Research Article |
Corresponding author: Wen-Bin Yeh ( wbyeh@nchu.edu.tw ) Academic editor: Michael Schmitt
© 2016 Chi-Feng Lee, Cheng-Lung Tsai, Alexander Konstantinov, Wen-Bin Yeh.
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, Tsai C-L, Konstantinov A, Yeh B-W (2016) Revision of Mandarella Duvivier from Taiwan, with a new species, new synonymies and identities of highly variable species (Insecta, Chrysomelidae, Galerucinae, Alticini). ZooKeys 568: 23-49. https://doi.org/10.3897/zookeys.568.7125
|
Taiwanese species of Mandarella Duvivier are compared on the basis of morphological and molecular evidence. Only three of eleven morphospecies are considered to be valid. Mandarella uenoi (Kimoto, 1969) is transferred from the genus Luperus Geoffroy. Stenoluperus taiwanus Kimoto, 1991 and S. kimotoi Döberl, 2001 are synonymized with M. uenoi. Taiwanese records of Stenoluperus tibialis Chen, 1942, S. nipponensis Laboissière, 1913, and S. potanini (Weise, 1889) were based on misidentifications and represent M. uenoi. The Taiwanese population previously erroneously identified as S. pallipes Gressitt and Kimoto, 1963 is here described as a new species, M. tsoui sp. n., Stenoluperus esakii Kimoto, 1969, S. matsumurai Takizawa, 1978, and M. taiwanensis Medvedev, 2012 are synonymized with M. flaviventris (Chen, 1942).
Flea beetles, alpine, molecular, taxonomic revision
Mandarella Duvivier, 1892 is a small genus of flea beetles containing five species (
More than 250 mountains exceed 3000 meters in Taiwan. Localities higher than 3000 m present cold and windy montane habitats (Figs
Although these Mandarella species can be separated by color patterns and relative lengths of antennomeres (
Exact label data are cited for type specimens and voucher ones of the described species; a double slash (//) divides the data on different labels and a single slash (/) divides the data in different rows. Other comments and remarks are in square brackets: [p] – preceding data are printed, [h] – preceding data are handwritten, [w] – white label, [y] – yellow label, [b] – blue label, [r] – red label, and [y] – yellow label.
Approximately 2500 specimens were examined for this study. Most of them either belong to the historic collections at
Genomic DNA was extracted from the meta-femora via QuickExtract DNA extraction kits (Epicenter Biotechnologies, Madison, WI). The protocol was modified according to
Gene | Primer | Sequence 5’→3’ | Size (bp) | References |
---|---|---|---|---|
COI | COI-Chry_F (+) | ACYAAYCAYAAAGAYATWGG | 689 | In this study |
COI-49_Chrysomelidae_F | CATAAAGATATTGGHACHTT | 683 | ||
COI-64_Chrysomelidae_F | ACHYTRTAYTTYATTTTYGG | 668 | ||
CI-731Coleoptera (-) | CCAAAAAATCAAAATAAATGTTG |
|
Sequences were edited in Bioedit 7.0 (
Phylogenetic inference of COI was conducted using neighbor-joining clustering (NJ) and Bayesian inference (BI). For NJ, Kimura two-parameter (K2P) was selected as the substitution model and 1000 replications of bootstrapping analyses were applied. For BI, the evolutionary hypothesis of nucleotide substitution inferred in jModelTest 0.1 (Posada 2008) using Bayesian Information Criterion (BIC) for the best-fit models of COI was TIM3 + I + G. BI of the COI gene was analyzed in MrBayes v. 3.2 (
A total of 131 specimens of Mandarella flea beetles were successfully amplified for COI, with 584 bp in this study. The average nucleotide compositions for G, A, T, C are 18.4%, 28.0%, 32.8%, 20.8%, respectively. All sequences have been deposited in GenBank (Suppl. material
Phylogenetic inferences based on COI gene using neighbor-joining (NJ) and Bayesian inference (BI) reveal that montane Mandarella flea beetles are monophyletic, with three lineages, each including several morphospecies (Fig.
Among Mandarella flea beetles, the interspecific genetic distances range from 16.2–22.6%, while the intraspecific divergence is 0.0–14.4%.
Stenoluperus tibialis:
Mandarella tibialis:
Luperus uenoi Kimoto, 1969: 39 (Taiwan).
Stenoluperus nipponensis:
Stenoluperus potanini:
Stenoluperus taiwanus Kimoto, 1991a: 14. New synonym
Mandarella taiwana:
Stenoluperus minor Kimoto, 1991b: 117 (nec Kimoto, 1977).
