﻿Three new species and five new records within the genus Lilioceris (Coleoptera, Chrysomelidae, Criocerinae) from China

﻿Abstract The Chinese species of Lilioceris are revised, and three new species are described from Tibet, China: Lilioceriszhentangensis Xu & Liang, sp. nov., Liliocerismedogensis Xu & Liang, sp. nov. and Lilioceriszayuensis Xu & Liang, sp. nov. Five species of Lilioceris are reported for China as new records: L.dromedarius (Baly, 1861), L.pulchella (Baly, 1859), L.semicostata (Jacoby, 1908), L.unicolor (Hope, 1831) and L.nepalensis Takizawa, 1989. Liliocerisseminigra (Jacoby, 1889) is proposed as a junior synonym of L.unicolor Hope, 1831. Redescriptions, habitus photographs, geographic distributions, host plants (if available) and habitats are provided for these species.


Introduction
Lilioceris Reitter, 1913 is the second largest genus of Criocerinae, containing approximately 160 species in the world and 60 species in China.Most of the species of Lilioceris are distributed in the Oriental region (Clavareau 1913;Monrós 1960) and the Chinese species are mainly distributed in the southern part of China (Bezděk and Schmitt 2017).
Tibet is the second largest autonomous region in China, with various climatic zones from the tropics to frigid mountains.However, Lilioceris in Tibet has been poorly investigated in the past: only four species have been recorded (Yang 2004).Lilioceris subpolita (Motschulsky) was obviously misidentified from Tibet and is excluded from its fauna (Bezděk and Schmitt 2017;Xu et al. 2021).In recent years seven expeditions were made in Tibet and Yunnan.These expeditions resulted in more than two thousand specimens of Criocerinae.Among Lilioceris collected in Dinggyê, Mêdog and Zayü were species new to science.We also identified five species, collected in Tibet and Yunnan that were originally recorded in India, Nepal or Cambodia.
The purpose of this paper is three-fold: to describe three new species from Tibet, to report five new distribution records from Tibet, Yunnan and Hainan, and to synonymize a species of Lilioceris.Diagnosis.Antennae nearly half as long as body length, antennomeres V-X quadrate.Pronotum with distinct anterior and posterior transverse impressions, pronotal disc with two rows of fine and irregular punctures in middle.
Head (Fig. 1).Vertex with shallow groove in middle, punctate and pubescent, almost smooth; frontoclypeal area triangular, lateral side of disc with sparse punctures and pubescence; labrum transverse, with sparse long pubescence; antennae nearly half as long as body, antennomeres I-IV nearly globular, antennomeres V-X 1.2 times as long as wide, III-XI densely pubescent and punctate.
Pronotum (Figs 1,19A).Anterior angle protruding, posterior angle not protruding; sides distinctly constricted in middle; anterior and posterior transverse impression distinct; middle of disc with two rows of fine and irregular punctures; basal transverse groove indistinct.Scutellum triangular and densely pubescent.
Abdominal sternites (Fig. 19C).Lateral transverse impressions distinct on sternites I-IV, area of transverse impressions and middle of sternites I-IV smooth, other areas with dense pubescence and punctures.
Legs (Fig. 2).Femora with dense pubescence on dorsal surface, with sparse pubescence on ventral surface, middle area widened.
Female reproductive organs (Fig. 28A-C).Tergites VIII and IX, and sternites VIII and IX of female sclerotized, posterior areas of tergite VIII and sternite VIII with pubescence and apodemes, spiculum gastrale Y-shaped and long; ovipositor with dense pubescence, distal part of ovipositor cylindrical, short, with protuberance; spermatheca simple and curved.
Etymology.The specific name zhentangensis refers to its type locality Zhêntang, Dinggyê County, Tibet.

Host plant and habitat (Figs 42-45).
