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A new species, a new combination, and a new record of Crossotarsus Chapuis, 1865 (Coleoptera, Curculionidae, Platypodinae) from China
expand article infoShengchang Lai, Ling Zhang§, You Li|, Jianguo Wang§
‡ Nanjing Forestry University, Nanjing, China
§ Jiangxi Agricultural University, Nanchang, China
| University of Florida, Gainesville, United States of America
Open Access

Abstract

This study describes a new species, Crossotarsus beaveri Lai & Wang, sp. nov., designates a new combination, C. brevis (Browne, 1975, comb. nov. from Platypus Herbst, 1793), and notes a new record, C. emorsus Beeson, 1937, from China. Genetic data from four genes indicate that the new species and C. brevis form a clade clustered with other Crossotarsus species. Molecular phylogeny and morphological characters support their taxonomic placement.

Keywords

Ambrosia beetle, Fujian, Jiangxi, molecular phylogeny, pinhole borer, taxonomy

Introduction

The genus Crossotarsus Chapuis 1865 was erected for 29 species of pinhole borers. Crossotarsus wallacei (Thomson, 1857) was designated as the type species of the genus (Hopkins 1914). Wood (1993) revised the genera of Platypodidae and placed Crossotarsus in the subfamily Platypodinae, tribe Platypodini. Crossotarsus is distinguished from other Platypodine genera primarily by the following combination of characters (Browne 1961; Wood 1993; Beaver and Sanguansub 2015): 1. Labial palps two-segmented, with basal segments fused in the midline; 2. Sexually dimorphic protibiae, the outer face of the protibia transversely carinate in the male and finely granulate in the female; 3. Pronotum without specialized mycangial pores in either sex; 4. Femoral grooves angulate at the anterior extremity and gently rounded behind; 5. Metacoxa strongly projecting with a deep vertical posterior face. But Wood’s (1993) generalisation that the female pronotum of Crossotarsus species has numerous mycangial pores is incorrect (Beaver 2004).

The catalog of Wood and Bright (1992) includes 118 species of Crossotarsus. As a result of taxonomic changes since that time, 116 species are currently recognised. Most species of Crossotarsus occur in the Oriental region, extending from India across Southeast Asia and Indonesia to Australia and the Pacific islands, and northward to Taiwan and Japan (Wood 1993). Crossotarsus externedentatus (Fairmaire 1849) is also widespread in the Afrotropical forests.

The Platypodinae have been almost entirely neglected in China. Only a few papers include original records of Crossotarsus from the country. Yin and Huang (1987) recorded three species: C. coniferae Stebbing, 1906, C. squamulatus Chapuis, 1865, and C. wallacei from Yunnan. Yin et al. (2002) added two species: C. externedentatus and C. terminatus Chapuis, 1865 from Hainan island, and Zhang et al. (2008) cited 13 species from China. After taxonomic changes (Beaver 2004; 2005; 2016; Bright 2014), the following 13 species are currently known from China: C. coniferae (Yunnan, Sichuan, Xizang); C. emancipatus Murayama, 1934 (Taiwan); C. externedentatus (Hainan, Taiwan); C. flavomaculatus Strohmeyer, 1912 (Taiwan); C. formosanus Strohmeyer, 1912 (Taiwan); C. niponicus Blandford, 1894 (Taiwan); C. piceus Chapuis, 1865 (Taiwan); C. saltatorinus (Schedl, 1954) (Fujian); C. sauteri (Strohmeyer, 1913) (Taiwan); C. simplex Murayama, 1925 (Taiwan); C. squamulatus (Yunnan); C. terminatus (Hainan, Yunnan, Xizang); C. wallacei (Hainan, Taiwan).

In this study, we describe a new species of Crossotarsus from China, give a new distribution record, transfer a previously described species to the genus, and provide molecular data of Chinese species for molecular phylogenetic analyses.

Materials and methods

Abbreviations used for collections

BMNH The Natural History Museum, London, United Kingdom.

JXAU College of Agricultural Sciences, Jiangxi Agricultural University, Nanchang, China.

KIZCAS Kunming Institute of Zoology, Chinese Academy of Sciences, Kunming, China.

NIAES National Institute of Agro-Environmental Sciences (ITLJ), Tsukuba, Ibaraki, Japan.

NMNS National Museum of Natural Science, Taichung, Taiwan.

