﻿A review of the semipunctata species group within the genus Lilioceris Reitter, 1913 (Coleoptera, Chrysomelidae)

﻿Abstract A new species group of Lilioceris Reitter, 1913 is proposed and reviewed, the semipunctata group. It includes eleven species: L.atrilateralis Kimoto & Takizawa, 1973, L.consentanea (Lacordaire, 1845), L.dentifemoralis Long, 1988, L.discrepens (Baly, 1879), L.jianfenglingensis Long, 1988, L.latissima (Pic, 1932), L.lianzhouensis Long, 2000a, L.rondoni Kimoto & Gressitt, 1979, L.rufometallica (Pic, 1923), L.semipunctata (Fabricius, 1801), and L.yuae Long, 2000b. Liliocerisdiscrepens and L.rondoni were new records from China. Two synonyms are proposed: Liliocerisxinglongensis Long, 1988, syn. nov. of L.consentanea (Lacordaire, 1845), and Criocerisrufimembris Pic, 1921, syn. nov. of L.semipunctata. An identification key, descriptions, habitus photographs, geographic distributions, host plants and habitats (if available), are provided for these species.


Introduction
Lilioceris Reitter, 1913 has a high number of species in Criocerinae Latreille, 1804, which to date contains approximately 150 species worldwide, and at least 70 species in Asia (Monrós 1960;Heinze and Pinsdorf 1962;Gressitt 1965;Warchałowski 2011;Bezděk and Schmitt 2017).The genus is widely distributed in tropical and subtropical parts of the world, with the highest species diversity found in the Oriental Region.There are many taxonomic works focusing on regional species of the genus (e.g., Jacoby 1904Jacoby , 1908;;Gressitt and Kimoto 1961;Heinze and Pinsdorf 1962, 1963, 1964;Kimoto and Gressitt 1979;Tishechkin et al. 2011;Warchałowski 2011;Xu et al. 2021), but still many similar species are difficult to identify based on existing keys, and more revisionary work on species group is needed.
At present, three species groups in Lilioceris have been recognized and reviewed: the L. impressa group (Tishechkin et al. 2011), the L. neptis group (Xu et al. 2021), and the L. sinica group (Xu and Liang 2022).Here, we established the fourth, the semipunctata group, including eleven species.Some species have long been confused and misused, and the characteristics listed by different researchers are different.For example, Kimoto and Gressitt (1979) listed simple characteristics of L. discrepens and L. latissima in a key, and provided illustrations for the sternum and episternum.However, we examined and compared the types (Figs 9,10) and found that their identifications were wrong.Therefore, these species need to be revised.
In this article, we compare types and re-identify syntypes, establish a new species group, propose two synonyms, and provide a key to aid identification.

Materials and methods
Specimens from several museums and collections were examined.The collections cited in this article are indicated by the following abbreviations: Except as noted, all specimens examined are deposited in IZCAS.Dry specimens were soaked in hot water for 1-2 h to soften the body.The abdomen was opened at its latero-apical margin and genitalia were pulled out using forceps.Genitalia were soaked in warm 10% KOH for 1 h, and dyed in Chlorazol Black E. The basal orifice of the median lobe was injected with 100% ethanol with a micro-injector until the internal sac was fully everted.The median lobe with its everted internal sac was photographed using a large depth-of-field 3D digital microscope (Keyence VHX-1000C), and finally edited in Photoshop © .A microvial with genitalia was pinned to the specimen from which the genitalia were removed for storage.

BSM
Body length (BL) was measured from the anterior margin of the labrum to the apex of the elytra; body width (BW) was measured along the greatest elytral width.Other methods of specimen observation and preparation follow previous publications (Tishechkin et al. 2011;Li et al. 2013).Morphological terminology follows Chou et al. (1993), Tishechkin et al. (2011) and Schmitt and Uhl (2015).