The host plant is Smilax menispermoidea A. DC., (Smilacaceae) according to our observations in Zhêntang town.Zhêntang is located in a deep valley at the southern part of the Himalayas.Warm and humid air currents from the Indian Ocean enter the valley frequently.Abundant rainfall and rugged topography in the valley make the biodiversity of Zhêntang very rich.This species occurs at the altitude of 2200 to 2800 m.The habitat is open, composed of tall trees, woody vines and weeds.Remarks.This species looks similar to L. cyanicollis (Pic, 1916) (our concept is based on a specimen determined by J. L. Gressitt, NHML), but is differentiated by its pronotum with distinct anterior and posterior transverse impressions; head, antennae, and legs without blue metallic luster.In L. cyanicollis, the pronotum only with a weak posterior transverse impression; head, antennae, and legs with a blue metallic luster.It is also similar to L. latissima (Pic, 1932) (based on a syntype studied, MNHN), but differs by the metasternum with a short strip of pubescence.In L. latissima, the metasternum is glabrous.Diagnosis.Elytra and abdomen brownish red, rest of body black.Antennae nearly half as long as body length, antennomeres V-X quadrate.Pronotal disc with two rows of fine punctures in middle.Elytral punctures sparse and absent on apical 1/3.
Head (Fig. 3).Vertex with shallow groove in middle, punctate and pubescent sparsely; frontoclypeal area triangular, lateral side of disc with sparse punctures and pubescence; labrum transverse, with sparse long pubescence; antennae nearly half as long as body, antennomeres I-IV nearly globular, antennomere II shortest, antennomeres V-X 1.2 times as long as wide, V-XI densely pubescent and punctate.
Pronotum (Figs 3,20B).Anterior angle protruding, posterior angle not protruding; sides distinctly constricted in middle; middle of disc with two rows of fine punctures; anterior and posterior transverse impression shallow, basal transverse groove indistinct.Scutellum triangular and densely pubescent.
Abdominal sternites (Fig. 20C).Lateral transverse impressions distinct on sternites I-IV.Lateral side of sternite I-IV and pygidium densely pubescent, other areas with sparse pubescence and punctures.
Female reproductive organs (Fig. 29A-C).Tergites VIII and IX, and sternites VIII and IX sclerotized, posterior areas of tergite VIII and sternite VIII with pubescence and apodemes, spiculum gastrale Y-shaped and short; ovipositor with dense pubescence, distal part of ovipositor cylindrical, short and with protuberance; spermatheca simple and curved.
Etymology.The specific name medogensis refers to its type locality Mêdog, Tibet, China.

Host plant and habitat (Figs 38-41).
Beetles were found to feed on Smilax ferox Wall.Ex Kunth (Smilacaceae) in Mêdog in the northernmost edge of the tropics (China, Tibet, Mêdog), with altitudes of ~ 1000 to 2000 m.The type locality Renqingbung temple is located on a mountain in Mêdog County, with high temperatures, high humidity and plentiful precipitation.Vegetation type is subtropical evergreen broadleaved forest.
Remarks.Lilioceris medogensis sp.nov.and L. zhentangensis sp.nov.are similar in their pronotia having anterior and posterior transverse depressions, which are easily distinguished from those of the other members of Lilioceris.However, L. medogensis sp.nov. is different from L. zhentangensis sp.nov.by the brownish red pronotum (Fig. 20A) and metasternum (Fig. 20B), only the lateral side of the metasternum is black; the lateroposterior corner of the metasternum is glabrous; and the posterior part of the dorsal sclerite of the male genitalia in dorsal view is widened (Fig. 23C).In L. zhentangensis, the pronotum and metasternum are black (Fig. 19A); the lateroposterior corner of the metasternum has a strip of pubescence (Fig. 19B); and the posterior part of the dorsal sclerite in dorsal view is parallel (Fig. 22C).In addition, their host plants and habitats are also different: L. medogensis lives on Smilax ferox in warmer and lower altitudes, while L. zhentangensis inhabits Smilax menispermoidea in colder, higher altitudes.Diagnosis.Antennae nearly half as long as body length, antennomeres V-IX quadrate.Pronotum without distinct anterior and posterior transverse impression, pronotal disc with four or five fine punctures in middle.Elytral punctures sparse and absent on apical 1/3.Metasternal disc almost glabrous, posterior margin with sparse pubescence.
