Calleida Latreille, 1824

Casale, Achille & Shi, Hongliang, 2018, Revision of the Oriental species of Calleida Latreille (sensu lato). Part 1: Introduction, groups of species, and species of six species groups (Coleoptera: Carabidae: Lebiini), Zootaxa 4442 (1), pp. -1--1: 4-8

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Calleida Latreille, 1824


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 View Figure , 32-34 View Figure ). 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 View Figure , 51 View Figure ). 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 View Figure ), or without copulatory pieces in C. puncticollis  sp. n. ( fig. 27 View Figure ). 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 View Figure , 74), in some species with a developed basal sclerotized plate (annulus receptaculi, figs. 55 View Figure , 60 View Figure ) or surface more or less whorled ( figs. 26 View Figure , 63 View Figure ); 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 View Figure ), and/or with a apical protuberance ( figs. 63 View Figure , 74). Spermathecal gland duct inserted at base of spermatheca, generally laterally inserted (e.g. fig. 39 View Figure ), exceptionally ventrally inserted in C. puncticollis  sp. n. ( fig. 26 View Figure ); 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 View Figure ) or longer ( fig. 63 View Figure ); atrium more or less protuberant, on the base of glandular area ( figs. 60 View Figure , 63 View Figure ). 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.












Calleida Latreille, 1824

Casale, Achille & Shi, Hongliang 2018



Latreille, 1824 : 132



Csiki, 1932 : 1439 Jedlička, 1963 : 432 Habu, 1967 : 121



Chaudoir, 1872 : 152



Casey, 1920 : 288



Jeannel, 1949 : 947



Jeannel, 1949 : 952



Jeannel, 1949 : 952