♂, 3♀ (TARI), same locality, 12.VI.2015, leg. C.-F. Lee; Miaoli: 1♂ (TARI), Hsuehchien (雪見), 7.VI.2013, leg. W.-B. Yeh; Nantou: 1♂ (TARI), Chingching (清境), 5.III.2007, leg. H.-C. Chen; 1♂ (TARI), Meifeng (梅峰), 2–4.VI.1980, leg. L. Y. Chou & C. C. Chen; 1♀ (TARI), same locality, 24–26.VI.1981, leg. K. S. Lin & W. S. Tang; 1♂ (TARI), Tatachia (塔塔加), leg. 21.VI.2009, leg. C.-F. Lee; Taichung: 1♂, 2♀ (TARI), Kukuan (谷關), 21.III.2014, leg. B.-X. Guo.
Elytra relatively narrow, 1.7–1.8 × longer than wide, entirely yellowish brown or black; disc smooth, lacking ridges; with sparse, fine punctures
Length 4.6–5.3 mm, width 2.0–2.4 mm. Body yellow, head and pronotum reddish brown, antenna blackish brown except ventral sides of five basal antennomeres, bases of femora and lateral margins of tibia black; tarsi darker in females (Fig.
Males of P. lui sp. nov. display great variation in color. Some are totally black except for yellowish brown legs and abdomens (Fig. F, G); a few individuals are mainly black but pronota are reddish brown (Fig.
Adults of P. lui sp. nov. are distinguished within the species group by unicolorous elytra. In males, the aedeagus (Fig.
Adults feed on leaves of Viburnum parvifolium Hayata (Fig.
The species is widespread at mid-altitudes (1,500–2,500 m) in central and southern Taiwan.
Dedicated to Mr Hsi-Feng Lu, the member of TCRT who collected most specimens of this new species.
Pyrrhalta shirozui
Kimoto, 1969: 26 (Taiwan);
Pyrrhalta (Pyrrhalta) shirozui:
Holotype
♀ (
Taiwan. Chiayi: 1♂ (TARI), Yushan (玉山), 1.VII.2015, leg. J.-C. Chen; Hualien: 2♀ (TARI) Hahuan Cross-Ridge (合歡越嶺古道), 4.VIII.2018, leg. H.-F. Lu; 1♂ (TARI), Pilu (碧綠), 29.VI.2018, leg. H.-F. Lu; Ilan: 6♂, 3♀ (TARI), Mingchi (明池), 25.V.2008, leg. M.-H. Tsou; 4♂, 2♀ (TARI), same but with “16.VIII.2008”; 1♀ (TARI), Ssuyuan (思源), 11.VIII.2014, leg. J.-C. Chen; 1♂ (TARI), Taipingshan (太平山), 26–28.VII.1983, leg. L. Y. Chou; 1♂, 1♀ (TARI), same locality, 8.VII.2008, leg. H.-J. Chen; 3♂, 6♀ (TARI), same locality, 25.V.2009, leg. C.-F. Lee; 1♂ (TARI), Yingtzuling (鶯仔嶺), 3.VI.2011, leg. Y.-L. Lin; Nantou: 1♀ (TARI), Meifeng (梅峰), 5–9.X.1980, leg. C. C. Chen & C. C. Chien; 1♂, 1♀ (TARI), Nengkaoshan (能高山), 18.X.2011, leg. J.-C. Chen; 1♀ (TARI), Tatachia (塔塔加), 20.VII.2009, leg. S.-F. Yu; 1♀ (TARI), same but with “leg. H. Lee”; 1♀ (TARI), same but with “C.-F. Lee”; 1♂ (TARI), same locality, 21.IX.2009, leg. C.-F. Lee; 3♂♂, 2♀♀ (TARI), Tsuifeng (翠峰), 12–14.IX.1984, leg. K. S. Lin and S. C. Lin; Pingtung: 8♂♂ (TARI), Jinshuiying (浸水營), 12.VIII.2010, leg. J.-C. Chen; 1♀ (TARI), Tahanshan (大漢山), 1.VIII.2009, leg. U. Ong; 1♀ (TARI), same locality, 19.VII.2012, leg. C.-F. Lee; 1♂ (TARI), same locality, 29.VI.2018, leg. Y.-T. Chung; Taichung: 1♂ (TARI), Hassenzan (= Pahsienshan, 八仙山), 4.VI.1942, leg. A. Mutura; 1♂ (TARI), Wuwoweishan (屋我尾山), 5.VI.2012, leg. J.