Head (Fig. 5).Vertex flat and without groove in middle, lateral side with sparse punctures and pubescence; clypeofrontal area triangular, lateral side of disc with sparse punctures and pubescence; labrum transverse, with sparse long pubescence; antennae nearly half as long as body, antennomeres I-IV nearly globular, antennomeres V-X 1.5 times as long as wide, III-XI densely pubescent and punctate.
Pronotum (Figs 6,21A).Anterior angle protruding, posterior angle not protruding; sides distinctly constricted in middle; without anterior and posterior transverse impression of disc; disc almost smooth and with four or five fine punctures in middle; basal transverse groove indistinct.Scutellum triangular and densely pubescent.
Abdominal sternites.Lateral transverse impressions distinct on sternites I-IV, area of transverse impressions smooth, middle of sternite I with dense pubescence, other areas of sternites I-IV with sparse pubescence and punctures (Fig. 21C).
Etymology.The specific name zayuensis refers to its type locality Zayü County, Tibet, China.
Host plant and habitat (Figs 46, 47).Beetles were collected feeding on Smilax longebracteolata J. D. Hooker (Smilacaceae) in Zayü.This species is con-fined to subtropical areas, at an elevation of ~ 1900 m.Xiongjiu village, Zayü has a mild climate and abundant rainfall.The habitat is secondary forest along a roadside, composed of tall trees, woody vines and many weeds.
Remarks.This new species is a member of the neptis species group, and can be keyed out in couplet 4 with L. cantonensis (Heinze, 1943) and L. neptis (Weise, 1922) in the key by Xu et al. (2021: 301).It is different from L. cantonensis by its middle femora being completely black; elytral strial punctures dense and large at the base, diminished posteriorly, and absent on the apical 1/3 or 1/4; apical portion of the dorsal sclerite of the male genitalia is rounded.In L. cantonensis, the middle femora are bicolored, black with the middle brownish red on their ventral surfaces, and the elytral strial punctures sparse and large in basal impression, but absent on the apical 1/2 or 1/3; apical portion of the dorsal sclerite of the male genitalia truncated.It is also different from L. neptis by the black middle femora; metasternum almost glabrous, with only the posterior margin with sparse pubescence.In L. neptis, the middle femora are brownish red, and the metasternum has a long strip of setae extending from the anterior to the posterior margin.
This species is only found at the type locality.It seems that the local population was very low.We explored this place three times, but only collected eight specimens.We also tried several other places in Zayü County, but no specimens of this species were found.Diagnosis.Antennae nearly half as long as body length, antennomeres VIII-X widened, twice as wide as long.Pronotal disc with two rows of fine punctures in middle.Elytra raised near suture at base, elytral punctures sparse and coarse, but absent at apex.

New record for China
Redescription.BL = 8.8 mm, BW = 4.5 mm.Antennomeres VIII-IX, head, femora and tibiae brownish, claws black, antennomeres I-VII, pronotum, elytra and abdomen brownish yellow, each elytron with big brownish marking in middle of lateral area, abdominal sternites except pygidium with three black markings in lateral side and middle.
Head (Figs 7,8).Vertex with groove in middle, punctate and pubescent densely; frontoclypeal area triangular, lateral side of disc with sparse punctures and pubescence; labrum transverse, with sparse long pubescence; antennae nearly 1/2 as long as body, antennomeres I-IV nearly globular, antennomere II shortest, antennomere V longest, antennomeres VI and VII length as long as width, antennomeres VIII-X widest, wide 2 times as long as length, antennomeres V-XI densely pubescent and punctate.
