Research Article |
Corresponding author: Hongliang Shi ( shihl@bjfu.edu.cn ) Academic editor: Borislav Guéorguiev
© 2018 Hongliang Shi, Achille Casale.
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:
Shi H, Casale A (2018) Revision of the Oriental species of Calleida Latreille (sensu lato). Part 2: the C. discoidalis species group (Coleoptera, Carabidae, Lebiini). ZooKeys 806: 87-120. https://doi.org/10.3897/zookeys.806.30051
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The C. discoidalis species group of the genus Calleida Latreille from Asia (in the sense of
Calleida , character evolution, Oriental, spermatheca, taxonomy
Calleida Latreille, 1824 is a genus of LebiiniCalleidina (Coleoptera, Carabidae) with rich species diversity mainly distributed in tropical and subtropical regions of the Americas, Sub-Saharan Africa and Southeast Asia (
As the second part of our contributions to the Oriental species of Calleida, the present paper is mainly dedicated to revision of the C. discoidalis species group. This group is defined as follows: abdominal sternite VII with four or more setae in males (two or more on each side), six or more setae in females (three or more on each side); abdominal sternite VII notched in males (Figs
At the beginning of our work, we included all Oriental species with multisetose abdominal sternite VII (four or more setae on abdominal sternite VII in males and six or more in females) in the C. discoidalis group. But C. puncticollis Shi & Casale, although with multisetose abdominal sternite VII, was recognized as very different from all other members both from external and genital characters. Thus, we erected a separate group to accommodate this species (
When studying morphologies of each species, we found that the female reproductive tract (especially the spermatheca) has very important value of inferring species relationships of the Oriental Calleida. Thus, a preliminary phylogenetic analysis was performed to solve the systematic position of each species of the non-monophyletic C. discoidalis group, and moreover to interpret female genitalic evolution and evaluate its taxonomic value in the genus.
This contribution was based primarily on the examination of Calleida specimens from different collections which are indicated with abbreviations:
CCA Collection of Achille Casale, Torino, Italy
CCCC Collection of Changchin Chen, Tianjin, China
CDG Collection of Augusto Degiovanni, Bubano di Mordano (Modena), Italy
CPB Collection of Peter Bulirsch, Prague, Czech Republic
CRS Collection of Riccardo Sciaky, Milano, Italy
HBUM Museum of Hebei University, Baoding, China
MNHN Muséum National d’Histoire Naturelle, Paris, France
NMNST National Museum of Natural Science, Taibei, China
NMPC Národní Přírodovědecké Muzeum, Prague, Czech Republic
SCAU South China Agriculture University, Guangzhou, China
SYUM Sun Yat-sen University Museum, Guangzhou, China
The methods, terminology and taxonomic treatment follow our previous work (
TL: body total length, from the anterior margin of clypeus to the apex of elytra, measured along the suture. L: overall length, from apex of mandibles to apex of elytra, measured along the suture. PL: length of pronotum, as linear distance from the anterior to the basal margin, measured along the midline. PW: maximum width of pronotum, as greatest transverse distance of pronotum. EL: length of elytra, as linear distance from the basal ridge to the apex, measured along the suture. EW: maximum width of elytra, as greatest transverse distance of two closed elytra.
1 | All intervals of elytra with strong metallic lustre, uniformly green, bluish or cupreous, without any trace of a dull reddish patch on disc | 2 |
– | Elytra metallic green or bluish, with a reddish patch on the posterior half of disc, sometimes the reddish patch vague or narrow, but at least the inner three intervals with metallic lustre very faint (Figs |
4 |
2 | Elytra with basal ridge incomplete, extended only from shoulder to the fourth interval; elytral disc metallic bluish green to bluish purple; basal three antennomeres reddish yellow, the rest ones distinctly darkened; femora distinctly bicolor, with basal part yellowish and apical part almost black (Fig. |
[1] Calleida fukiensis Jedlička |
– | Elytra with basal ridge complete, extended from shoulder to the parascutellar stria; elytral disc metallic green to cupreous; antennae and femora uniformly reddish, or with apical antennomeres and apex of femora only weakly darkened; abdominal sternite VII with eight or more setae in females (Fig. |
2 |
3 | Head and pronotum dark brown, without trace of metallic lustre (Figs |
[2] Calleida piligera Shi & Casale, sp. n. |
– | Dorsal side, including head and pronotum, evenly metallic green (Fig. |
[3] Calleida cochinchinae Casale & Shi, sp. n. |
4 | Head, pronotum and legs brownish, markedly darkened; elytra dark metallic green, with obvious reflections of cupreous red or purple on lateral and apical areas (Figs |
[4] Calleida yunnanensis Shi & Casale, sp. n. |
– | Head, pronotum, and legs red to reddish yellow; elytra bright metallic green or a little bluish, not reddish or cupreous on lateral or apical areas. Pronotum more transverse (ratio WP/PL = 1.17–1.24), elytra short and wider (EL/EW = 1.52–1.65). Philippine Islands | 5 |
5 | Head, pronotum and elytral patch dark reddish; elytral disc metallic bluish green; elytral patch usually wider, occupying the inner five or six intervals (Figs |
[5] Calleida discoidalis Heller |
– | Head, pronotum and elytral patch reddish yellow; elytral disc bright metallic green; elytral patch usually narrower, occupying the inner three to five intervals (Figs |
[6] Calleida luzonensis Casale & Shi, sp. n. |
Callida onoha ab. fukiensis Jedlička, 1953: 146 (type locality: China: “Fukien”; holotype deposited in ZFMK), unavailable name.