-C. Chen; Taipei: 2♂♂ (TARI), Fengkueitsui (風櫃嘴), 21.X.2007, leg. M.-H. Tsou; 6♂, 3♀ (TARI), Hsiaoyukeng (小油坑), 24.V.2008, leg. M.-H. Tsou; 4♀ (TARI), same but with “12.X.2008”; 9♀ (TARI), same locality and collector, reared from larvae, 21–29.III.2009; 1♀ (TARI), 5.XI.2006, Shihlin (士林), 5.XI.2006, leg. H.-T. Cheng; 2♂, 6♀ (TARI), Yangmingshan (陽明山), 12.V.2007, leg. M.-H. Tsou; 1♀ (TARI), same but with “27.V.2007”; Taitung: 3♂, 1♀ (TARI), Hsiangyang (向陽), 2.VII.2009, leg. S.-F. Yu; 1♀ (TARI), Liyuan (栗園), 19.VI.2013, leg. C.-F. Lee; 1♀ (TARI), Motien (摩天), 23.V.2011, leg. C.-F. Lee; Taoyuan: 4♂, 10♀ (TARI), Lalashan (拉拉山), reared form larvae, 27.IV.2009, leg. C.-F. Lee; 1♂, 9♀ (TARI), same but with “28.V.2009”; 1♀ (TARI), same locality, 15.VII.2009, leg. H.-J. Chen; 1♀ (TARI), Tamanshan (塔曼山), 25.VIII.2008, leg. H. Lee.
Length 4.9–6.8 mm, width 2.4–3.4 mm. Body color (Fig.
Diagnostic characters of Pyrrhalta shirozui Kimoto A antenna, male B antenna, female C aedeagus, dorsal view D ditto, lateral view E apex of tibia of middle leg, male F abdominal ventrite VIII G gonocoxae H spermatheca I abdominal ventrite V, female J abdominal ventrite V, male K tarsi of middle leg, male.
Some specimens have a black stripe along the entire suture of the elytra (Fig.
adults of P. shirozui Kimoto are easily recognized by the characteristic color patterns on the elytra and sparse, coarse elytral punctures, as well as diagnostic shape of the aedeagus differing from all other species of Pyrrhalta.
Larvae and adults feed on leaves of Viburnum formosanum (Hance) Hayata, V. foetidum var. rectangulatum Rehder, V. integrifolium Hayata, V. luzonicum Rolfe, V. taitoense Hayata, and V. urceolatum Siebold and Zucc.
The following life cycle information is based on Mr Mei-Hua Tsou’s (TCRT) observations (Lee and Cheng 2010). Females deposited single eggs in crevices of small twigs (Fig.
This species is widespread in lowlands (0–1,500 m) in northern Taiwan and mid-altitudes (1,500–2,500 m) in central Taiwan.
Pyrrhalta kobayashii
Kimoto, 1974: 25;
Holotype
♀ (
Taiwan. Nantou: 1♀ (TARI), Huakang (華岡), 20.VII.2017, leg. J.-C. Chen; Taichung: 1♀ (TARI), Pilu (畢祿), 2.VII.2008, leg. M.-H. Tsou; Taitung: 1♀ (TARI), Hsiangyang (向陽), 12.VII.2012, leg. J.-C. Chen.
(females). Length 6.2–6.3 mm, width 3.2 mm. Body yellow (Fig.
Habitus of Pyrrhalta kobayashii Kimoto and P. lineatipes (Takei) A P. kobayashii, female, dorsal view B ditto, ventral view C ditto, lateral view D P. lineatipes, lectotype, dorsal view E P. lineatipes, paralectotype, dorsal view F P. lineatipes, type labels G P. humeralis, from Taiwan, female, dorsal view H ditto, ventral view I ditto, lateral view.
The color pattern of adults of P. kobayashii (Fig.