Pronotum (Figs 7,8).Anterior angle protruding, posterior angle not protruding; sides slight constricted in middle; middle of disc with two rows of fine punctures; anterior and posterior transverse impression indistinct, basal transverse groove very weak.Scutellum triangular and densely pubescent.
Elytra (Figs 7,8).Suture at base with raised hump; humeri protruding, humeral groove and basal impression distinct; striae with very sparse and coarse punctures, puncture absent at elytral brownish black marking area, intervals smooth; epipleura slightly raised, with row of fine punctures.
Mesosternum pubescent; apical portion of mesosternal process narrow and flat, obliquely pointed, not horizontally connected with metasternum.Metasternal disc and metepisternum densely pubescent.
Abdominal sternites.Lateral transverse impressions distinct on sternites I-IV, area of transverse impressions smooth, other areas with dense pubescence and punctures.
Legs.Femora with dense pubescence on dorsal surface, with sparse pubescence on ventral surface, middle area widened.
Male genitalia.Unknown.Female reproductive organs (Fig. 31A-C).Tergites VIII and IX, sternites VIII and IX sclerotized, posterior areas of tergite VIII and sternite VIII with pubescence and apodemes, spiculum gastrale Y-shaped and distinctly widen in distal part; ovipositor with dense pubescence, distal part of ovipositor cylindrical, long and with protuberance; spermatheca greatly convoluted.
Host plant and habitat.A host plant is unknown.A single specimen in IZ-CAS was collected by Meiying Lin when beating vegetation.The collecting site, Hudiegu, is located in a tropical area in Wuzhi Shan of Hainan Province, with high temperatures, high humidity, and plentiful precipitation.Vegetation type is tropical evergreen broadleaved forest.
Remarks.This species is very similar to Lilioceris gibba (Baly, 1861) (based on a syntype studied, NHML) but differs from the latter by antennomeres V-X being twice as wide as long (in L. gibba, antennomeres V-X as wide as long).In addition, the pronotum and elytra are yellow in L. dromedarius (dark brown in L. gibba).Diagnosis.Antennae ~ 1/3 as long as body, antennomeres V-X cylindrical.Pronotum with distinct posterior transverse impression, pronotal disc with two rows of fine punctures in middle, scutellum pubescent.Elytral punctures fine, diminishing posteriorly, but not absent.Lateral side of metasternite with long narrow strip of pubescence.

New records
Redescription.BL = 11.0 mm, BW = 5.0 mm.Antennae, head, pronotum, scutellum, prosternum, mesosternum, legs, and half of first abdominal sternite black, with blue metallic luster, elytra and remainder of abdominal sternite brownish red.Head (Figs 9,10).Vertex with deep groove in middle, sparsely punctate and pubescent in lateral area; frontoclypeal area triangular, lateral side of disc with sparse punctures and pubescence; labrum transverse, middle of anterior margin concave, disc with sparse punctures and pubescence; antennae nearly 3/5 length of body, antennomeres I-IV nearly globular, antennomere II shortest, antennomeres V-XI cylindrical, 3 times as long as wide.
Pronotum (Figs 9,10).Anterior angle protruding, posterior angle not protruding; sides constricted in middle; anterior and posterior transverse impression distinct; disc with irregular fine punctures in middle; scutellum triangular and densely pubescent.
Mesosternum pubescent; apical portion of mesosternal process narrow and flat, obliquely pointed, not horizontally connected with metasternum.Lateral side of metasternite with a long narrow strip of pubescence extending from the lateroposterior corner to anterior margin.Metepisternum densely pubescent.
Abdominal sternites.Lateral transverse impressions indistinct on sternites I-IV.Lateral side of sternite I-IV with densely pubescent, other areas with sparse pubescence and punctures.
Legs.Femora with dense pubescence in dorsal surface, with sparse pubescence on ventral surface, middle area widened.Claws distinctly asymmetrical, outer one is longer than inner one.