Stenoluperus kimotoi Döberl, 2001: 383. (replacement name for Stenoluperus minor Kimoto, 1991). New synonym
Luperus uenoi. Holotype ♂ (
Stenoluperus minor. Paratype: 1♂ (
Stenoluperus taiwanus. Paratypes: 1♀ (
Stenoluperus nipponensis. 1♂ (
Stenoluperus potanini. 1♀ (
Stenoluperus tibialis. 1♂ (
Male. Body size, relative lengths of antennomeres, and color patterns extremely variable, separated into six forms:
Form A (formerly identified as M. uenoi): Length 3.1–3.5 mm. General color metallic blue except antenna, leg, and abdomen black (Figs
Diagnostic characters of Mandarella uenoi. 20 Male antenna, form A, typical 21 Male antenna, form A, elongate 22 Male antenna, form B 23 Male antenna, form C 24 Male antenna, form D, typical 25 Male antenna, form D, variation 26 Male antenna, form E 27 Male antenna, form F 28 Penis, dorsal view 29 Penis lateral view 30 Gonocoxae 31 Ventrite VIII 32 Spermatheca.
Form B (formerly identified as M. potanini): Similar to form A, but body larger, length 3.7–4.1 mm; elytron without longitudinal ridges (Figs
From C (formerly identified as M. nipponensis): Length 3.8–4.1 mm. Similar to form B (Figs
Form D (formerly identified as M. tibialis): General color metallic blue but antennae and legs yellowish brown, coxa and femora metallic blue except apex (Figs
Form E (formerly identified as M. kimotoi): General color metallic blue but antennae and legs yellowish brown (Figs
Form F (formerly identified as M. taiwana): Similar to form E, but larger, length 3.4–3.7 mm. Antennae longer, about 1.1X longer than body; antennomere III much longer than III, III to X extremely slender; ratio of length of antennomeres II to XI 0.4 : 1.0 : 1.5 : 1.6 : 1.5 : 1.6 : 1.5 : 1.5 : 1.4 : 1.6; ratio of length to width from antennomeres II to XI 1.5 : 3.9 : 6.1 : 6.4 : 6.5 : 6.7 : 6.3 : 7.1 : 6.7 : 6.7 (Fig.
Pronotum 1.4–1.6 times as broad long, quadrate, disc with scattered fine punctures, sometimes with feeble lateral depressions. Elytra 1.7–1.8 times as long as broad, parallel-sided, disc with dense, irregular, coarse punctures. First tarsomeres of front and middle legs extremely variable, extremely swollen, either elongate swollen or apically swollen. Posterior margin of last abdominal ventrite rounded, with two small incisions. Penis (Figs
Females. Length 3.5–3.7 mm, width 1.5–1.7 mm. Similar to male; head weakly constricted behind eyes. First tarsomeres of front and middle legs normal and not swollen. Gonocoxae (Fig.
Mandarella nipponensis, M. tibialis, and M. potanini were misidentified as M. uenoi. Mandarella nipponensis possesses a wide and asymmetric penis (Figs
Adults are abundant at high altitudes during spring and summer. The first author collected more two hundred specimens in three hours in Yuanfeng, Nantou county on June 12, 2015. They were resting on leaves of various plants and produced very small feeding scars.
Endemic to Taiwan.
A total of 1008 specimens was examined (Supplementary file 2: Mandarella uenoi, specimens examined).
Stenoluperus pallipes: Kimoto, 1969: 39 (Taiwan); Kimoto, 1989: 253 (additional records in Taiwan). Misidentification
Holotype ♂ (
(n = 354). Holotype ♂ (
Male. Length 3.3–4.1 mm, width 1.3–1.7 mm. General color (Figs
Female. Length 4.3–4.7 mm, width 1.8–1.9 mm. Similar to male (Figs
This new species is similar to M. pallipes but the latter lacks lateral depressions and ridges on the elytra.
Host plant. Adults are closely associated with Stachyurus himalaicus Hook. f. & Thomson ex Benth. (Stachyuraceae), which is sympatric with Dercetina itoi Kimoto, 1969 and D. shirozui Kimoto, 1969.
This new species is named after Mr. Mei-Hua Tsou, a member of the TCRT and the first to collect this new species.
Endemic to Taiwan.
Stenoluperus flaviventris Chen, 1942: 67 (China: Jiangxi);
Stenoluperus esakii Kimoto, 1969: 40. New synonym
Stenoluperus matsumurai Takizawa, 1978: 128. New synonym
Mandarella matsumurai:
Stenoluperus itoi Kimoto, 1991b: 116 (nec Stenoluperus itoi Chûjô, 1966).
Mandarella taiwanensis Medvedev, 2012: 427 (replacement name for Stenoluperus itoi Kimoto, 1991). New synonym
Stenoluperus flaviventris. The holotype was reportedly deposited at the Institute of Zoology, Chinese Academy of Sciences, China but could not be found (Yong-Ying Ruan, pers. comm. 8 October 2015).