Callida fukiensis Jedlička, 1963: 437, available name.
Calleida
suensoni
Kirschenhofer, 1986: 324 (type locality: China: Zhejiang; holotype deposited in ZMUC, paratypes in
C. fukiensis, holotype: female, examined through photo (Fig.
Calleida fukiensis Jedlička. 1 Habitus and labels of holotype (female, Fukien,
C. fukiensis: This species was originally described based on a single specimen (holotype by monotypy) from “Kuatun, Fukien” (= Guadun, Fujian) deposited in ZFMK (
C. suensoni: This species was described based on the male holotype and eleven paratypes, with type locality: China, Tien-Mu-Shan (= Tianmushan, Zhejiang), 30°23'N, 119°37'E. We did not examine the holotype deposited in ZMUC, but the two examined paratypes (one of them from the type locality) plus one additional male specimen from the type locality, and the original description with male genitalia illustrations (Kirschenhofer, 1986) are enough to recognize this species as identical to C. fukiensis. It is noticeable that the original literature erroneously reported that this species has two apical setae on the abdominal sternite VII in males, and four in females. But actually it has four (two on each side) setae in males, and usually six (three on each side) in females.
From the examination of abundant material for this species, including the type material cited above, we proved that the morphological features of these two taxa are coincident. So we treat C. suensoni Kirschenhofer, 1986 as a junior synonym of Calleida fukiensis Jedlička, 1963.
Fujian: 1 female, “Kuatun. Fukien. China. 27.6.46. (Tschung Sen.)” [type locality], “fukiensis Jedl. Det. Ing. Jedlička” (NMPC). 1 male, “Fujian, Chong’an, Xingcun, Sangang, 740 m”, “1960.V.17, leg. Pu Fuji” (
With the character states of the Calleida discoidalis species group, but different from all other known species by the combination of the following features: (1) elytra metallic bluish green to bluish purple; (2) antennae and femora distinctly bicolor: basal three antennomeres reddish, the following darkened, femora with basal half reddish, apical blackish; (3) elytral basal border only extended from the shoulders to interval 4; (4) abdominal sternite VII with four setae in males (two on each side), six setae in females (three on each side, in some specimens with one additional seta on one side). This species is unique amongst all known Asiatic Calleida species by its incomplete elytral basal ridge. From all other species of the C. discoidalis group, and from all other Calleida species in China, C. fukiensis can be easily distinguished by its bluish elytra and bicolored femora.
General features as in Fig.
Colour: Head, pronotum and scutellum reddish orange to maroon; palpomeres and apex of mandible dark brown to blackish; apical palpomere light yellow at apex; the basal three antennomeres, and base of fourth yellow reddish, distal antennomeres dark brown, the second to fourth antennomeres a little darkened in some specimens; elytra bluish green to bluish purple, with marked metallic reflection and sutural area darker; epipleura metallic dark blue; ventral side orange yellow; legs yellowish to reddish brown, with apical third of femur and base of tibia blackish; tarsomeres dark brown on the dorsal side.
Lustre and microsculpture: Head and pronotum shiny, with highly effaced microsculpture; elytra shiny, with fine but distinct isodiametric reticulate microsculpture and marked metallic lustre.
Head: Smooth or very finely and sparsely punctate; frons with oblique, faint wrinkles at sides; supraorbital furrows deep, interrupted before the level of hind edge of eyes; genae longer than the half length of eyes; temporae swollen, gradually narrowed towards the neck constriction; apical labial palpomere securiform, truncate at apex in males, less tumid and not truncate in females; mentum with lateral lobes triangular, the inner margins oblique; median tooth obtuse, with two short setae, inserted in the middle part of the tooth.
Pronotum: Transverse-cordiform (ratio PW/PL = 1.13–1.18), greatest width at approximate anterior third; lateral margins arcuate near the middle, slightly sinuate anteriorly to the posterior angles; posterior angles almost rectangular or slightly acute in some specimens; lateral expansions narrow; disc slightly convex, with sporadic transverse wrinkles and fine punctures.
Elytra: Elongate (ratio EL/EW = 1.64–1.68), with basal ridge only extended from the shoulders to the fourth interval; striae moderately deep, finely punctate, punctures gradually weakened in the apical part of elytra; intervals flat, finely and sparsely punctate; the eighth interval slightly tumid near apex; umbilicate series of composed of 15 pores along stria 8; apical truncation straight or slightly concave; outer apical angle thickened but not angulate.
Ventral side: Prosternum, lateral area of metasternum, and metepisterna finely pubescent; abdominal sternites with sparse and short accessory setae; sternite VII with four setae in males (two on each side), six setae in females (seldom with one additional seta on one side) (
Male genitalia (Figs
Female genitalia (reproductive tract Fig.
