Calleida Latreille, 1824

Calleida Latreille, 1824: 132 . Type species: Carabus decorus Fabricius, 1801 [nec Panzer, 1800] from North America (designated by Andrewes, 1939) = Calleida cordicollis Putzeys, 1845 .

Callida Latreille et Dejean,1824, sensu: Csiki, 1932: 1439; Jedlička, 1963: 432; Habu, 1967: 121.

Spongoloba Chaudoir, 1872: 152 . Type species: Calleida fulgida Dejean, 1831 from North America (designated by Habu, 1960).

Lecalida Casey, 1920: 288 . Type-species: Lecalida pimalis Casey, 1920 from North America = Calleida platynoides Horn, 1882 .

Callidiola Jeannel, 1949: 947 . Type species: Calleida chloroptera Dejean, 1831 from Asia.

Auchenotricha Jeannel, 1949: 952 . Type species: Callidiola marginalis Jeannel, 1949 from Madagascar.

Goniocallida Jeannel, 1949: 952 . Type species: Calleida fastuosa Klug, 1833 from Madagascar.

Taxonomic treatment. Calleidina are here treated in the narrow sense of Casale (1998), Ball & Hilchie (1983), Ball & Bousquet (2001) (= Callidides of Chaudoir, 1872, pars; Calleidini of Jeannel, 1949, pars; Callidina of Habu, 1967, pars; Callidini of Basilewsky, 1984; Agrina “Calleidida group” of Erwin, 2004), as a subtribe of the tribe Lebiini of the subfamily Harpalinae, in agreement with the family-group ranks codified by the ICZN.

The Calleidina is a very diverse assemblage of genera and species of carabid beetles distributed in all continents, with their highest diversity in intertropical areas. Most of them are specialized arboreal beetles, tied to foliage in forest canopies.

Main features of Calleidina are elongate, narrow, normally (but not always) macropterous body. The most specialized arboreal species present the characters made evident by several authors (see Erwin & Pogue, 1988; Stork, 1987), common to other Carabidae specialized for living in canopy: bright colours; depressed head capsule; hemispheric eyes; deeply lobed fourth tarsomeres; numerous subtarsal setae, and pectinate tarsal claws; obsolete integumental microsculpture, and often metallic, polychromous integuments.

Main diagnostic characters of Calleidina (auctorum) in respect to other Lebiini are the following, in a peculiar combination (Casale, 1998): 1, body glabrous or scarcely pubescent (in the lativittis species group); 2, head without suborbital setae; 3, mandibles short and widened near base, normally with evident outer scrobes; 4, apical labial palpomere moderately or markedly dilated; 5, pronotum cordiform, rectangular or sub-quadrate, with straight or slightly convex, not medially lobate base; 6, fifth elytral interval without setiferous pore in general (exceptionally with additional pores); 7, tarsi of “dilated form" (sensu Habu, 1967), more or less depressed, dorsally furrowed, the fourth tarsomere strongly bilobed in arboreal species, stout in Anomotarus and some terrestrial taxa; 8, claws pectinate; 9, in male pro- and mesotarsi with the three basal tarsomeres dilated and clothed beneath with two rows of whitish ventral adhesive hairs.

Median lobe of aedeagus pleuropic left, with short preapical orifice; endophallus normally with an evident copulatory piece, or flagellum and/or spines (unarmed only in a few species). Spermatheca elongate, sac-like or ductus-like, with or without a long basal ductus, and with developed gland inserted in different positions; female gonocoxite II without lateral and median (inner) ensiform setae, without preapical sensory furrow and nematiform setae, and with numerous short apical and preapical spinae and setae (reduced in or virtually absent from only some Australian Demetrida species).

Monophyly and relationships. Reconstruction of the phylogeny of Lebiini as a whole, or of different subtribes, has been attempted by several authors (for an overview and synthesis see Ball, 1982; Ball & Hilchie, 1983; Basilewsky, 1984; Casale, 1998).

In a recent contribution based on molecular data, Ober & Maddison (2008) showed that Lebiini (in the current sense of authors) are not monophyletic. Nevertheless, they stressed that the clade including Agrina, Physoderina, Metallicina and Calleidina is strongly supported, appears in most phylogenies and is largely concordant with the calleidine phylogeny proposed by Casale (1998), based on cladistic analysis of morphological features. Furthermore, new data show—as recently suggested by Shi et al. (2013) from the morphological study of all genera of this subtribe—that Physoderina, including Physodera, Lachnoderma and some others, are more closely related to Calleidina than to Metallicina or Agrina.