Unknown.
The species occurs at scattered localities at mid-altitudes (1,500–2,500 m) in central and southern Taiwan.
Galerucella lineatipes Takei, 1916: 35 (Japan: Gumma).
Galerucella humeralis Chen, 1942: 17 (China: Guanxi, Liaoning). syn. nov.
Pyrrhalta humeralis: Nakane & Kimoto, 1961: 21 (Japan: Okinawa island);
Pyrrhalta (Pyrrhalta) humeralis:
Gallerucella lineatipes. Lectotype ♂ (
Galerucella humeralis. Presumably deposited at the
China. Fujian: 1♀ (
Length 6.0–7.9 mm, width 2.9–4.1 mm. Body color (Fig.
Adults of P. lineatipes (Takei) (Fig.
Mr. Takei sent specimens to Dr. Matsumura for identification. He wrote a new species name on the identification card, Galerucella lineatipes sp. n., but that name was never published. Later,
Although Pyrrhalta lineatipes feed on leaves of Viburnum spp., it does not belong to the P. shirozui species group due to a number of apomorphies in adults and arrangement of eggs. Pyrrhalta lineatipes differs from members of the P. shirozui species group with its symmetrical aedeagus (Fig.
Viburnum sp. (
The overwintering eggs of P. lineatipes were deposited into the twigs of the hostplants (Fig.
China (Anhui, Fujian, Gansu, Guandong, Guanxi, Helongjiang, Hubei, Hunan, Jiangxi, Jilin, Liaoning, Shaanxi, Sichuan, Zhejian;
Pyrrhalta ohbayashii
Kimoto, 1984: 46;
Holotype
♀ (
Taiwan. Kaohsiung: 1♀ (
Length 4.5–4.6 mm, width 1.9–2.1 mm. Body color (Fig.
Adults of P. ohbayashii Kimoto (Fig.
Adults feed on leaves of Prunus phaeosticta var. phaeosticta (Hance) Maxim. (Fig.
The species is widespread at lowlands (0–1,500 m) in northern and southern Taiwan.
Pyrrhalta takizawai
Kimoto, 1996: 32;
Holotype
♀ (
Taiwan. Hsinchu: 1♂ (TARI), Feifengshan (飛鳳山), 5.III.2009, leg. S.-F. Yu; 1♀ (TARI), Kuanhsi (關西), 21.VI.2009, leg. W.-T. Liu; 4♂, 8♀ (TARI), same locality, 24.VII.2010, leg. H. Lee; 2♂, 1♀ (TARI), Peitelaman (北德拉曼), 26.VI.2008, leg. H. Lee; 1♂ (TARI), Shihlu trail (石鹿古道), 23.VIII.2014, leg. Y.-L. Lin; 1♀ (TARI), Talu trail (大鹿林道), 26.VIII.2012, leg. Y.-L. Lin; 1♂ (TARI), Wufeng (五峰), 17.III.2009, leg. S.-F. Yu; 1♂ (TARI), Tahunshan (大混山), 1.III.2009, leg. M.-H. Tsou; Ilan: 2 ♂ (JBCB,
Diagnostic characters of Pyrrhalta ohbayashii Kimoto A antenna, male B antenna, female C aedeagus except apex, dorsal view D apex of aedeagus, dorsal view E aedeagus, lateral view F abdominal ventrite VIII G apex of tibia of middle leg, male H gonocoxae I spermatheca J abdominal ventrite V, female K abdominal ventrite V, male L tarsi of middle leg, male.
Length 10.4–12.3 mm, width 4.3–5.4 mm. Body dark brown or blackish brown (Fig.
Adults of P. takizawai Kimoto are similar to those of P. igai Kimoto and P. meihuai sp. nov. in having large, brown bodies but differ by the sparse pubescence on the pronotum (vs. dense pubescence on pronotum in P. igai and P. meihuai sp. nov.), sparse, coarse punctures on elytra (vs. dense, coarse punctures on elytra in P. meihuai sp. nov.; sparse, fine punctures on elytra in P. igai). The form of the aedeagus, gonocoxae, and female abdominal ventrite VIII are also diagnostic.
Larvae and adults feed on leaves of Celtis sinensis Pers. (Cannabaceae).