Endemic to China, but widespread and known from several provinces of South China: Zhejiang, Fujian, Jiangxi, Henan, Hubei, Hunan, Guangdong, Guangxi, Guizhou, Shaanxi. C. fukiensis is one of the most widely spread Calleida species in China (Map
We examined one male in NMPC labeled as “Swan-ping. Mongolei”, referring to Shuo-ping Fu, an old name for the region around Youyu county (39.98N, 112.47E, 1300 m) in northernmost Shanxi. This locality is far from all other confirmed localities of this species. We consider it to be a dubious record and was not included in the distribution map.
This species was mainly found in evergreen broad-leaf forest of southern China. Some specimens were collected on vegetation or by light trap.
Taiwan, Taoyuan county, Siling (24.65N, 121.42E, 1100 m).
Holotype, male, “Taiwan, Taoyuan, Fuhsing, Siling; leg. Changchin Chen, 1995.V.28, C.C.C.C.” (NMNST, Fig.
Calleida piligera sp. n. 9 Habitus and labels of holotype (male, Taiwan, NMNST) 10 Habitus (female paratype, Taiwan,
In Latin, piliger means setose. The specific name is referred to the remarkable high number of setae on the abdominal sternite VII in both sexes.
With the character states of the Calleida discoidalis species group, but differing from all other known species by the combination of: (1) elytra uniformly metallic green or cupreous green, without discal patch; (2) head and pronotum brownish, without metallic lustre; (3) abdominal sternite VII with eight or more setae in both males and females; (4) pronotum only with a few fine punctures along the median furrow.
Compared with Calleida fukiensis, the new species differs by the complete elytral basal ridge that reaches the parascutellar stria, the much higher number of setae on abdominal sternite VII, and the different body colour.
In this species group, the new species is somewhat similar to C. yunnanensis sp. n., but differs from that species by the elytra lacking any trace of discal reddish patch, and males with more than eight setae on the terminal ventrite. Amongst the other Calleida species of China, the new species is similar to C. klapperichi Jedlička, 1963 in general appearance, but can be readily distinguished by the number of setae on abdominal sternite VII.
General features as in Figs
Colour: Head dark brown, mouth part and antennae yellowish brown; pronotum dark brown, with lateral expansions yellowish; scutellum reddish brown; elytra uniformly metallic green, usually with cupreous reflection, without reddish discal patch; elytral suture and lateral margins yellowish brown; epipleura yellowish brown; ventral side and legs yellowish brown, femora darkened in some specimens.
Lustre and microsculpture: Head without microsculpture; pronotum mostly without distinct microsculpture, except some fine transverse meshes near the discal transverse wrinkles, and faint isodiametric meshes on the middle part of the basal area; elytra with distinct microsculpture in an isodiametric mesh.
Head: Moderately convex; frons with a few very fine punctures and oblique wrinkles laterally, distinct or very faint; supraorbital furrows deep, extended to the level of posterior edge of eyes; temporae not swollen, gradually narrowed towards the neck; genae shorter than the half length of eyes; antennae reaching the basal fifth of elytra; terminal labial palpomere strongly securiform with truncate apex in males, only slightly expanded in female; mentum lateral lobes with outer margins slightly arcuate, inner margins oblique, mentum tooth near triangular, rounded at apex, with two short setae inserted in middle part of the tooth.
Pronotum: Transverse (ratio PW/PL = 1.19–1.28), with its maximum width at about the anterior third; lateral margins gradually arcuate near the middle, straight or slightly sinuate before the posterior angles which are obtusely rounded; lateral expansions widened, explanate in front; disc slightly convex, with distinct transverse wrinkles and with a few fine punctures along only the median furrow; median furrow distinct, but interrupted before both the anterior and posterior margins.
Elytra: Elongate (ratio EL/EW = 1.65–1.75), with basal border complete, reaching the parascutellar stria; striae distinct, finely punctate, the punctures gradually weakened in the apical part; intervals slightly convex, finely and sparsely punctate; intervals without additional setigerous pores; third to fifth intervals slightly depressed at the basal fourth; eighth interval slightly tumid at apex; umbilicate series of 15–16 pores along eighth stria; apical truncation slightly concave; lateral margins distinctly thickened at the outer apical angles, which are obtusely rounded.
Ventral side: Prosternum, lateral area of metasternum, and metepisterna with fine pubescence; abdominal sternites with dense and long accessory setae; abdominal sternite VII with 8–12 setae in both male and female (four to six on each side) (Fig.
Male genitalia (Figs
Female genitalia (reproductive tract Fig.
Widely distributed in several provinces of south China: Taiwan, Shaanxi, Shanghai, Guangdong, Guangxi, Guizhou and Sichuan (Map
From many important morphological aspects, C. piligera sp. n. is very peculiar in the C. discoidalis species group: (1) the base of spermatheca has an evident annulus receptaculi, surface not whorled; in contrast spermatheca is without such structure and surface more or less whorled in all other known species; (2) terminal ventrite with four or more setae on each side in males; in contrast usually with two (exceptionally three) setae on each side in males for all other known species; and (3) gonocoxite II subulate, apex narrowed and sharp in contrast to gonocoxite II with apex more or less oblique truncated in all other known species. The similarities of spermatheca lead us to hypothesize a relationship of the new species to the C. terminata species group, in which all known species have spermatheca with annulus receptaculi and without whorled surface (
In most specimens, the elytra are metallic green, with distinct cupreous reflection (Figs
S. Vietnam: “Cochinchina”.
Holotype, female, “MUSEUM PARIS Cochinchine, Baudouin d’Aulne 1897”, “Calleida cochinchine” (MNHN, Fig.