Male genitalia.Unknown.Female reproductive organs (Fig. 32A-C).Tergites VIII and IX, sternites VIII and IX sclerotized, posterior areas of tergite VIII and sternite VIII with pubescence and apodemes, spiculum gastrale X-shaped and short; ovipositor with dense pubescence, distal part of ovipositor cylindrical, short, and with protuberance; spermatheca simple and folded.
Distribution (Fig. 37).China (Tibet); India.Host plant and habitat.A host plant is unknown.This species was collected in an environment between Hanmi and Lage according to the collector.Hanmi and Lage are two small courier stations on an old trail from Doxong La pass to Baibung town in Mêdog County, at altitudes of 2000-3000 m.They are located in a subtropical area, with a primary forest composed of large trees, woody vines and many shrubs.
Head (Figs 11,12).Vertex raised, with deep groove in middle, sparsely punctate and pubescent in lateral area; frontoclypeal area triangular, lateral side of disc with sparse punctures and pubescence; labrum transverse, middle of anterior margin concave, disc with sparse punctures and long pubescence; antennae nearly as long as body length, antennomeres I-III nearly globular, antennomere II shortest, antennomeres IV-XI cylindrical, 3 times as long as wide.
Mesosternum pubescent; apical portion of mesosternal process narrow and flat, obliquely pointed, not horizontally connected with metasternum.Metasternite with scattered and sparse pubescence.Metepisternum sparsely pubescent.
Abdominal sternites.Lateral transverse impressions indistinct on sternites I-IV.Lateral sides of sternites I-IV with denser pubescence, other areas with sparse or scattered pubescence.
Legs.Femora with dense pubescence on dorsal surface, with sparse pubescence on ventral surface, middle area widened.Claws distinctly asymmetrical, outer one longer than inner one.
Female reproductive organs (Fig. 33A-C).Tergites VIII and IX, sternites VIII and IX sclerotized, posterior areas of tergite VIII and sternite VIII with pubescence and apodemes, spiculum gastrale X-shaped and long; ovipositor with dense pubescence, distal part of ovipositor cylindrical, short and with protuberance; spermatheca simple and curved.
Distribution (Fig. 37).China (Tibet); India.Host plant and habitat.Host plant is unknown.Specimens of this species in IZCAS were collected in Hanmi by Ye Liu and Xiaodong Yang when sweeping the vegetation canopy.Hami is located in the subtropical area of Mêdog, with primary forest composed of large trees, woody vines and many shrubs.
Remarks.This species is very similar to Lilioceris flavipennis (Baly, 1859) (based on a syntype studied, NHML), but differs from the latter by the pronotum being slightly constricted in middle, anterior and posterior angles not protruding; elytral striae regular.In L. flavipennis, pronotum is strongly constricted in middle, anterior and posterior angles are strongly protruding; elytral striae are irregular.27601°N, 97.85190°E, 902 m, 2020.5.26, Hongbin Liang and Yuan Xu coll. (IZCAS).

Lilioceris unicolor
Diagnosis.Antennae ~ 1/3 as long as body, antennomeres V-XI quadrate.Pronotum without anterior and posterior transverse impression, pronotal disc with one or two rows of punctures in middle.Elytral punctures large, not diminishing posteriorly, intervals slightly convex on apical 1/4; epipleura raised, with row of fine punctures.
Redescription.BL = 8.0 mm, BW = 3.8 mm.Body brownish red.Head .Vertex with shallow groove in middle, sparsely punctate and pubescent in lateral area; frontoclypeal area triangular, disc with sparse punctures and pubescence; labrum transverse, disc with sparse long pubescence; antennae nearly 1/3 length of body, antennomeres I-IV nearly globular, antennomere II shortest, antennomeres V-X strongly widened, 2 times as wide as long.
Pronotum .Anterior angle protruding, posterior angle not protruding; sides constricted in middle; anterior and posterior transverse impression absent; middle of disc with one or two rows of fine punctures.Scutellum triangular and pubescent.