Sternoluperus esakii. Holotype ♀ (
Stenoluperus itoi. Paratype: 1♂ (
Stenoluperus matsumurai. Holotype ♂ (
1♀ (
Color patterns and relative lengths of antennae separated into four forms:
Form G (formerly identified as M. flaviventris): General color (Figs
Form H (formerly identified as M. esakii): Similar to form G, but antennae and legs dark brown (Figs
Form I (formerly identified as M. matsumurai): General color metallic blue but antennae, legs, and abdomen yellowish brown; seven apical antennomeres darkened antennomeres (Figs
Form J (formerly identified as M. taiwanensis): Color pattern similar to form I, but abdomen blackish brown. In male, antennae 1.3X longer than body (Fig.
Male. Length 3.6–4.6 mm, width 1.6–2.1 mm. Head strongly constricted behind eyes. Pronotum 1.4 times as broad long, quadrate, disc with dense and coarse punctures as on elytra, lacking lateral depressions. Elytra 1.7 times as long as broad, parallel-sided, disc with dense, irregular, coarse punctures. First tarsomeres of front and middle legs swollen. Posterior margin of last abdominal ventrite truncate, with two small incisions. Penis (Figs
Female. Length 3.9–5.4 mm, width 1.9–2.6 mm. Similar to male; but head weakly constricted behind eyes. First tarsomeres of front and middle legs normal and not swollen. Gonocoxae (Fig.
Although Mandarella flaviventris is highly variable in color patterns, it is characterized by the small third antennomere (3rd antennomeres ≤1.3 times as long as 2nd antennomere). Some black individuals of M. uenoi also have small 3rd antennomeres, similar to M. flaviventris but their abdomens are black (yellow abdomens in M. flaviventris).
Like Mandarella uenoi, adults rested on leaves of various plants and left small feeding scars.
China (Fujian, Jiangxi), Taiwan.
Totally 717 specimens were studied (Suppl. material
1 | Lateral depression and ridge present on each elytron; antennae, legs, and abdomen yellow, and antennomere III much longer than antennomere II (1.6 times) | M. tsoui sp. n. |
– | Lateral depression and ridge absent from each elytron; individuals with yellow antennae, legs, and abdomen, antennomere III slightly longer than antennomere II (1.3 times) (form I of S. flavivientris) | 2 |
2 | Individuals with black or blackish legs, abdomen black, in males antennomere III from slightly longer to much longer than antennomeres II (≥1.3 times); individuals with yellow legs, antennomere III much longer than antennomere II (≥2.0 times); tectum of penis apically tapering, apex of endophallic sclerite bifurcate and acute | M. uenoi (Kimoto, 1969) |
– | Individuals with black or blackish legs, abdomen yellow, in males antennomere III shorter than antennomere II (0.8 times); individuals with yellow, antennomere III slightly longer than antennomere II (1.3 times); tectum of penis membranous and invisible, apex of endophallic sclerite with dense marginal setae | M. flaviventris (Chen, 1942) |
A total of 11 species within Mandarella has been reported from Taiwan. Molecular analyses based on the COI sequences and morphological studies, including male aedeagi, revealed that only three species exist in Taiwan (Fig.
Morphological characters (i.e., body sizes, color patterns, relative lengths between antennomeres) used previously to identify different morphospecies likely are a reflection of their altitudinal distributions induced by local adaptations. For example, in the Hohuanshan mountains, the black forms A/B was mainly collected at an elevation of 3422 m (Hohuanshan Mt. Peak), while at 2756 m (Yuanfeng), the form D with yellow legs and darkened basal femora, is dominant with only seven out of 239 specimens representing the form A. Phylogenetic inferences of these Mandarella flea beetles also revealed local adaptation. The variable morphological forms recognized in the M. uenoi and M. flaviventris lineages may represent a more complicated scenario and related to evolutionary processes. Additional specimens from different montane areas are required to elucidate their phylogeographic histories and address the local adaptations of body size, body color, and the length of antennomeres.
We thank the curators mentioned above for allowing us to study the type and voucher specimens. We thank the Taiwan Chrysomelid Research Team for assistance in collecting material, including Hou-Jay Chen, Jung-Chang Chen, Yi-Ting Chung, Bo-Xin Guo, Hsueh Lee, Wen-Chuan Liao, Mei-Hua Tsou, and Su-Fang Yu. We especially thank Ta-Hsiang Lee for photos of specimens, Hsueh Lee and Mei-Hua for photos in the field. We are grateful to Prof. Christopher Carlton (Louisiana State Arthropod Museum, USA) for reviewing the manuscript. This work was supported by grants from National Chung Hsing University and Agricultural Research Institute, Council of Agriculture, Executive of Yuan, ROC (
Collection information, taxon ID, and accession numbers of COI gene for each flea beetle
Data type: DNA alignment
Explanation note: DNA Submission in GenBank.
Mandarella uenoi, other material examined
Data type: Occurence
Explanation note: 1008 specimens were examined. The localities, dates, and depositories were recorded.
Mandarella flaviventrites, other material examined
Data type: Occurence
Explanation note: 707 specimens were examined. The localities, dates, and depositories were recorded.