Results
The Lilioceris semipunctata species group of Lilioceris Reitter, 1913 Diagnosis.Species of this group share the following characters: 1) body almost brownish red, some with metallic luster, some of the lateral metasternum or base of abdominal sternites dark; 2) antennae more or less flattened; 3) pronotum without distinct transverse impression; 4) scutellum densely pubescent; 5) elytral punctures diminishing posteriorly and absent at 1/2, 1/3, or 1/4 of apex; 6) apical portion of mesosternal process narrow, obliquely pointed, not horizontally connected with metasternum; 7) medium to large size, body length more than 7.5 mm.
Remarks.This species is very similar to L. semipunctata according to the original description and photographs of the holotype, but differs by the black femora with its middle brownish red in ventral view; sides of the abdomen are black.In L. semipunctata, the femora are brownish red; sides of the abdomen are brownish red.For this species, we have not collected sufficient specimens; therefore, the species is only listed and not treated in detail.
Diagnosis.Anterior angles of pronotum rounded, pronotal disc with two rows of fine punctures.Humeral groove of elytra indistinct, punctures sparse and diminishing posteriorly, absent on apical 1/3.Femora of mid-and hind legs without tooth.Lateral metasternum with a wide strip of pubescence.
Pronotum (Figs 3,5,37B).Anterior angles distinctly rounded, not protruding, posterior angles slightly protruding; sides strongly constricted in the middle; disc slightly raised; middle of disc with two rows fine punctures.Scutellum triangular and pubescent.
Mesosternum pubescent.Lateral metasternum with a long strip of pubescence, extending from the posterior to anterior margin.Metepisternum densely pubescent (Fig. 37C).
Leg (Fig. 27C).Femora with dense pubescence on the dorsal surface, with sparse pubescence on the ventral surface, without tooth.
Host plant and habitat (Fig. 69A, B).This species fed on Cycas revoluta Thunb.according to observation of HBL in Diaoluo Shan (Hainan).Diaoluo is one of the

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The body color of this species is variable.The abdomen and legs of the specimens from Hainan are brownish red, while the specimens from Yunnan are completely black.We have compared the genitalia of specimens from Hainan and Yunnan and found no differences.We compared the type of Lilioceris xinglongensis Long, 1988 (Figs 3, 4) with that of L. consentanea, and found no significant differences; the two species are therefore treated as conspecific.Lilioceris dentifemoralis Long, 1988: 231 (China: Hainan).Diagnosis.Anterior angles of pronotum slightly protruding, pronotal disc with two or three irregular rows fine punctures.Humeral groove of elytra distinct, punctures of elytra diminishing posteriorly, absent on apical 1/4.Femora of mid-and hind legs with tooth.Lateral metasternum with a wide strip of pubescence.
Pronotum (Figs 7,38B).Anterior angles slightly protruding, posterior angles not protruding; sides constricted in the middle; disc slightly raised; middle of disc with two or three irregular rows fine punctures.
Abdominal sternite (Fig. 28A).Lateral transverse impressions big and distinct on sternites 1-3, other areas of sternites 1-3 and all of sternites 4 and 5 densely pubescent.Leg (Fig. 28C).Femora with dense pubescence on the dorsal surface, nearly smooth on the ventral surface, femora of hind legs with a distinct tooth, and middle legs with a weak tooth.
Host plant and habitat.Host plant is unknown.We visited Jianfengling of Hainan where all Chinese specimens were collected, located in a subtropical area.The habitat is mixed primeval forest, orchards, and farmland with high temperatures, plentiful precipitation, and plenty of sunlight.The type locality, Tianchi, refers to a lake area surrounded by shrubs and tall trees.
Pronotum (Figs 9,39B).Anterior angles protruding, posterior angles not protruding; sides slightly constricted in the middle; disc flat; middle of disc with four or five irregular rows of fine punctures.Scutellum triangular and pubescent.
Leg (Fig. 29C).Femora with dense pubescence on the dorsal surface, with sparse pubescence on the ventral surface, without tooth.
Distribution (Fig. 67).China (Yunnan, Guangxi); Laos; Thailand; Vietnam.New record from China Host plant and habitat (Fig. 70A, B).This species is fed on Smilax sp. according to our observations in Yunnan.One locality of this species in Maku village (Yunnan, Gongshan, Dulongjiang) is situated at the subtropics.The habitat is primeval forest, which is characterized by both high temperatures and humidity, plentiful precipitation, but without much sunlight.The forests are composed of tall trees, woody vines, and weeds.
Remarks.The body size of this species is very large for the genus.It is similar to L. latissima, but differs by the pronotal disc with four or five irregular rows of fine punctures, and the lateral metasternum with a long arcuate strip of pubescence (in L. latissima, the pronotal disc has two rows of fine punctures, and the lateral metasternum is smooth).
Lilioceris discrepens was described by Baly (1879) from Laos, subsequently listed with simple characteristics in a key by Kimoto and Gressitt (1979: 220) and an illustration of the sternum and episternum (Kimoto and Gressitt 1979: 224, fig. 14j), but we examined the types (Figs 9, 10) and found that the identification was wrong.The lateral metasternum has a long strip of pubescence, rather than the sparse pubescence as illustrated by Kimoto and Gressitt (1979).61A-C).Tergites 8 and 9, sternites 8 and 9 sclerotized, posterior areas of tergite 8, sternite 8, and apodemes with pubescence, spiculum gastrale Y-shaped and long; ovipositor with dense pubescence, distal part of ovipositor cylindrical, long and with a protuberance; spermatheca simple and hooked.
Host plant and habitat.Unknown.