Adults were collected from Taipei City Zoo, January 19, 2008 and transferred to the laboratory for rearing. Females began to deposit an average of 10–20 eggs in single egg mass (Fig.
The species is widespread at lowlands (0–1,500 m) in Taiwan.
1 | Antenna extremely slender, antennomeres III–V more than 3.0 × longer than wide | 2 |
– | Antenna long or stout, antennomeres III–V less than 3.0 × longer than wide | 8 |
2 | Antennae and legs black; elytra yellow with black margins | 3 |
– | Antennae and legs yellowish brown; part of elytra green, or yellowish brown elytra with brown longitudinal stripes | 4 |
3 | Elytra with dense, fine punctures, and black stripes along suture; tibiae entirely black (Fig. |
P. kobayashii Kimoto |
– | Elytra with sparse, coarse punctures, black stripes and spots variable; tibiae yellowish brown with lateral margin black (Fig. |
P. shirozui Kimoto |
4 | Elytra at least partly green, without brown longitudinal stripes | 5 |
– | Elytra yellowish brown, with brown longitudinal stripes (Figs |
P. tahsiangi sp. nov. |
5 | Elytra with longitudinal ridges, apically brown | 6 |
– | Elytra smooth, lacking longitudinal ridges, apices green | 7 |
6 | Elytra with coarse punctures and sparse pubescence (Fig. |
P. gressitti Kimoto |
– | Elytra with fine punctures and dense pubescence (Fig. |
P. viridipennis Kimoto |
7 | Elytra green with yellow lateral margin (Fig. |
P. taiwana Kimoto |
– | Elytra green with wide brown band along suture (Figs |
P. houjayi sp. nov. |
8 | Pronotum with three large black spots, one at middle, two laterally | 9 |
– | Pronotum without black spots | 13 |
9 | Body flattened; elytra metallic green (Fig. |
X. yuae sp. nov. |
– | Body convex; elytra brown, reddish brown, or dark brown | 10 |
10 | Body reddish brown; elytra with five pairs of black spots, one pair near base, two pairs near middle, two pairs at apical 1/3 (Fig. |
11 |
– | Body brown or dark brown, elytra with black stripes at humeral calli | 12 |
11 | Antennomere III elongate, 4.5 × longer than wide, apically expanded in males (Fig. |
P. maculata Gressitt & Kimoto |
– | Antennomere III short, 2.7–3.4 × longer than wide, antennomere IV with small tubercle in males (Fig. |
P. tsoui Bezděk & Lee |
12 | Small species, 4.3–5.0 mm in length; elytra relatively narrow, 1.7 × longer than wide, disc with dense coarse punctures, with one additional pair of longitudinal dark stripes between humeral calli and suture (Fig. |
P. jungchani sp. nov. |
– | Large species, 6.0–7.9mm in length; elytra relative broad, 1.5 × longer than wide, disc with sparse fine puncture, lacking longitudinal dark stripes between humeral calli (Fig. |
P. lineatipes (Takei) |
13 | Smaller species, less than 6.5 mm in length | 14 |
– | Larger species, more than 6.5 mm in length | 21 |
14 | Elytra with ridges | 15 |
– | Elytra smooth, lacking ridges | 17 |
15 | Elytra with regular dark spots between ridges (Fig. |
P. ishiharai Kimoto |
– | Elytra unicolorous, without dark spots | 16 |
16 | Smaller species, 3.3–3.7 mm in length; elytra relatively broad, 1.5 × longer than wide (Fig. |
P. wulaiensis sp. nov. |
– | Larger species, 4.5–4.6 mm in length; elytra relatively narrow, 1.7–1.8 × longer than wide (Fig. |
P. ohbayashii Kimoto |
17 | Elytra relatively narrow, 1.7–1.8 × longer than wide, entirely yellowish brown or black, disc with sparse, fine punctures (Fig. |
P. lui sp. nov. |
– | Elytra relatively broad, 1.4–1.6 × longer than wide, entirely reddish brown, or yellowish brown with black margin and suture, disc with dense, coarse punctures | 18 |
18 | Body entirely reddish brown | 19 |
– | Elytra yellow or partly yellow | 20 |
19 | Legs reddish brown (Fig. |
P. formosanensis sp. nov. |
– | Legs black (Fig. |
P. semifulva (Jacoby) |
20 | Elytra entirely yellowish brown (Fig. |
P. meifena Kimoto |
– | Elytra yellowish brown with black margin and suture, sometimes black band along suture enlarged or with additional transverse black bands (Fig. |
P. discalis Gressit & Kimoto |
21 | Larger species, 10.4–12.3 mm; elytra with sparse coarse punctures (Fig. |
P. takizawai Kimoto |
– | Smaller species, 7.3–8.7mm; elytra with dense fine punctures | 22 |
22 | Body black (Fig. |
P. alishanensis sp. nov. |
– | Body brown | 23 |
23 | Discs of pronotum and elytra with reticulate microsculpture (Fig. |
P. igai Kimoto |
– | Discs of pronotum and elytra smooth, lacking reticulate microsculpture (Fig. |
P. meihuai sp. nov. |
The taxonomic relationship of Pyrrhalta, Tricholochmaea, and Xanthogaleruca has been controversial for many decades.