NZMC National Zoological Museum of China, Institute of Zoology, Chinese Academy of Science, Beijing, China.

RAB Private collection of Roger A. Beaver, Chiang Mai, Thailand.

RIFID Research Institute of Forest Insect Diversity, Namyangju, South Korea.

SYU Museum of Biology, Sun Yat-sen University, Guangzhou, China.

USNM National Museum of Natural History, Washington D.C., USA.

ZIN Zoological Institute. Russian Academy of Sciences, St. Petersburg, Russia.

Adults of the new species were collected by log dissection. The samples were immediately preserved in tubes containing 99.9% ethyl alcohol, which were stored at −20 °C for DNA extraction and examination. Specimens were examined using an Olympus SZX160 stereoscopic zoom microscope. Photographs were taken with a KEYENCE VHX-6000 Digital Microscope System. All photos were further adjusted and assembled with Adobe Photoshop CS6. Body length was measured between the anterior margin of the pronotum and the elytral apex (head not included).

Genomic DNA was extracted from the adult head. The total genomic DNA was extracted from each individual using the Ezup Column Animal Genomic DNA Purification Kit (Sangon Biotech Co. Ltd). Amplification of four gene fragments (COI, EF-1α, CAD, 28S) was made by PCR, using primers (Table 1) and cycling conditions previously described (Jordal et al. 2011). The PCR products were sent to Sangon Biotech Co. Ltd (Shanghai, China) for sequencing, and the sequences were analyzed using the software DNAstar. Additional information on Crossotarsus material was collected by the authors in China or downloaded from NCBI (The National Center for Biotechnology Information) (Table 2). Concatenated DNA sequence data from Jordal (2013) were analysed in MrBayes v. 3.2.6 (Ronquist et al. 2012). Partitions and models were estimated by PartitionFinder 2 (Lanfear et al. 2017) and ModelFinder (Kalyaanamoorthy et al. 2017) respectively in PhyloSuite (Zhang et al. 2020), GTR+G+I were selected for each partition. 10 million generations were run, with 25% of the generations as burn-in. PSRF close to 1.0 and standard deviation of split frequencies below 0.01 were accepted.

Table 1.

Gene fragments targeted for PCR and the primers used. Sequencing primers were identical to those used in PCR.

Gene Primer name Annealing Primer sequence Reference
COI S1718 46 5'-GGAGGATTTGGAAATTGATTAGTTCC-3' Jordal et al. 2011
A2411 5'-GCTAATCATCTAAAAACTTTAATTCCWGTWG-3'
28S S3690 55 5'-GAGAGTTMAASAGTACGTGAAAC-3' Jordal et al. 2011
A4394 5'-TCGGAAGGAACCAGCTACTA-3'
EF-1α S149 52 5'-ATCGAGAAGTTCGAGAAGGAGGCYCARGAAATGGG-3' Jordal et al. 2011
A1043 5'-GTATATCCATTGGAAATTTGACCNGGRTGRTT-3'
CAD CAD for4 50 5'-TGGAARGARGTBGARTACGARGTGGTYCG-3' Jordal et al. 2011
CAD rev1mod 5'-GCCATYRCYTCBCCYACRCTYTTCAT-3'
Table 2.

Material used for phylogenetic analyses, including their GenBank accession numbers.

No. Taxon Country CAD COI EF-1α 28S Reference
1 C. beaveri sp. nov. China: Jiangxi LC616080 LC613149 LC613157 This study
2 C. brevis (Browne, 1975) China: Yunnan LC616086 LC613154 LC616520 LC613163 This study
3 C. chalcographus Schedl, 1972 Papua New Guinea KR261163 KR261313 Jordal 2015
4 C. emorsus Beeson, 1937 China: Yunnan LC616087 LC613155 LC613164 This study
5 C. externedentatus (Fairmaire, 1849) China: Yunnan LC616083 LC613152 LC616518 LC613160 This study
6 C. externedentatus (Fairmaire, 1849) Tanzania KR261162 KR261312 KR261216 Jordal 2015
7 C. externedentatus (Fairmaire, 1849) Madagascar KR261166 KR261316 KR261275 KR261218 Jordal 2015
8 C. fractus Sampson, 1912 Papua New Guinea KR261165 KR261315 KR261274 Jordal 2015
9 C. minusculus Chapuis, 1865 Papua New Guinea HQ883809 HQ883669 HQ883739 HQ883579 Jordal 2015
10 C. niponicus Blandford, 1894 China: Sichuan LC613156 LC613165 This study
11 C. nitescens Schedl, 1979 Australia KR261161 KR261311 KR261272 Jordal 2015
12 C. sauteri (Strohmeyer, 1913) China: Jiangxi LC616081 LC613150 LC616516 LC613158 This study
13 C. squamulatus Chapuis, 1865 China: Yunnan LC616084 LC613153 LC613161 This study
14 C. terminatus Chapuis, 1865 China: Jiangxi LC616082 LC613151 LC616517 LC613159 This study
15 C. wallacei (Thomson, 1857) China: Yunnan LC616085 LC616519 LC613162 This study
16 P. contaminatus (Blandford, 1894) China: Jiangxi LC387560 LC383433 LC387562 LC386151 Lai et al. 2019

