Revision of the Theopea genus group (Coleoptera, Chrysomelidae, Galerucinae), part III: Descriptions of two new genera and nine new species

Abstract This publication treats species within Theopea and closely allied genera that were not covered in the previous two revisions. Three species of Theopea Baly, 1864 are treated herein, with T. bicolor Kimoto, 1989 and T. mouhoti Baly, 1864 redescribed, and T. bicoloroidessp. nov. described. A new genus that we consider closely related to Theopea, Pseudotheopeagen. nov., is described. This new genus can be recognized with the presence of reticulate microsculpture on the vertex of the head and pronotum and presence of an apical spine on each metatibia. The following species are transferred to Pseudotheopea as new combinations: Theopea aeneipennis Gressitt & Kimoto, 1963, T. azurea Gressitt & Kimoto, 1963, T. clypealis Medvedev, 2015, T. nigrita Medvedev, 2007, T. smaragdina Gressitt & Kimoto, 1963, T. similis Kimoto, 1989, and T. subviridis Medvedev, 2012. Theopea subviridis Medvedev, 2012 is regarded as new synonym of Pseudotheopea similis (Kimoto, 1989). In addition, six new species of Pseudotheopea are described: P. borerisp. nov. from India, P. gressittisp. nov. from Philippines, P. hsingtzungisp. nov. from Laos, P. kimotoisp. nov. from Laos, Thailand, and Vietnam, P. leehsuehaesp. nov. from Laos, and P. sufangaesp. nov. from Taiwan. A second new genus regarded as closely related to Pseudotheopea, Borneotheopeagen. nov., can be recognized by possessing uniform antennae in both sexes and lacking an apical spine on each metatibia. Two new species of Borneotheopea are described from Borneo: B. jaklisp. nov. and B. kalimantanensissp. nov.


Remarks.
Populations from Laos and southwest China were misidentified. They represent Theopea bicoloroides sp. nov. (see below). Kimoto, T. bicoloroides sp. nov., and T. mouhoti Baly are characterized by their reddish brown elytra. Theopea bicolor and T. bicoloroides sp. nov.  can be easily separately from T. mouhoti (Fig. 1G-I) by the dark brown or blackish brown head, prothorax, and scutellum, and indistinct ridges in the elytra. Besides, males of T. bicolor and T. bicoloroides sp. nov. possess a median elongate sclerite internally in the aedeagus that is covered with transverse rows of hair-like setae (Figs 2C,D;3C,D). This differs from those of T. mouhoti, which lacks such hair-like setae (Fig. 4C,D). Theopea bicolor differs from T. bicoloroides sp. nov. by the relatively slender antennae (Fig. 2A) in males (length to width ratios of antennomeres V-X more than 3.0 in T. bicolor, relatively broader antenna (Fig. 3A), less than 3.0 in T. bicoloroides sp. nov.), the narrowly rounded apex of the ventral surface of the aedeagus (Fig. 2E) (broadly rounded apex of aedeagus in T. bicoloroides sp. nov. (Fig. 3E)), endophallic sclerite broad and without longitudinal groove in lateral view (Fig. 2D) (dorso-ventrally flattened and with longitudinal groove in lateral view in T. bicoloroides sp. nov. (Fig. 3D)), and slender notch at apex of gonocoxae (Fig. 2G) (broad notch at apex of gonocoxae in T. bicoloroides sp. nov. (Fig. 3G)).
Diagnosis. Theopea bicolor Kimoto, T. bicoloroides sp. nov., and T. mouhoti Baly are characterized by their reddish brown elytra. Theopea bicolor and T. bicoloroides sp. nov.  can be easily separately from T. mouhoti by the dark brown or blackish brown head, prothorax, and scutellum, and indistinct ridges on the elytra .
Variations. Males of Laos have apically tapering and truncate apex of aedeagus and its ventral surface have median notch not extending into basal opening (Fig. 4F).
Diagnosis. Theopea mouhoti Baly, T. bicolor Kimoto, and T. bicoloroides sp. nov. are characterized by their reddish brown elytra. Theopea mouhoti  can be easily separately from Theopea bicolor and T. bicoloroides sp. nov.  by the reddish brown head, prothorax, and scutellum, and distinct and convex ridges on the elytra. Further, males of T. mouhoti have median elongate internal aedeagal sclerites without transverse rows of hair-like setae (Fig. 4C,D Head. Eyes moderately large. Anterior part of head not modified or modified (strongly excavated and modified in P. costata group). Frontal tubercles prominent, narrow, usually produced at inner anterior angle. Penultimate maxillary palpomere not greatly swollen, apical palpomere conical. Vertex with reticulate microsculpture.