The name is derived from the former name of the southern province of Vietnam of the former French empire (1862–1954), as indicated in the original label of the holotype.
This brilliant new species is distinct amongst all Asiatic Calleida species for: (1) abdominal sternite VII with five setae on each side in females (males unknown); (2) head, pronotum and ventral side uniformly metallic green, with marked cupreous reflection; (3) elytra rather elongate; (4) elytral apical margin strongly concave, with margins thickened at the outer apical angles but not angulate. C. cochinchinae can be easily distinguished from all other species in the C. discoidalis species group by its special coloration, but might be confused with C. viet Casale & Shi, also from South Vietnam, which has similar completely metallic greenish dorsal surface. Different from that species, the new species has a more elongate shape (EL/EW = 1.84, contrasting to 1.68 in C. viet), less prominent elytral outer apical angles, and multisetose abdominal sternite VII (males unknown, but supposed multisetose also).
General features as in Fig.
Colour: Head, pronotum, elytra (epipleura included), and ventral side uniformly metallic green with marked cupreous reflection, more evident at the elytral base and apex; clypeus and labrum blackish, palpomeres dark brownish with yellowish apex for the terminal segments; antennae and legs reddish yellow; legs with faint metallic green reflection; apex of femora, and all tarsomeres dark brownish.
Lustre and microsculpture: Dorsal surface rather shiny and polished; head and pronotum with vanished microsculpture; elytra with faint but distinct microsculpture in isodiametric meshes.
Head: Slightly convex, almost impunctate; frons with very faint transverse wrinkles at sides; supraorbital furrows moderately deep, vanished anteriorly to the half of the inner edge of eyes; temporae barely swollen, narrowed towards the neck; genae short, as long as the half length of eyes. Antennae short, only slightly exceeding the humeral angles of elytra. Terminal labial palpomere markedly dilated in females (probably securiform in males); mentum lateral lobes with outer margins straight, the inner margins oblique; mentum tooth obtusely truncate at apex, with two short setae inserted at the middle part of tooth.
Pronotum: Roundish-cordate (ratio PW/PL = 1.14), with its maximum width at about the anterior third; lateral expansions moderately wide; lateral margins reflexed and arcuate at the middle, distinctly sinuate before the posterior angles which are obtuse, not pointed at apex. Disc slightly convex, with deep transverse wrinkles and with a few punctures along the median furrow; median furrow deep, widened at base and reaching the posterior margin.
Elytra: Elongate (ratio EL/EW = 1.84), with basal ridge complete, extended to the parascutellar stria; striae deep, finely punctate; intervals flat, very finely and sparsely punctate; intervals 7–8 moderately tumid at apex; umbilicate series of 13 (right elytron) to 15 (left elytron) pores along the eighth stria; apical truncation oblique, markedly concave, margins thickened at outer apical angels, which are obtusely prominent, not angulate.
Ventral side: Glabrous; in the female holotype, abdominal sternite VII with five setae on each side, apical margin almost straight, slightly excised at middle. Male sternum unknown.
Male genitalia: Unknown.
Female genitalia (reproductive tract Fig.
Only known from the single holotype female from Vietnam: “Cochinchina”, without further information on the type locality in the label (Fig.
From the rather elongate habitus and peculiar female genital characters, C. cochinchinae sp. n. is different from all other known species in the C. discoidalis species group, but surprisingly accords with the C. lativittis species group. Amongst all examined species of Asiatic Calleida, only C. cochinchinae and species in the C. lativittis group (two species with female genitalia examined of three species of the group;
Yunnan, Pu’er city, Caiyanghe (22.60N, 101.12E, 1700 m).
Holotype: Male, “Yunnan, Simao, Caiyanghe national reserve, 1700 m, 2007.VII.28, leg. Zhao Yongshuang” (
Calleida yunnanensis sp. n. 22 Habitus and labels of holotype (male, Yunnan,
The name of the new species refers to its type locality: Yunnan, China.
With the character states of the C. discoidalis species group, but different from all other known species by the combination of: (1) elytra metallic green, with evident lustre of cupreous red or purple; (2) elytral discal reddish patch not well defined; (3) head, disc of pronotum and legs brownish, markedly darkened; (4) elytra with several very small additional setae on odd intervals; (5) abdominal sternite VII with two (seldom three) setae on each side in males, four or more in females; (6) median lobe of aedeagus with a cuneiform projection on the ventral side.
In this species group, C. yunnanensis sp. n. is somewhat similar to the two species (C. discoidalis and C. luzonensis) from the Philippines based on the presence of a reddish patch on elytral disc. But from the other two, C. yunnanensis can be distinguished by: (1) the darker color on head, pronotum, legs and ventral side; (2) the presence of cupreous reddish hue on elytra; (3) narrower pronotum and elytra; and (4) the characters of aedeagus which has a peculiar projection on the median lobe ventral surface.
In southern Yunnan, C. yunnanensis sp. n. is sympatric with C. quadricollis Straneo and one undescribed species belonging to the C. splendidula species group. These three species are similar in general features and coloration, but C. yunnanensis sp. n. can be readily distinguished by its higher number of setae on the terminal ventrite in both sexes.