Results

Crossotarsus beaveri Lai & Wang, sp. nov.

Figures 1, 2

Type material

Holotype. Male, China: Jiangxi Province, Ganzhou City, Longnan County, Jiulianshan national nature reserve of Jiangxi, Hualu Village, 24°37'19"N, 114°29'57"E, 2.VII.2020, log dissection, host Paulownia fortunei, Shengchang Lai leg. (deposited in NZMC IOZ(E)225775).

Allotype. Female, same data as holotype (deposited in NZMC IOZ(E)225776).

Paratypes. 6 males, 6 females, same data as holotype, but host Phoebe zhennan and Liquidambar formosana (5 males, 5 females JXAU; 1 male, 1 female NZMC); 11 male, 6 females, as holotype except: Xunwu County, Xiangshan Town, Congkeng Village, 24°54'20"N, 115°52'44"E, ca 650m, 15.IX.2017, log dissection, host Castanopsis fargesii and Vernicia montana, Shengchang Lai leg. (10 males, 5 females JXAU; 1 male, 1 female RAB); 6 males, 6 females, as holotype except: Xunwu County, Liuche Town, Luanluozhang, 24°40'41"N, 115°44'9"E, ca 640 m, 22.VIII.2017, log dissection, host Castanopsis carlesii, Shengchang Lai leg. (5 males, 5 females JXAU; 1 male, 1 female RAB); 38 males, 38 females, China: Fujian Province, Zhangzhou City, Yunxiao County, Xiahe Town, Qigaoqi Village, 24°1'31"N, 117°10'36"E, 8.VII.2019, log dissection, host C. carlesii, Ling Zhang leg. (2 males, 2 females BMNH; 2 males, 2 females KIZCAS [KIZ0121459–0121462]; 2 males, 2 females NIAES; 2 males, 2 females NMNS; 2 males, 2 females RAB; 2 males, 2 females RIFID; 2 males, 2 females SYU; 2 males, 2 females USNM; 2 males, 2 females ZIN; 20 males, 20 females JXAU).

Figure 1. 

Male of Crossotarsus beaveri sp. nov. A dorsal view B head C lateral view D declivity. Scale bars: 0.5 mm.

Figure 2. 

Female of Crossotarsus beaveri sp. nov. A dorsal view B head C lateral view D declivity. Scale bars: 0.5 mm.

Description

Male. 3.58–4.01 mm long (mean = 3.78; n = 20); 2.75–2.95 times as long as wide. Head and pronotum dark brown, disc of elytra reddish brown becoming dark brown, declivity of elytra nearly black.

Head. Frons flat, slightly shining, with irregular large punctures; finely, sparsely punctured above the epistoma, bearing bristly, erect, long setae, weakly concave, smooth around short median line, upper part of frons with scattered, coarse punctures, the punctures with moderate, semierect, dorsally directed setae. Antennal scape clavate with scattered, forwardly directed hairs on apical half; club oval, flattened, evenly covered with short setae. Labial palps two-segmented, with basal segments fused along the midline.

Pronotum. About 1.2 times longer than wide, shining, no mycangial pores, the lateral femoral grooves angulate anteriorly, pronotum widest in front of the grooves, with finely, scattered, irregular punctures, a few semierect backwardly pointed hairs close to anterior margin, median line extending about 1/4 from base.

Scutellum. Depressed below level of elytra, with a median longitudinal groove between lateral carinae.