Elytra .Humeri protruding, humeral groove distinct, basal impression indistinct; striae with large punctures, punctures not diminishing posteriorly, intervals slightly convex on apical 1/4; epipleura raised, with row of fine punctures.Abdominal sternites.Lateral transverse impressions indistinct on sternites I-V, lateral sides of sternites I-V with dense pubescence, only with sparse pubescence in middle.
Legs.Femora with dense pubescence on dorsal surface, with sparse pubescence on ventral surface, middle area widened.
Female reproductive organs (Fig. 35A-C).Tergites VIII and IX, sternites VIII and IX sclerotized, posterior areas of tergite VIII and sternite VIII with pubescence and apodemes, spiculum gastrale Y-shaped and long; ovipositor with dense pubescence, distal part of ovipositor cylindrical, short and with protuberance; spermatheca convoluted.
Host plant and habitat.We collected two adults of L. unicolor on Dioscorea sp.(Dioscoreaceae) in the villages of Jiemao and Longbazhai of Yunnan Province, but feeding was not observed, so their host plant needs confirmation.Jiemao and Longbazhai are located in a subtropical area, at elevations of 800-900 m.These two places have a mild climate and abundant rainfall.The habitat in Jiemao is a large secondary forest, composed of tall trees, woody vines, shrubs and weeds.The habitat in Longbazhai is a very small secondary forest of several thousand square meters, surrounded by crop fields.
Remarks.We examined the lectotype and a non-type specimen of L. unicolor present at NHML.The specimens in IZCAS from Yunnan are not significantly different from the lectotype.The punctures on pronotum are variable: the lectotype has two rows of punctures in the middle of the pronotum, the non-type specimen in NHML and the specimen in IZCAS have only one row of punctures in the middle of the pronotum.The male genitalia of our specimen from Yunnan (Fig. 26C, D) are identical to those of the lectotype in NHML (Tishechkin et al. 2011: 80, fig. 30).
We also examined a syntype of L. seminigra in NHML, and no significant morphological difference was found from the lectotype of L. unicolor, except for the bi-coloration on the elytra of L. seminigra.When Jacoby (1889: 153) described L. seminigra, he noticed a variant: "Var.Elytra entirely fulvous".In four Chinese specimens in IZCAS, the elytrae are all brownish-red in two specimens of L. unicolor; but the apical 2/3 of each elytron is black, 1/3 of base is brownish-red in one specimen of L. seminigra, and completely brownish-red in another specimen.The female reproductive organs of these two species were dissected and compared, and no significant differences were found.Therefore, we conclude that L. unicolor and L. seminigra are conspecific, and elytral bi-coloration in L. seminigra is simply a variation.In addition, antennomeres V-X in this species are flat, strongly transverse, and the outer distal angle of antennomeres V-VII is protruding.These characteristics are unique among Lilioceris species.Takizawa, 1989 Figs 17, 18, 27A-D, 34A-C, 37, 48-51 Lilioceris nepalensis Takizawa, 1989: 327 (Nepal: Bagmati, holotype        Diagnosis.Antennae ~ 2/3 as long as body, antennomeres V-X cylindrical.Pronotum with distinct posterior transverse impression, pronotal disc smooth.Elytral punctures sparse on basal half but absent on apical half.Metasternite smooth.
Mesosternum pubescent; apical portion of mesosternal process narrow and flat, obliquely pointed, not horizontally connected with metasternum.Metasternum and metepisternum smooth.
Legs.Femora with dense pubescence on dorsal surface, with sparse pubescence on ventral surface.Claws distinctly asymmetrical, outer one longer than inner one.

Figure 36 .
Figure 36.Collecting sites of three new Lilioceris species.

Figure 37 .
Figure 37. Collecting sites of five new records of Lilioceris species in China (distributions outside China are not marked here because of the lack of precise locality data).