Diagnostic characters of Asiatic Calleida species (in the widest sense, including Callidiola of Authors). The genus Calleida as a whole shows the character states of the subtribe Calleidina (see above). The main features of Asiatic species are the following (see also Habu, 1967, for the Japanese species):

Head wide, eyes large and convex; two supraorbital setae on each side present. Antennae slender, densely pubescent from fourth segment; the third segment longer than the fourth. Labrum transverse, dilated towards apex, somewhat sinuate at apex, with six setae; apical maxillary palpomere almost cylindrical or slightly tumid near the middle, truncate at apex; apical labial palpomere tumid or securiform (usually more dilated in males than in females), obliquely truncate at apex; mentum deeply emarginate, with two rather short setae at base of median tooth, median tooth present, rounded or somewhat emarginate at apex, epilobes distinct; ligula dilated at apex, truncate or slightly sinuate at apex, with two setae, exceptionally with some additional setae; paraglossae rather wide, not or slightly extending beyond ligula, glabrous.

Pronotum cordiform, sub-quadrate or slightly transverse; lateral margins more or less widely explanate. One mid-lateral seta and one basolateral seta on each side present.

Elytra weakly convex or depressed, glabrous or pubescent (in the lativittis species group); striae moderately or markedly impressed. The third interval with two discal and one apical setiferous pores; exceptionally with additional pores on the fifth and seventh intervals; basal pore present. Apical margin not or sinuate, beaded; outer apical angle obtuse, widely rounded, or acutely prominent in the C. terminata species group; apical sutural angle not dentate.

Metathoracic wings fully developed in all species. Legs short or moderately elongate; protibiae with cleaning spur fairly distant from the ventral margin. Tarsi of dilated form (sensu Habu, 1967), glabrous on the dorsal side; the basal three tarsomeres of meso- and metatarsi longitudinally grooved at middle on the dorsal side; the fourth tarsomere deeply bilobed, its lobes not or slightly widened at apex. Pro- and mesotarsi in males with the three basal tarsomeres clothed beneath with two rows of whitish ventral adhesive setae. Tarsal claws pectinate on the inner side.

Metepisterna longer than wide; abdominal sternum VII with apical margin moderately or deeply notched in the middle in males, not notched in both male and female only of C. puncticollis sp. n., with a variable number of setae on each side (see above, and the following key to the species groups).

Male genitalia: aedeagus slender, not distinctly or moderately bent, pleuropic left (figs. 21, 32-34). Ventrolateral margin of median lobe straight, undulate (figs. 75-77), or in some Asiatic species irregularly explanate ventrally and, in lateral aspect, forming one or two distinct ventral lobes (figs. 45, 51). Endophallus generally with two (one in some of the C. terminata species group) spine-like narrow chitinized copulatory pieces, close to each other, exceptionally with one elongate and bent flagellum in C. gressittiana sp. n. (fig. 21), or without copulatory pieces in C. puncticollis sp. n. (fig. 27). Left paramere wide, right paramere distinctly narrower than left paramere, apex very short, truncate or notched.

Female genitalia: Female reproductive tract with spermatheca sac-like or digitiform, in several examined Asiatic species with a basal projection on which the spermathecal gland duct is inserted (figs. 63, 74), in some species with a developed basal sclerotized plate (annulus receptaculi, figs. 55, 60) or surface more or less whorled (figs. 26, 63); spermathecal pedicel (basal duct) more or less expanded in the basal half, slender in the apical half, shorter or longer than the spermatheca, in some species curved or curly (fig. 63), and/or with a apical protuberance (figs. 63, 74). Spermathecal gland duct inserted at base of spermatheca, generally laterally inserted (e.g. fig. 39), exceptionally ventrally inserted in C. puncticollis sp. n. (fig. 26); gland duct long and slender, generally longer than spermatheca; glandular area slightly or abruptly inflated, generally about as long as the duct, exceptionally much shorter (fig. 39) or longer (fig. 63); atrium more or less protuberant, on the base of glandular area (figs. 60, 63). Term for female reproductive tract mainly follow Liebherr & Will (1998). Gonocoxite I of ovipositor long and wide; gonocoxite II, as a rule in Calleidina, without preapical sensory furrow and nematiform setae or marginal ensiform setae; apex obliquely truncate, with developed apical fringe of membranous extension; outer margin with one or two setae at sub-apical area, inner margin with a few hairs or setae.