Elytra. About 2.0 times as long as wide, about 1.4 times as long as pronotum. Surface of disc smooth, shining, striae distinctly impressed for almost their entire length, except striae 6 and 7, other striae with circular, distinct, shallow punctures, the bases of striae 1 and 2, striae 3 and 4, respectively, conjoint, more impressed; interstriae slightly raised on disc, interstriae 1, 3 and 5 distinctly raised and conjoint at base, interstriae 8 and 9 fused at apex of disc, forming ventral, rounded angle; cylindrical declivity obliquely truncate, acutely margined all around except at sutural apex, strongly concave, forming a cup-like structure, surface shining, with 4 rows of longitudinal granules bearing erect, long, golden setae, a row of sparse, medially directed, erect golden setae at the inner margin of declivity, elytral apex broadly emarginate, the main emargination approximately U-shaped, about as wide as deep, extending about 1/3 of the height of the declivity, at its inner end a much smaller, V-shaped second emargination (Fig. 1A, D).

Figure 3. 

Male of Crossotarsus brevis (Browne) A dorsal view B head C lateral view D declivity. Scale bars: 0.5 mm.

Protibia. Five transverse carinae at tibial apex, transverse rugae at base.

Abdomen. Abdominal ventrites 1–4 moderately finely punctured, with irregular rows of erect, short hairs at both sides posteriorly, ventrite 5 strongly concave at middle, with dense, large, circular punctures.

Female. 3.64–4.42 mm long (mean = 3.96 mm; n = 20); 2.79–2.93 times as long as wide. Head and pronotum brown, disc of elytra reddish brown becoming dark brown to apex.

Head. Similar to male, but frons flatter, very shining, smooth, with shallow, small punctures; finely, sparsely punctured above the epistoma, bearing bristly, erect, long setae; very shallowly concave in median line, upper part of frons with scattered, shallow, small punctures, the punctures with moderate, semierect, dorsally directed setae.

Pronotum. Similar to male.

Elytra. About 1.8 times as long as wide, about 1.5 times as long as pronotum, sides subparallel. Similar to male, but disc of elytra shining, with dense, longitudinal, semierect, backwardly pointed hairs at apex and declivity, striae weakly impressed, interstriae smoother, declivity vertical, a few irregularly granules, sparsely hairy.

Protibia. Three transverse carinae at tibial apex, fine, confused granules at base.

Abdomen. Surface of abdominal ventrites smooth, rounded, sparsely hairy, ventrite 5 without concavity, punctures shallow.

Etymology

The species is named for Roger A. Beaver to honor his contributions to the study of platypodines and scolytines.

Host plants

Castanopsis carlesii (Hemsl.) Hayata, C. fargesii Franch. (Fagaceae), Liquidambar formosana Hance (Altingiaceae), Phoebe zhennan S.K.Lee & F.N.Wei (Lauraceae), Paulownia fortunei (Seem.) Hemsl. (Paulowniaceae), Vernicia montana Lour. (Euphorbiaceae).

Distribution

China (Jiangxi, Fujian).

Diagnosis

The species is placed in Crossotarsus because it possesses a combination of characters similar to that cited in the introduction.

Crossotarsus beaveri is very similar to Crossotarsus brevis (Browne, 1975) (new combination, see below) and Crossotarsus platypoides (Browne, 1955). They can be easily distinguished from other Crossotarsus species by the male elytral apex truncate with a large, circular, concave declivity. The elytral apex of male of C. beaveri and that of C. brevis possess a deep, acutely margined declivity, with a broad, almost circular, apical emargination.