Pronotum quadrate or transverse, 1.2-1.3× as wide as long, broadest at middle, with pair of discal depressions. Anterior pronotal border absent. Lateral margins rounded or subparallel. Disc with reticulate microsculpture. Elytra. Surface almost glabrous (with scattered erect setae on apical part only) except P. similis (Kimoto); punctate and striate, usually with longitudinal ridges between two longitudinal rows of punctures, sometimes ridges reduced or absent in part. Epipleura gradually narrowed to apex. Disc with reticulate microsculpture. Legs. Procoxae globular, prosternal process reduced to thin depressed ridge but apically expanded, procoxal cavities closed. Protarsomere I more or less swollen. Metatibia simple, with apical spine. Length of metatarsomere I nearly equal to following tarsomeres combined. Tarsal claws appendiculate with basal tooth small and rounded. Metatarsomere I simple.
Abdomen. Last ventrite apically trilobate. Aedeagus always ventrally flattened, apex with shallow notch. Ventral surface with wide groove, with a constriction formed by two small triangular sclerites that are elongate in some species. Internal sac with median elongate sclerite, divided into two parts; sometimes with single or paired hook-like or longitudinal and apically tapering sclerites.
Differential diagnosis. This new genus possesses the following characters shared with Theopea Baly: the punctures on the elytra are striate, with ridges between two longitudinal rows of punctures; spaces between longitudinal rows of punctures broader when ridges are reduced or absent. But Pseudotheopea gen. nov. differs from Theopea by the presence of reticulate microsculpture on the vertex and pronotum (lacking reticulations in Theopea), with apical spine of metatibia (absent in Theopea), and antennomeres III-X usually longer and curved in males (antennomeres III-X usually swollen or modified in males of Theopea). Genitalic characters that distinguish males of Pseudotheopea from those of Theopea include the relatively longer tectum (> 0.5× as long as aedeagus) and divided median elongate endophallic sclerite in Pseudotheopea (relative shorter tectum and < 0.5× as long as aedeagus and the intact median elongate endophallic sclerite in Theopea). In females, the gonocoxae are convergent apically in Pseudotheopea (divergent in Theopea).
Remarks. All Theopea species (11 species) from East Asia studied by Lee and Bezděk (2018) and T. costata (Allard) (Lee and Bezděk 2019) are transferred to this new genus. Twelve additional species are recognized as members of Pseudotheopea gen. nov. including five species transferred from Theopea and seven new species. Two species groups are proposed here (Table 1).
Etymology. This new genus is named for its similarity with the genus Theopea Baly.
Redescription. Length 6.5-6.6 mm, width 2.4-2.5 mm. Body color  reddish brown or yellowish brown, but antennomeres III-XI more or less darker, elytra greenish bronze. Frontoclypeus  transverse and deeply excavated between eyes in males, concavity as wide as interspace between eyes; apical margin produced anterior, with clusters of hair-like setae at middle and sides, and convex at sides; with one pair of erect and slender sclerites at center, close to each other, apices rounded; with short, erect, and rounded sclerites insides at middle and sides of basal margin, margined with long hair-like setae and with longitudinal ridges at middle of basal margin. Antennae filiform in males, ( Elytra elongate, parallel-sided, 1.9× longer than wide; disc with dense, coarse punctures, arranged into longitudinal rows, with one indistinct longitudinal ridge between two longitudinal rows of punctures. Tarsomeres I of front legs slightly swollen in males; subparallel in females. Aedeagus (Fig. 7C-E) slender, 7.2× longer than wide; apex with shallow notch; tectum elongate, from apical 1/20 to basal 1/5; dorso-ventrally flattened, slightly curved in lateral view, angular at apical 1/7, straight from apex to apical 1/7; triangular sclerites small; internal sac with elongate, endophallic sclerite complex, 0.5× as long as aedeagus, composed of three sclerites, apical piece as long as basal piece, 0.45× long as entire sclerite; median piece shortest, 0.1× long as entire sclerite; with one elongate, apically sclerite located near base of apical piece; with one pair of short hook-like sclerite at sides. Gonocoxae ( Fig. 7G) elongate, both gonocoxae fused from basal 1/4 to apical 1/3; apices convergent and narrowly rounded, each gonocoxa with eight setae along lateral margin from apex to apical 1/6, with lateral processes at basal 2/5. Ventrite VIII (Fig. 7F) elongate and well sclerotized; disc with several long setae at sides and near apical margin, and with dense, short setae along apical margin; spiculum extremely slender. Receptacle of spermatheca (Fig. 7H) tightly joined with pump, pump slender and strongly curved; proximal spermathecal duct deeply inserted into receptacle, narrow and short.