Other genera of Calleidina in Asia. Several species of other different genera of Calleidina are recorded from Asia: in particular, members of the genera Agatus Motschulsky, 1845, Anomotarus Chaudoir, 1875, Glycia Chaudoir, 1842, Lipostratia Chaudoir, 1872, Merizomena Chaudoir, 1872, Paraglycia Bedel, 1904, Plochionus Dejean, 1821 and Stenocallida Jeannel, 1949 (the latter recorded only from Yemen) (see Kabak, 2003, all cited as Agrina, and Kabak, 2017, as Calleidina). Genera Anchista Nietner, 1856 and Taicona Bates, 1873 have been moved into Physoderina (Shi et al., 2013) (see below)

Six genera have Palaearctic and/or Afrotropical distributions. Two genera ( Plochionus and Anomotarus) have pantropical distributions, with a few species distributed in the Oriental Region. Some of them are widely distributed.

Sibling Asiatic genera and species attributed or similar to Calleidina, but members of other groups of carabid beetles. As recalled by Casale (1998), with emphasis on the Neotropical Region and Africa, in tropical forests of different continents, species of different carabid genera present the same patterns of size, shape and colours, which in some Lebiini involve possible batesian and müllerian mimetic complexes with other beetles (not only Carabidae) in the same area. These convergences make the study of this group particularly interesting in tropical environments, but are of no help in the study of Calleidina from the taxonomic standpoint, and make their discrimination difficult. The possible significance of this kind of convergence has been also debated in different previous papers on mimicry in carabids of different groups (see, for a comment, Ball, 1979: 612). This occurs in Asia with several members of the platynine genus Euplynes Schmidt-Göbel, 1846 .

Therefore, it is not astonishing that some Asiatic carabid taxa have been attributed, in the past and even recently, to Calleidina, but actually belong to other Lebiomorph groups, or to Platynini . Among others, we should recall these examples:

Lebiini . Two genera, Anchista and Taicona, previously placed in Calleidina (see Habu, 1967; Kabak, 2003; Lorenz, 2005), have been moved into Physoderina (Shi et al., 2013). Endynomena and Lachnoderma as well, previously treated as Calleidina (Habu, 1967), are currently treated as Physoderina (Lorenz, 2005; Shi et al., 2013).

Hexagoniini . Calleida cyclops Matsumura, 1910 from Ako (Japan), later attributed to Ophionea Klug, 1821 (Odacanthina), is currently treated as member of the genus Hexagonia Kirby, 1825 (Hexagoniini) (Habu, 1982; Lorenz, 2005).

Platynini . 1. Callidagonum Lorenz, 1998 (replaced name for Callidula Jedlička, 1955; type species: Callidula pallida Jedlička, 1955). The type species of this genus, from Sikkim, presents the same colour pattern and shape as Calleida splendidula . The genus was correctly attributed to “ Agonidae ” by Jedlička (1955) in its original description.

2. Several species of the platynine genus Euplynes Schmidt-Göbel, 1846 are Calleida -like (for instance: E. callidoides [Chaudoir, 1879]). The recently described genus Pseudocalleida Kirschenhofer, 2010 (type species: P. pahangensis Kirschenhofer, 2010, from W Malaysia: Pahang; type material not examined), after the characters provided in the original description and illustration (elytra with apical margin not truncate, tarsal claws smooth on the inner side: see Kirschenhofer, 2010), is not a member of Lebiini Calleidina, but is very probably a junior synonym of Euplynes .

Life history and larval features. Both adults and larvae of Calleidina live on leaves and trunks of various plants, attacking mainly Lepidoptera larvae; many tropical species are canopy specialists.

Habu (1967) provided an exhaustive synthesis of the life history of some Asiatic species, based on observations in the field made by Andrewes, Dammerman, Ito, Leefmans, Minamikawa, Ramachandra Rao, Umeya, Takenaka, and by himself. In particular: Calleida splendidula (Fabricius) was often observed to destroy some caterpillars in Southeast Asia. It devours the tea leaf-rollers, Cydia leucostoma Meyrick (Tortricidae), Caloptilia theivora (Walsingham) (Gracillariidae) and Homona magnanima Diakonoff (Tortricidae) in Java and it is predaceous on larvae and pupae of Nephantis serinopa Meyrick (Xyloryctidae) in India. In Malaysia, both adults and larvae of C. splendidula search for their prey, the tea leaf-rollers, inside the folded leaves. This species is predaceous on Hyblaea puera Cramer (Hyblaeidae), Pyrausta machaeralis Walker and Nephopteryx rhodobasalis Hampson (Pyralidae), and on the bamboo defoliator Algedonia coclesalis (Walker) (Pyralidae) . It was also observed destroying the tea leaf-roller Homona magnanima Diakonoff (Tortricidae), and is predaceous on Prodenia litura Fabricius (Noctuidae) in Taiwan.