Key to the species of Crossotarsus with a circular, truncate elytral declivity

1 Male elytral apex truncate, with a circular, shallow, concave, bluntly margined declivity; sutural apex of declivity slightly dehiscent without apical emargination. Female smaller and stouter, 2.60–2.70 mm long, 2.70–2.75 times as long as wide C. platypoides Browne
Male elytral apex truncate, with a circular, deep, concave, acutely margined declivity, with a broad, almost circular, apical emargination. Female larger and more elongate, 3.00–3.90 mm long, 2.79–3.44 times as long as wide 2
2 Male striae weakly impressed on disc of elytra (Fig. 1A); declivity gradually, obliquely truncate, its face shining, cylindrical, apex rounded with a double sutural emargination, borders of inner emargination weakly elevated, outer emargination forming pointed angles; surface of declivity with 4 longitudinal rows of granules, bearing erect, long golden setae (Fig. 1D). Female frons flat, more shining, smoother, very shallowly concave in median line; dense, shallow, small punctures bearing semierect hairs on upper part; almost flat above the epistoma below median line (Fig. 2B); striae weakly impressed on disc of elytra (Fig. 2A). 3.64–3.90 mm long C. beaveri sp. nov.
Male striae moderately impressed on disc of elytra (Fig. 3A); declivity abruptly, vertically truncate, its face subnitid, cylindrical, apex rounded with a double sutural emargination, borders of inner emargination distinctly elevated and dilated, outer emargination forming obtuse angles; surface of declivity with sparse, obscure granules, bearing erect, long golden setae (Fig. 3D). Female frons slightly shining, reticulate, very distinctly concave, smooth around median line; dense, deep, large punctures bearing semierect hairs on upper part; weakly, irregularly impressed above the epistoma below median line (Fig. 4B); striae moderately impressed on disc of elytra (Fig. 4A). 2.96–3.44 mm long C. brevis (Browne)

Crossotarsus brevis (Browne, 1975), comb. nov.

Figures 3, 4

Platypus brevis Browne: Beaver and Browne 1975: 306.

Dinoplatypus brevis Browne: Beaver 1998:184.

Material examined

7 males, 5 females (JXAU); 1 male, 1 female (RAB): China: Yunnan Province, Xishuangbanna Dai Autonomous Prefecture, Jinghong City, Damanmi Village, 22°02'50"N, 100°48'27"E, ca 580 m, 20.I.2018, log dissection, host unknown, Shengchang Lai leg.

Figure 4. 

Female of Crossotarsus brevis (Browne) A dorsal view B head C lateral view D declivity. Scale bars: 0.5 mm.

Taxonomy

The specimens in the RAB were identified by comparison to a paratype C. brevis, which is also in the RAB. Browne put this species in Platypus Herbst, noting that the apical emargination of the elytra was rather similar to that of Platypus caliculus Chapuis 1865 (Beaver and Browne 1975). In fact, C. brevis has the typical characters of Crossotarsus: labial palps two-segmented, with basal segments fused in the midline, whereas Platypus has the labial palps three-segmented, with separate basal segments. Beaver (1998) transferred the species from Platypus to Dinoplatypus Wood following Wood’s (1993) attempt to split the genus Platypus. Wood diagnosed Dinoplatypus largely on the basis of the circular, truncate, elytral declivity of the male, with the sutural apex emarginate. However, this is an adaptive character of the declivity which has evolved independently more than once in the Platypodinae, as it has in the Scolytinae (Hulcr et al. 2015). Molecular phylogenetic study also shows that the few morphological characters used by Wood (1993) to erect several groups of Neotropical and Indo-Malayan/Australasian species in Platypodini to new genera are not sufficiently diagnosable for all those groups (Jordal 2015).

Browne (1961) and Beaver and Sanguansub (2015) suggested that the adult generic characters of primary value in Crossotarsus included the structure of the labial and maxillary palps, the form of the pronotum, the sexual dimorphism of the protibia, and various modifications of the abdominal sternites in the male. Based on the two-segmented labial palps, the lateral pronotal emarginations angulate anteriorly, the pronotum without mycangial pores, and the sexual dimorphism of the protibiae, Platypus brevis belongs in the genus Crossotarsus, and is here transferred to that genus.

Distribution

Thailand (Beaver and Liu 2013). New to China (Yunnan).

Host

Castanopsis sp. (Fagaceae) (Beaver and Liu 2013).

New record

Crossotarsus emorsus Beeson, 1937

Figures 5, 6

Crossotarsus emorsus Beeson, 1937: 87.

Material examined

4 males, 1 female (JXAU) China: Yunnan Province, Xishuangbanna Dai Autonomous Prefecture, Jinghong City, Nabanhe River Watershed National Nature Reserve, Guomenshan, ca 1030 m, 22°14'46"N, 100°36'10"E, 27.I.2018, log dissection, host Dalbergia assamica, Shengchang Lai leg.; 1 male, 1 female (RAB); 1 male (JXAU) China: Yunnan Province, Xishuangbanna Dai Autonomous Prefecture, Jinghong City, Damanmi Village, ca 580 m, 22°02'50"N, 100°48'27"E, 20.I.2018, log dissection, host Cassia siamea, Shengchang Lai leg.