Diagnosis. Pseudotheopea aeneipennis (Gressitt and Kimoto) is characterized by its color pattern: reddish brown body with bluish or greenish metallic elytra ( Fig. 5A-C). The aedeagus is characterized by its broadly rounded apex, with one additional elongate dorsal sclerite near base of apical piece, and with one pair of small hook-like sclerites at sides near apex of median apical piece .

Pseudotheopea boreri
Etymology. This new species is dedicated to Matthias Borer (Curator, NHMB), who encouraged the first author to focus his research on leaf beetles.
Variation. One male collected from Dalat has a smaller body (5.3 mm long, 2.2 mm wide) and the convex area on the elytra is indistinct and with longitudinal ridges (Fig. 20D), antennomeres IV-VI are curved (VII-XI lost, Fig. 20C), and the frontoclypeus lacks a concavity.

Diagnosis.
Pseudotheopea leehsuehae sp. nov. (Fig. 18F, H) is similar to P. clypealis (Medvedev) (Fig. 8D, F) based on the convex, distinct ridges and weak indistinct ridges intertwined on the elytra, but P. leehsuehae sp. nov. can be separated from P. clypealis by its metallic purple body (Fig. 18F, H) (golden green body in P. clypealis (Fig.  8D, F)), and concavity covered with a membranous sclerite and with one pair of erect rounded sclerites (Fig. 13C, D) (without such structures in P. clypealis (Fig. 9C, D)). Males of P. leehsuehae sp. nov. are characterized by their extremely elongate aedeagi (11.7× longer than wide) and one pair of hook-like sclerites arising from the middle of the ventral surface of the apical piece of the endophallic sclerite complex (Fig. 21C-E).

Types. Holotype
Diagnosis. Pseudotheopea sufangae sp. nov. (Fig. 5F-H) is similar to P. azurea (Gressitt and Kimoto) (Fig. 5D, E) based on distinct but not convex ridges on the elytra. It differs by possessing a broad concavity between the eye with a median ridge (Fig. 19C,D) (narrow concavity between eyes and without ridge in P. azurea (Fig. 6A)). Males of P. sufangae sp. nov. are characterized by its incised aedeagal apex (with notch in other species), and presence of only one additional elongate dorsal sclerite near the base of the apical piece of the aedeagus (Fig. 23C-E). Females of P. sufangae sp. nov. are similar to those of the sympatric species, P. sauteri. Both lack sexually dimorphic characters but female P. sufangae differ in having the frons elevated above the clypeus (Fig. 19E) (frons as same height as clypeus in females of P. sauteri (Fig. 19F)).
Etymology. The new species is dedicated to Mrs. Su-Fang Yu, who is a member of the Taiwan Chrysomelid Research Team (TCRT) for her contribution to the diversity of leaf beetles.
Diagnosis. Pseudotheopea nigrita (Medvedev) is easily recognized by its characteristic black color and dense setae on the elytra. In addition, a number of autapomorphic characters among genitalic structures are diagnostic, including the circular notch of the aedeagal apex; broad tectum, characteristic shape of the ventral sclerite, the recurved and combined lateral processes of the gonocoxae, and strongly curved proximal spermathecal duct of spermatheca.
Remarks. Most setae are missing from the elytra of the holotype (Fig. 18E), but these setae are dense on the elytra (Fig. 18F) of other specimens examined.
Variation. Reticulate microsculpture on pronotum is more or less reduced on specimens from Vietnam. A specimen from South Vietnam is metallic green in colour.
Diagnosis. Pseudotheopea similis (Kimoto) is similar to P. irregularis (Takizawa) based on the indistinct ridges on the elytra and metallic blue body. This species is characterized by the presence of dense, short erect setae on the elytra. Males of P. similis are similar to those of P. collaris in the presence of paired dorsal aedeagal sclerites, but differ in the bifurcate and symmetrical apices  (apices obliquely truncate apex in those of P. collaris).
Remarks. Pseudotheopea similis is easily recognized by the presence of dense erect setae on the elytra, which is also found on the holotype of Theopea subviridis. Such a characteristic feature supports their synonymy.
Distribution. Laos, Vietnam.  Head. Eyes moderately large. Anterior part of head not modified. Frontal tubercles prominent, narrow, usually produced at inner anterior angle. Penultimate maxillary palpomere not greatly swollen, apical palpomere conical. Vertex with reticulate microsculpture. Antenna 11-segmented, filiform and slender, uniform in both sexes; antennomere II very short, III long, 2.0-3.5× longer than II, 0.6-0.8× as long as I, 2.8-3.4× as long as wide. Pronotum square or transverse, 1.1-1.2× as wide as long, broadest at middle, with pair of discal depressions. Anterior pronotal border absent. Lateral margins rounded or subparallel. Disc with reticulate microsculpture. Elytra. Surface almost glabrous (with scattered erect setae on apical part only); punctate and striate, usually with longitudinal ridges between two longitudinal rows of punctures, sometimes ridges reduced or absent in part. Epipleura gradually narrowed to apex. Disc with reticulate microsculpture.