Tricholochmaea was described by
The Pyrrhalta genus complex badly requires comprehensive revision based on molecular data of species from the whole distributional area. The revision of Taiwanese species supports inclusion of Tricholochmaea as part of the Pyrrhalta semifulva species group within Pyrrhalta. This species group also comprises maculate species traditionally classified in Pyrrhalta (cf.
Some characters presumed to be important for generic diagnosis are not supported by the present study. The apical spur of the middle leg in males appears across whole genus and species groups, or in some species within different groups, including Xanthogaleruca; Pyrrhalta gressitti, P. tahsiangi sp. nov., and P. viridipennis within the P. gressitti species group; P. maculata, P. tsoui, P. formosanensis sp. nov., and P. ishiharai within the P. semifulva species group; and P. jungchani sp. nov. and P. shirozui within the P. shirozui species group. Some of these species have tarsomere I of the middle leg modified, including P. tahsiangi sp. nov. within the P. gressitti species group; P. maculata, P. formosanensis sp. nov., and P. ishiharai within the P. semifulva species group; P. jungchani sp. nov. and P. shirozui within the P. shirozui species group. Groups based on other morphological characters such as the ratio of length vs. width for each antennomere and elytra; sizes and genitalic characters in both sexes are more diagnostic for sorting species within the genus. Such groupings are corroborated by phylogenetic relationships of host plants and shared feeding behaviors. Members of Xanthogalerucae feed on leaves of Ulmus species or Zelkova serrata (Ulmaceae), those of the Pyrrhalta gressitti species group feed on leaves of leaves of Rhododendron species or Vaccinium randaiense (Ericaceae), those of the P. meifena species group feed on leaves of Acer species (Sapindaceae), those of the P. semifulva species group feed on flowers of Meliosma rhoifolia (Sabiaceae) or species of Rosaceae, and those of the P. shirozui species group feed on leaves of Viburnum species (Adoxaceae). This suggests that information about host plants and feeding behaviors may be helpful in grouping species of Pyrrhalta.
Species richness of Pyrrhalta may be underestimated based on the following reasons. Most Pyrrhalta species are monophagous; for example, four species of the P. meifena species group feed on Acer species (Sapindaceae), of which six species are found in Taiwan (
We are grateful to the Taiwan Chrysomelid Research Team (TCRT) for assistance in collecting material, including Jung-Chang Chen, Hou-Jay Chen, Yi-Ting Chung, Bo-Xin Guo, Hsueh Lee, Wen-Chuan Liao, His-Feng Lu, Mei-Hua Tsou, and Su-Fang Yu. We especially thank Chi-Lung Lee, and Hsing-Tzung Cheng for photos of specimens, Hsueh Lee, Ta-Hsiang Lee, His-Feng Lu, Mei-Hua Tsou, and Su-Fang Yu for their field photography, Chih-Kai Yang for identification of host plants, and Haruo Takizawa for providing information about P. lineatipes (Takei). This study was supported by the Ministry of Science and Technology MOST 109-2313-B-055-003. We 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.