General features as in Figs
Colour: Head dark reddish brown, vertex distinctly darker, approximate piceous; mouth parts and antennae yellowish brown, terminal palpomere with light yellow apex. Pronotum dark brown, with lateral expansions yellowish brown and disc slightly lighter along the median furrow; scutellum reddish brown. Elytra metallic green with marked cupreous reddish or purplish reflection, more evident in the apical area; metallic reflection gradually fainter toward the suture, forming a vague discal reddish brown patch almost without metallic lustre on the apical half or two thirds, not reaching elytral base, widest near elytral apical third or fourth, occupying the inner three or four intervals, sometimes the fifth interval also partly brownish; lateral margins, elytral suture, and epipleura brownish with faint metallic reflection. Ventral side reddish brown to dark brown, with metacoxae and adjacent metasternal area generally lighter; legs brown, femora gradually darker to apex, tibiae infuscate at base, tarsomeres yellowish brown.
Lustre and microsculpture: Dorsal surface moderately shiny and polished; head and pronotum with vanished microsculpture; elytra with faint microsculpture in isodiametric meshes, more evident in females.
Head: Slightly convex, almost impunctate; frons with oblique wrinkles aside, weakly defined or very faint; supraorbital furrows moderately deep, vanished at half of the inner edge of eyes; temporae moderately swollen, gradually narrowed towards the neck; genae longer than the half length of eyes; antennae reaching the basal fifth of elytra; terminal labial palpomere strongly securiform with truncate apex in males, also dilated but less so in females; mentum lateral lobes with outer margins straight, inner margins oblique; mentum tooth obtuse with apex truncate or slightly rounded, with two short setae inserted at the base of tooth.
Pronotum: Sub-quadrate (ratio PW/PL = 1.07–1.13), with its maximum width near anterior third; lateral expansions moderately wide; lateral margins arcuate at the middle, more or less sinuate before the posterior angles; posterior angles rectangular or obtuse, sometimes slightly pointed at apex; disc weakly convex, sometimes with very faint transverse wrinkles and a few punctures along the median furrow; median furrow distinct, but not reaching anterior nor posterior margins.
Elytra: Elongate (ratio EL/EW = 1.72–1.75), with basal border complete, extended to the parascutellar stria; striae distinct, finely punctate, with punctures gradually weakened in the apical part; intervals flat, finely and sparsely punctate; the third, fifth and seventh intervals each with more than 10 very small additional pores, slightly larger than interval punctures, each bearing one short seta, setae a little longer in the basal area of intervals; the eighth interval distinctly tumid near apex; umbilicate series of 15–16 pores along the eighth stria; apical truncation straight or very weakly concave; lateral margins distinctly thickened at the outer apical angles, which are obtusely rounded.
Ventral side: Mostly glabrous, lateral areas of prosternum at apex and metasternum with a few short setae; abdominal sternites sparsely pubescent. Abdominal sternite VII with two (the holotype bearing three setae on the right side) setae on each side in males, the inner seta placed a little forward; with four to six setae on each side in females, with the outer second seta placed a little forward; apical margin of abdominal sternite VII distinctly notched in males, straight in females.
Male genitalia (Figs
Female genitalia (reproductive tract Fig.
Known from two localities of southern and southwest Yunnan, and northern Laos (Luang Namtha) (Map
We examined one female from northern Laos (Fig.
Callida
discoidalis
Heller, 1921: 529 (type locality: Philippines: Mindanao, Davao; holotype deposited in
Holotype, male, “Davao Mindanao Baker”, “7247”, “1920/3”, “discoidalis. Typus”, “Staatl. Museum für Tierkunde. Dresden” (
Calleida discoidalis Heller. 32 Habitus and labels of holotype (male, Mindanao,
1 female, “Davao Mindanao Baker”, “Ex Mus. Coll. Agric. Phil. Is.”, “Callida discoidalis Heller compared with type H.E.A.”, “H.E. Andrewes Coll. B. M. 1945-97.” (
C. discoidalis is distinct within the Calleida discoidalis group for its evident reddish spot on the inner five or six intervals of the elytra, markedly contrasting with the lateral metallic bluish green intervals, and for having the head, antennae, pronotum, legs and ventral side completely reddish yellow. C. discoidalis is similar to C. luzonensis n. sp., from which is distinct by several characters stressed in the key to species (and see diagnosis of C. luzonensis below).
C. discoidalis could be confused with C. splendidula, sympatric in the Philippines, for their similar habitus and colour, and the evident reddish patch on the elytral disc in both species. It is, however, easily recognized by the latter for the multisetose abdominal sternite VII (in C. splendidula, the abdominal sternite VII bears only one seta on each side in males, two in females), generally larger size, and for the different shape of aedeagus.
The original description provided by
General features as in Figs
Colour: Head, antennae, pronotum, underside and legs reddish yellow. Elytra dark metallic green, with trace of bluish; disc with an evident reddish patch, one-half to two-thirds as long the elytra length, widest in apical third of elytra, occupying the inner five or six intervals; reddish patch not reaching elytral base, narrowed to apex, only with the first interval reddish at apex.
Lustre and microsculpture: Moderately shiny; head and pronotum with obsolete microsculpture; elytra with faint but distinct microsculpture in isodiametric meshes.