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Remarks.The species is most similar to L. yuae, but differs by the antenna being brownish red, without a metallic luster, and the antennomeres 7-10 as long as wide (Fig. 30D); the lateral transverse impressions are larger on the abdominal sternites 1-4 (Fig. 30A); the ventral sclerite is long and thin (Fig. 54C,  D). (in L. yuae, the antenna is brownish red with a blue metallic luster, antennomeres 7-10 are distinctly longer than wide (Fig. 36D); lateral transverse impressions are very small on all sternites (Fig. 36A); and the ventral sclerite is shorter and thicker (Fig. 55C, D)).
Leg (Fig. 31C).Femora with dense pubescence on the dorsal surface, with sparse pubescence on the ventral surface, without tooth.
Host plant and habitat.This species is fed on Smilax corbularia Kunth according to our observation in Yunnan.One locality of this species in Baihua Shan (Yunnan, Xishuangbanna, Jinghong) is situated at subtropics.This habitat is mixed primeval forest and farmland, with high temperatures, plentiful precipitation, and plenty of sunlight.
Remarks.Lilioceris latissima can be distinguished from other species in this group by the smooth metasternum (Fig. 41C) and middle of pronotal disc with three or four rows of fine punctures (Fig. 41B).
This species was described by Pic (1932) from Tonkin, Vietnam.Subsequently, Kimoto and Gressitt listed simple characteristics in a key (1979: 220) and provided an illustration for the sternum and episternum (1979: 224, fig. 141).We examined the types (Figs 13, 14) and found that their identification was wrong: the lateroposterior corner of the metasternum of the type is glabrous, rather than densely pubescent as illustrated by Kimoto and Gressitt (1979).Furthermore, the distributions in Thailand and Laos listed by Kimoto and Gressitt need confirmation, and we therefore exclude them in this study.Lilioceris lianzhouensis Long, 2000a: 262 (China: Guangzhou).
Pronotum (Figs 15,42B).Anterior angles slightly protruding, posterior angles not protruding; sides constricted in the middle; disc slightly raised; middle of disc with two or four irregular rows of fine punctures.
Leg (Fig. 32C).Femora with dense pubescence on the dorsal surface, nearly smooth on the ventral surface, without tooth.
Distribution (Fig. 68).China (Hunan, Yunnan, Guangxi, Guangdong, Hainan).Host plant and habitat.Host plant is unknown.One collecting place, Hongkan Reservoir in Baisha, Hainan, is located in a subtropical area.The habitat is a lake area surrounded by primeval forests with high temperature and humidity.