Legs. Procoxae globular, prosternal process apically expanded, procoxal cavities closed. Protarsomere I more or less swollen. Metatibia simple, without apical spine. Length of metatarsomere I nearly equal to following tarsomeres combined. Tarsal claws appendiculate with basal tooth small and rounded. Metatarsomere I simple.
Females. Antennae slender, unmodified. Protarsomere I not modified. Posterior margin of last ventrite regularly rounded, without incisions. Spermatheca with small receptacle and C-shaped pump. Gonocoxae with split and convergent apex, apical part usually with eight long setae, base bifurcate. Ventrite VIII longitudinal, with long setae at sides and short setae along apical margin, spiculum 2.4× as long as ventrite VIII.
Differential diagnosis. This new genus possesses the following characters shared with Theopea and Pseudotheopea gen. nov.: the punctures on the elytra are striate and ridges are present between two longitudinal rows of punctures; the spaces between two longitudinal rows of punctures are broader when ridges are reduced or absent. Borneotheopea gen. nov. is similar to Pseudotheopea gen. nov. based on the presence of reticulate microsculpture on the vertex and pronotum (lacking reticulate microsculpture in Theopea) and convergent apices of the gonocoxae in females (diverge apices in those of Theopea). However, Borneotheopea gen. nov. can be separated from other genera by the antennomeres III-X not modified in males (antennomeres III-X usually longer and curved in those of Pseudotheopea gen. nov.); absence of an apical spine on the metatibia (presence of apical spine on the metatibia in Pseudotheopea gen. nov.); swollen or modified in those of Theopea); broader aedeagus, < 6.0× longer than wide (> 7.0× longer than wide in Pseudotheopea gen. nov. and > 6.0× longer than wide in Theopea); the ventral surface entirely sclerotized and unmodified (with deep groove, short hollow area, hollow area in Theopea, or wide groove in Pseudotheopea gen. nov.); and with the undivided median elongate endophallic sclerite in males (divided median elongate sclerite in Pseudotheopea gen. nov.  IX.1997, leg. S. Jákl. Paratypes. 16♂♂, 2♀♀ (JBCB), same data as holotype.
Diagnosis. Borneotheopea jakli sp. nov. is easily distinguished from the other member of the genus, B. kalimantanensis sp. nov., based on the indistinct ridges on the elytra and metallic blue ventral surface  (distinct ridges on the elytra and yellowish brown ventral surface in B. kalimantanensis sp. nov. (Fig. 26E, F)). Males of B. jakli sp. nov. are also easily separated from those of B. kalimantanensis sp. nov. by the aedeagal apex directed ventrally and lacking angular processes (Fig. 27D) (the apex directed anteriorly and with angular process at apical 1/6 of aedeagus in B. kalimantanensis sp. nov. (Fig. 28C)), short, broad tectum (Fig. 27C) (extremely slender tectum in B. kalimantanensis sp. nov. (Fig. 28B)), absence of setae at apex of median elongate sclerite  (presence of clustered setae at apex of median elongate sclerite in B. kalimantanensis sp. nov. (Fig. 28B, C)).
Female unknown. Variation. Specimens from Sabah have slender lateral sclerites of the internal sac (Fig. 28E).
Diagnosis. Borneotheopea kalimantanensis sp. nov. is easily distinguished from the other member of the genus, B. jakli sp. nov., based on the distinct ridges on the elytra and yellowish brown ventral surface (Fig. 26E, F) (indistinct ridges on the elytra and metallic blue ventral surface in B. jakli sp. nov. (Fig. 26A-D). Males of B. kalimantanensis sp. nov. are also easily separated from those of B. jakli sp. nov. by the anterior directed apex and angular process at apical 1/6 of the aedeagus (Fig. 28C) (the apex of aedeagus directed ventrally and without angular processes in B. jakli sp. nov. (Fig.  27D)), extremely slender tectum (Fig. 28B) (short and wide tectum in B. jakli sp. nov. (Fig. 27C)), presence of clustered setae at apex of median elongate sclerite (Fig. 28B,C) (without setae at apex of median elongate sclerite in B. jakli sp. nov. (Fig. 27C-E)).
Etymology. The new species is named for its type locality. Distribution. Indonesia: South Kalimantan; Malaysia: Sabah.
Key to the species of Pseudotheopea gen. nov. and Borneotheopea gen. nov.