Head: Smooth; genae short, moderately swollen; neck constriction evident.
Pronotum: Wider than head, cordate, markedly transverse (ratio PW/PL = 1.17–1.23). Lateral margins widened and arcuate in front, reflexed and sinuate in the basal third; lateral expansions wide; posterior angles obtuse; disc with shallow transverse wrinkles. In one examined specimen, exceptionally the left side bears two antero-lateral setae.
Elytra: Moderately elongate (ratio EL/EW = 1.62–1.65), depressed, with basal border complete, extended to the parascutellar stria; striae deep, finely punctate; intervals convex at base, flattened on disc; intervals 7–8, slightly tumid at apex; apical margin obliquely truncate, with outer apical angles slightly thickened at the outer apical angles, which are obtusely rounded.
Ventral side: Abdominal sternites with sparse, short but evident pubescence. Abdominal sternite VII with two setae on each side in males, four on each side in females.
Male genitalia: As in Figs
Female genitalia: Not examined.
Only known from the Mindanao Island in the Philippines (Map
We examined five other specimens from Philippines in the collection of
Philippines, Luzon, Nagtipunan (16.22N, 121.60E).
Holotype: male, “Philippines-E Luzon Nagtipunan, Quirino V-2014” (CCA, Figs
Calleida luzonensis sp. n. 36 Habitus of holotype (male, Luzon, CCA) 37 Labels of holotype 38 Habitus (female paratype, Luzon,
The new species is named after its type locality: Luzon Island, Philippines.
C. luzonensis sp. n. is distinct amongst all Asiatic Calleida species but close to C. discoidalis for: (1) elytra with evident reddish patch on the inner three to five intervals, markedly contrasting with the lateral metallic green intervals; (2) head, antennae, pronotum, legs and underside uniformly yellow reddish; (3) terminal ventrite with two setae on each side in males, three or more setae in females. The new species is closest to C. discoidalis from Mindanao Island, both having very similar external features. Comparing with C. discoidalis, C. luzonensis sp. n. has the following differences: (1) elytral discal reddish patch usually more reduced, narrower and more prolonged; (2) elytra metallic region generally lighter and more vivid; (3) body size generally a little larger (L = 10.5–11.0mm). The following differences in the median lobe of aedeagus support these two species as distinct: (1) in C. luzonensis sp. n. the median lobe smaller in size, shorter and stouter, length about 1.9 mm; in C. discoidalis median lobe length about 2.6 mm; (2) in C. luzonensis sp. n. apical lamina wider and less developed, length 0.6 time as the basal width, apex not thickened in lateral aspect, versus in C. discoidalis, apical lamina longer and more developed, length 0.7 times the basal width, apex a little thickened in lateral aspect; (3) left margin markedly prominent near middle in dorsal aspect in C. luzonensis sp. n., and evenly curved in C. discoidalis.
As with C. discoidalis, C. luzonensis might be confused with C. splendidula, a sympatric species in the Luzon Island (Philippines) based on their similar habitus and colour, and the evident reddish patch on the elytral disc in both species. Moreover, in one female paratype from Dindin, Isabela (Fig.
The new species is, however, easily recognized from the latter by the larger size, the multisetose abdominal sternite VII (in the species belonging to C. splendidula group, abdominal sternite VII bears only one seta on each side in males, two in females), and for the different aedeagal shape.
General features as in Figs
Colour: Head, antennae, pronotum, underside and legs bright reddish yellow. Elytra bright metallic green, disc with an evident reddish patch, generally one-half to two-thirds as long as the elytra length, widest in apical third of elytra, occupying the inner three to five intervals; reddish patch not reaching elytral base, narrowed to apex, only with the first interval reddish at apex. In one examined specimen (Fig.
Lustre and microsculpture: Dorsal surface moderately shiny and polished; head and pronotum with vanished microsculpture; elytra with faint but distinct microsculpture in isodiametric meshes.
Head: Slightly convex, almost impunctate; supraorbital furrows shallow, vanished a little behind the anterior edge of eyes; temporae moderately swollen, gradually narrowed towards the neck; genae shorter than half the length of eyes; antennae reaching the basal fifth of elytra; terminal labial palpomere strongly securiform with truncate apex in males, also dilated much less so in females.
Pronotum: Wider than head, transverse-cordate (ratio PW/PL = 1.19–1.24), depressed on disc. Lateral margins widened and arcuate in front, reflexed and sinuate in the basal third; lateral expansions wide; basal foveae wide, very deep; posterior angles obtuse; disc with shallow transverse wrinkles.
Elytra: Moderately elongate (ratio EL/EW = 1.52–1.65), slightly widened in the posterior third, depressed; basal border complete, extended to the parascutellar stria; striae punctate; intervals subconvex at base, flattened on disc, sparsely punctate; the eighth intervals slightly tumid near apex; apical margin obliquely truncate, weakly concave, with margin slightly thickened at the outer apical angle, which is obtusely rounded.
Ventral side: Abdominal sternites with sparse and short pubescence. Abdominal sternite VII with two setae on each side in males, three to five setae on each side in females.
Male genitalia: As in Fig.
Female genitalia (reproductive tract Fig.