Figure 5. 

Male of Crossotarsus emorsus Beeson A dorsal view B head C lateral view D declivity. Scale bars: 0.5 mm.

Diagnosis

C. emorsus is similar to C. terminatus but can be distinguished using the characters given in Table 3.

Table 3.

Diagnostic characters separating Crossotarsus emorsus and Crossotarsus terminatus.

C. emorsus C. terminatus
Body size Male size 4.56–4.80 mm long, 3.20–3.42 times as long as wide; Male size 3.32–3.40 mm long, 2.78–3.00 times as long as wide;
Female size 4.8–5.34 mm long, 3.38–3.43 times as long as wide. Female size 3.44–3.58 mm long, 2.87–2.93 times as long as wide.
Frons Male frons almost flat, with shallower, irregularly placed punctures; circularly concave in median line. Male frons coarser, with deeper, irregularly placed punctures; linearly concave in median line.
Female frons almost flat, without concave around median line. Female frons concave forming a big, circular impression around concave median line.
Elytra Male without lateral emargination at declivity base, semicircular lateral borders with serrated, lateral tubercles. Male with lateral emargination at declivity base, semicircular lateral borders rounded, without distinct serrated, lateral tubercles.

Distribution

Myanmar, Thailand, Laos (Beaver and Liu 2013; Beaver 2016). New to China (Yunnan).

Host

The species is recorded from trees in the families Lecythidaceae, Fabaceae, Sterculiaceae and Verbenaceae (Beeson 1937), and is presumably polyphagous (Beaver 2016). Host plants recorded here are: Senna siamea (Lam.) H.S.Irwin & Barneby and Dalbergia assamica Benth. (Fabaceae).

Figure 6. 

Female of Crossotarsus emorsus Beeson A dorsal view B head C lateral view D declivity. Scale bars: 0.5 mm.

Molecular data

The phylogenetic tree for analyzing the evolutionary relationships of 13 taxa including the ingroups (Crossotarsus species) and the outgroups (P. contaminatus) was constructed based on four genes (Fig. 7). The BI tree shows the new species (C. beaveri) and the new combination (C. brevis) forming a clade, with high node support. These group with Schedl’s (1972a)Crossotarsi coleoptrati’ (C. fractus Sampson, 1912, C. squamulatus and C. terminatus) and cluster with all remaining Crossotarsus species. It confirms that the taxonomic changes and the relationship of C. brevis and C. brevis are correct. It also indicates that C. emorsus, C. fractus, C. squamulatus and C. terminatus should be considered distinct species (as by Beaver and Liu 2013), and not considered synonyms or subspecies (Schedl 1972a).

Figure 7. 

Tree topology resulting from Bayesian analysis of four genes. Posterior probabilities are given on the nodes. New species and new combination in boldface.

Discussion

Crossotarsus beaveri is clearly related to C. brevis. They are the sister lineage to the Crossotarsi coleoptrati group, not the genus Dinoplatypus. This is a good example that declivity in males usually is an adaptive character and not of generic significance. We consider that the morphologically diagnosable characters of the genus Crossotarsus should refer to the summary by Browne (1961) and Beaver and Sanguansub (2015, 2020), as introduction.

The genus Crossotarsus is one of the largest genera of Platypodinae, with more than 100 species. Although there are 13 previously recorded species of Chinese Crossotarsus (Yin and Huang 1987; Yin et al. 2002; Zhang et al. 2008), many additional species have been reported from countries neighboring China (Beaver and Shih 2003; Goto 2009; Beaver and Liu 2013; Beaver 2016) which still have not been found in China. This strongly indicates that many more species remain to be discovered, especially on the Chinese mainland. Crossotarsus is monophyletic in the latest molecular phylogeny (Jordal 2015). There are few molecular data for the genus in GenBank, less than 10% of the whole. More taxonomic samples are needed.

Acknowledgements

We thank Roger Beaver for reviewing earlier manuscript drafts and kindly commenting on the status of the new species. We also appreciate the help provided by Jiaxin Liao and Yufeng Cao in our fieldwork. This research was funded by grant CARS-33-BC2-JX01 from Survey of Bark Beetles on the Rubber Tree in China Project and the National Natural Science Foundation of China (no. 31160380, 31360457, 31760543).

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