Andrewes (1931b) reported that two specimens of C. pallipes Andrewes in Dhobalwala were taken under the bark of Mangifera indica and under dry stems of Croton tiglium, respectively. Later Andrewes (1933), from data provided by C.F.C. Beeson, reported that C. rapax Andrewes in northern India feeds on small caterpillars, and goes into hibernation in November. He also reported that C. splendidula, C. pallipes and C. rapax are all predaceous on the bamboo defoliator Algedonia coclesalis (Walker) at Dehra Dun (India).

More recently, Pillai & Nair (1990) reported C. splendidula as predaceous on the coconut leaf eating caterpillar, Opisina arenosella Walker (Crytophasidae), describing a method for rearing this carabid beetle in laboratory. On average, an adult beetle consumed one fully grown larva of the xyloryctid coconut pest Opisina arenosella every three days, and larvae consumed 11–13 second-instar prey during their development. Adults lived for 6–14 months in laboratory cages. Moore (2001) confirmed that this species consume larvae and pupae of O. arenosella on palms in SE Asia. Muraleedharan et al. (1992) reported C. nilgirensis Straneo (identification to be confirmed) as active predator of Cydia leucostoma Meyrick (Tortricidae) on tea in southern India. Hondô (2012) described a similar predaceous way of life on lepidopterous larvae in nine species of Parena ( Lebiini: Metallicina) in Japan.

In Japan (Habu, 1967), C. onoha Bates and C. lepida Redtenbacher hibernate under the bark of trees in the adult stage. C. onoha is one of the species which hibernate under the bark of Zelkova serrata . It passes the winter under the bark of Japanese cedar at about 1.5 m of height of trunks, and overwintering beetles in spring are in lower abundance in deciduous forests without Japanese cedars near or within them, than in those with some Japanese cedars.

In tropical forests of Asia, Calleida species don’t hibernate, but are not active in the dry season. Several species are attracted by light.

Larval features of some species of the western hemisphere have been illustrated by different authors (see, among others, Larson, 1969; Zhou & Goyer, 1993), and agree with those of other Lebiini, with some peculiar features. Concerning the Asiatic species, Gardner (1927) described the larva of C. splendidula, adding that this is predaceous on Pyrausta machaeralis Walker (Pyralidae) and Hyblaea puera Cramer (Hyblaeidae) in India.

Preliminary notes about the distribution, diversity and biogeography of the genus Calleida in Asia. As noted by Casale (1998), the number of Calleida species described so far from Asia is relatively low, compared to the high number of species known from continental Africa, Madagascar, and the Neotropical Region. Several new Asiatic species are described in this contribution, and others will be described in the following ones. The general distribution and biogeography of the genus in the Oriental Region will be illustrated and discussed later, as the revision of all taxa will be completed. Nevertheless, we can preliminarily underline the main general patterns of distribution as they appear in our study.

1 The two groups with highest species diversity, C. splendidula and C. chloroptera groups, show distribution ranges much wider than other species groups. In fact, some species are spread over all the entire Oriental Region: C. splendidula, C. lepida, and C. sultana for instance.

2 Members of some species groups have strictly allopatric distribution, showing examples of geographical replacement: in the C. lativittis species group, C. lativittis Chaudoir in E and N India, while its adelphotaxon C. excelsa Bates from Yunnan to Myanmar and Thailand, and C. jelineki sp. n. from S India; in the C. chloroptera species group, all species are strictly allopatric, with some very widely distributed and others localized to small areas.

3 Some species are endemic to islands of the Malay and Indonesian Archipelago. Several species are endemic to Borneo, including two rare and isolated new species ( C. gressittiana Casale & Shi sp. n. and C. borneensis Shi & Casale sp. n). Only two species ( C. discoidalis Heller in Mindanao and one undescribed species in Luzon) are endemic to the Philippines. One species, C. chloroptera, is endemic to Java and one undescribed species is endemic to Sulawesi. But in islands east to the Wallace Line, species diversity is relatively low, only represented by a few species of two species groups: C. splendidula and C. chloroptera groups.

4 Although generally fully winged, some species are known so far each from very restricted areas: C. viet Casale & Shi sp. n. only from southern Vietnam for instance; some others are very rare in collections, but relatively widely spread: C. puncticollis Shi & Casale sp. n. is only known from eight specimens but distributed in a rather wide range of central and southern China.