Only known from eastern Luzon Island (Quirino and Isabela provinces) in the Philippines (Map
As a conclusion of the preliminary character analysis (see remarks under each species and species groups in text and our previous contribution:
A total of 24 characters were selected, which included nine female genital characters, four male genital characters, four sexual dimorphic characters, and seven external characters. Two of the characters were multistate, whereas the others were binary. Although all characters were unordered in the phylogenetic analysis (without demonstration of character polarities), the supposed plesiomorphic states were coded “0”. A total of 18 species containing representatives of all nine species groups (defined in
1 Spermathecal basal projection: (0) absent, (1) present.
2 Spermatheca basal sclerotized plate (annulus receptaculi): (0) absent, (1) present.
3 Whorl on spermatheca: (0) strong, (1) absent or very faint.
4 Apical protuberance of spermathecal pedicel: (0) absent, (1) present.
5 Spermathecal pedicel shape: (0) near straight, (1) strongly curved or curled.
6 Spermathecal pedicel length: (0) less than or subequal to spermatheca, (1) longer than spermatheca.
7 Spermathecal gland duct insertion: (0) ventrally, (1) laterally.
8 Spermathecal glandular area: (0) not or weakly incrassate, (1) distinctly inflated.
9 Gonocoxite II apex: (0) subulate, (1) oblique truncate.
10 Ventral lobe on aedeagus: (0) absent, (1) present.
11 Male aedeagus venter: (0) plain or lobed, (1) ridged or markedly concaved.
12 Primary (ventral) copulatory piece of endophallus: (0) shorter than half length of median lobe, (1) longer than length of median lobe, flagellum-like, (2) absent or very weakly defined.
13 Secondary (dorsal) copulatory piece of endophallus: (0) well chitinized, (1) absent or very weakly defined.
14 Apex of abdominal sternite VII in males: (0) straight, (1) notched.
15 Setae on each side of male abdominal sternite VII: (0) one seta, (1) normally two, exceptionally three setae, (2) three or more setae.
16 Setae on each side of female abdominal sternite VII: (0) two setae, (1) three or more setae.
17 Female terminal labial palpomere: (0) same as in males, securiform, (1) less dilated than in males.
18 Antennomeres I–III except primary setae: (0) glabrous, (1) with accessory setae.
19 Pronotum anterior angles: (0) glabrous, (1) setose.
20 Elytra: (0) uniformly metallic, (1) disc with reddish patch.
21 Elytral apical margin: (0) straight or weakly concaved, (1) strongly concaved.
22 Elytral outer apical angles: (0) rounded or obtuse, (1) sharply angulate.
23 Elytral margins on apical outer angles: (0) more or less thickened, (1) not thickened.
24 Ratio elytra length/width: (0) between 1.5 to 1.75, (1) greater than 1.8.
Characters matrix for Asiatic Calleida and the out-group Anomotarus stigmula. “?” = missing data.
Taxa / Character | 000000000111111111122222 |
123456789012345678901234 | |
Anomotarus stigmula (Chaudoir) | 000000011002000010000010 |
C. gressittiana Casale & Shi | ?????????001110011100010 |
C. puncticollis Shi & Casale | 000000000002102100000010 |
C. excelsa Bates | 000000111000010011110011 |
C. jelineki Casale & Shi | 000000111000010010100011 |
C. corporaali Andrewes | 011000101100110010001100 |
C. viet Casale & Shi | 011000101100110010001100 |
C. borneensis Shi & Casale | 101111101000010010001000 |
C. doriae Bates | 101100100010010010000000 |
C. lepida Redtenbacher | 101101100010010010000000 |
C. sultana Bates | 101101100010010010100000 |
C. cf. splendidula (Fabricius) | 101011101000010010010000 |
C. tenuis Andrewes | 101011101000010010000000 |
C. onoha Bates | 101001101000010010000000 |
C. luzonensis sp. n. | 100000101000011110010000 |
C. cochinchinae sp. n. | 0000001?1??????100010011 |
C. piligera sp. n. | 011000100000012110000000 |
C. fukiensis Jedlička | 100000101000011110000000 |
C. yunnanensis sp. n. | 100000101100011110010000 |
Most of the materials used in the phylogenetic analysis were cited in the present and our previous contribution (
Anomotarus stigmula (Chaudoir): India (Andhra Pradesh)
Calleida lepida Redtenbacher: China (Jiangxi)
Calleida sultana Bates: China (Yunnan)
Calleida cf. splendidula (Fabricius): China (Guangxi)
Calleida tenuis Andrewes: Malaysia (Sabah)
Calleida onoha Bates: Japan (Okinawa)
The phylogeny reconstruction was performed using WIN-PAUP* Version 4.0b10 with the following parameters: Optimality criterion = parsimony; all characters were unordered; starting tree(s) was obtained via stepwise addition; addition sequence: random; number of replicates = 1000; number of trees held at each step during stepwise addition = 10; branch-swapping algorithm: TBR; steepest descent option not in effect; initial ‘MaxTrees’ setting = 100; ‘MulTrees’ option in effect; topological constraints not enforced; trees unrooted; bootstrap method with heuristic search; and number of bootstrap replicates = 1000. Branches with bootstrap values greater than 50% were maintained.