Lilioceris rondoni
Pronotum (Figs 17,43B).Anterior angles protruding, posterior angles not protruding; sides slightly constricted in the middle; disc flat; middle of disc with four or five rows of fine punctures.Scutellum triangular and pubescent.
Leg (Fig. 33C).Femora with dense pubescence on the dorsal surface, with sparse pubescence on the ventral surface, without tooth.
Host plant and habitat (Fig. 71A, B).The host plant of this species is Smilax bracteata Presl according to our observation in Xishuangbanna.It shares the same habitat with L. latissima in Baihua Shan of Xishuangbanna, Yunnan.
Remarks.This species is most similar to L. semipunctata, but differed by the strongly convex vertex (Fig. 43A); the disc of pronotum has three or four rows of fine punctures (Fig. 43B); the posterior part of the dorsal sclerite is slightly widened in dorsal view, quadrate at the apex (Fig. 48C).In L. semipunctata, the vertex is slightly convex (Fig. 44A); the disc of the pronotum has two rows of fine punctures (Fig. 44B); and the posterior part of the dorsal sclerite is parallel in dorsal view, rounded at apex (Fig. 50C).
This species has obvious sexual dimorphism: the middle of the abdominal sternites have a dense pubescence in the male, but the middle of these abdominal sternites are smooth in the female.
Host plant and habitat (Fig. 73A, B).Host plant of this species is Smilax bracteata Presl.and Lilioceris semipunctata shares the same habitat as L. latissima according to our observations in Baihua Shan of Xishuangbanna, Yunnan.
Remark.Fabricius (1801: 472) described Lilioceris semipunctata based on specimens from Sumatra.We examined three syntype photographs of L. semipunctata.They obviously belong to two different species: the specimen with the identification label and locality is here designated as lectotype (Figs 21,22).The other two specimens (TYPE / zmuc 00031032; TYPE / zmuc 00031031) with no label and no locality are here excluded from the type series.These two specimens with a glabrous metasternum are clearly different from the lectotype, but identical to Lilioceris latissima.Lacordaire (1845: 559) redescribed Lema semipunctata based the specimens from Java, and indicated that Lema dehaanii (Guérin-Méneville, 1844) was completely similar to L. semipunctata, and moved it to the genus Crioceris.Then Kimoto and Gressitt (1979: 230) synonymized L. dehaanii with L. semipunctata, and subsequent researchers followed this treatment.
We also examined a syntype of Lilioceris rufimembris (Pic, 1921) in NHHN, and no significant morphological differences were found from the lectotype of L. semipunctata, except for two rows fine punctures on pronotum (Fig. 23), which are absent in the lectotype of L. semipunctata (Fig. 21).In IZCAS, there are more than 150 specimens from Yunnan that perfectly match these two types of punctures on the pronotum.We compared all these specimens and found that there were no significant morphological differences including male genitalia.Therefore, we conclude that L. semipunctata and L. rufimembris are conspecific, and that the pronotal punctures in L. semipunctata can be variable.Distribution (Fig. 68).China (Guangxi, Guangdong, Hainan).Host plant and habitat (Fig. 74A-D).This species feeds on Smilax lanceifolia Roxb.according to our observation in Hainan.One locality of this species, near a road of Gaofeng village (Hainan, Baisha), is situated at the subtropics.The habitat is secondary forests, which is characterized by high temperature and humidity, plentiful precipitation, and much sunlight.The forests are open and composed of tall trees, woody vines, and many weeds.Remarks.Lilioceris yuae is very similar to L. cyanicollis but differs by the body color and pubescence on the metasternum; the body of L. yuae is brownish red, and only the head, antenna, leg, and lateral metasternum have a very weak blue metallic luster; lateroposterior corner of metepisternum with sparse pubescence.In L. cyanicollis, the elytra and abdomen are brownish red, pronotum is brownish red or blue, and the other areas are blue with a distinctly blue metallic luster; and the metepisternum is glabrous.However, the male genitalia and the female reproductive organs of these two species are very similar.We temporarily treat L. yuae as distinct species, as more material is needed to conclusively elucidate their relationship.

Figure 67 .
Figure 67.Distribution map of Lilioceris spp.(all marked location information is derived from the labels of available specimens).

Figure 68 .
Figure 68.Distribution map of Lilioceris spp.(all marked location information is derived from the labels of available specimens).