Both the equal weighting (EW) method and the successive weighting (SW) method were used in the phylogeny reconstruction. But, because of a limited number of characters, several branches in the cladogram with EW method were not expanded. So, only the cladogram generated with the SW analyses is presented in Fig.
In the cladogram generated with the SW method, the monophyly for each species group was corroborated with the exception of the C. discoidalis group which was posited as a polyphyletic group (blue color branches in the cladogram). Several relationships among species groups were suggested with relatively high reliabilities.
In our previous contribution (
The polyphyletic C. discoidalis group was composed of three isolated lineages. Lineage I containing only one known species, C. cochinchinae, was suggested as a relatively early branch in the cladogram. A monophyletic or paraphyletic group of lineage I + C. lativittis group was presumed, supported by the similarities on female reproductive tract shape (type II) and elongate elytra (character 24). Lineage II containing only one known species, C. piligera, was suggested as the sister group of the C. terminata group by a moderately high bootstrap value (= 66). Such relationship was also well founded by the presence of the basal plate on female reproductive tract (type IV). Lineage III containing four known species (three species selected in the phylogenetic analysis, which belong to the C. discoidalis group). The monophyly of lineage III was well supported (= 81) and can be also inferred by the similar female reproductive tracts (type IV).
Five species groups and two lineages of the C. discoidalis group formed a monophyletic clade (= 70), containing more than 80% described species of Asiatic Calleida. Under this clade, the C. splendidula group, C. borneensis group, C. doriae group, and C. chloroptera group are closer to each other than to the other branches. The closer relationships among them were also supported by their similar female reproductive tracts (type V).
The monophyly of these four groups and their relationships were unresolved in the cladogram, but it was suggested that the C. doriae group + C. chloroptera group are monophyletic (= 81), and C. borneensis is close to C. splendidula group.
The character evolution analysis was based on the cladogram obtained with the SW analysis. In this cladogram, only the eight characters (1–8) of female reproductive tract were analyzed. Character evolution was marked on the branches. For homoplastic transformations, the maximum parsimonious assumption was accepted with parallelisms priority to reversals. Spermathecae for all available species contained in the phylogenetic analysis were illustrated in Fig.
Possible character evolutions for female reproductive tracts in Asiatic Calleida species. Solid spots represent apomorphies; empty circles represent homoplasy. Five types of spermathecae are shown in different colors on taxa: type I (cyan), type II (red), type III (purple), type IV (blue), type V (green).
Three of the eight female reproductive tract characters were considered having apomorphic conditions (solid spots in Fig.
The remaining five characters had homoplastic or ambiguous transformations (empty circles in Fig.
Five types of female reproductive tracts were recognized corresponding to five lineages in the cladogram (different colors in Fig.
Type I Spermatheca distinctly whorled; spermathecal gland duct ventrally inserted; spermathecal base and pedicel apex not modified. Type I was supposed to be a very special and isolated type in Asiatic Calleida, only represented by the C. puncticollis group. In this species, also the absence of copulatory piece in endophallus is unique.
Type II Similar to type I, but spermathecal gland duct laterally inserted. Present in the C. lativittis group and lineage I of the C. discoidalis group. The type II also showed several primary characters, but can be separate from type I by the different position of spermathecal gland duct insertion.
Type III Spermatheca not or very faintly whorled; spermatheca with a well sclerotized basal plate (annulus receptaculi); spermathecal pedicel narrow and short. Present in the monophyletic clade of C. terminata group + C. discoidalis group lineage II. Type III is well recognized for its highly modified annulus receptaculi. The modified spermathecal base and the not or very faintly whorled spermatheca suggest a relationship to the type V.
Type IV Spermatheca distinctly whorled, with a distinct basal projection; spermathecal pedicel short and partly expanded; atrium very weakly defined. Present in lineage III of the C. discoidalis group.
Type V Present in the monophyletic clade of C. splendidula group + C. borneensis group + C. doriae group + C. chloroptera group, containing more than half of described Asiatic Calleida species. Type V is a highly modified and diverse type, recognized by the not or very faintly whorled spermatheca, strongly elongate spermathecal pedicel, and the presence of spermathecal basal projection. The elongate spermathecal pedicel is curved or curled, and/or has an apical protuberance in some taxa. The variable spermathecal pedicel in type V could be important in species definition of the C. splendidula group.
The phylogeny reconstruction partly supported the monophyly of most species groups, but the C. discoidalis group was proved to be polyphyletic with three isolated lineages. Some relationships amongst species groups were solved, but many branches had relatively low bootstrap value, while some others were unexpanded. Because of the above insufficiency, we did not tend to revise the definition of species groups on the phylogeny results, and just regard it as a very preliminary attempt to reveal species relationships.
The results of character evolution analysis showed that the female reproductive tract has very important taxonomic value in Calleida. Five distinct types of female reproductive tracts were recognized, corresponding to four monophyletic and one paraphyletic branches in the cladogram. Future studies are expected to solve character transformation polarity and better evaluate taxonomic value, when more materials will be examined, such as Afrotropical and Neotropical Calleida species, and allied genera of Calleidina such as the African genus Lipostratia Chaudoir and the Australian genus Demetrida White.
We wish to thank the following curators and colleagues for access to materials under their care: Mr Changchin Chen (CCCC), Mr Augusto De Giovanni (CDG), Dr David Wrase (