Biodiversity in tropical rainforests: Calleida Dejean, 1825 at the BIOLAT Biological Station, Rio Manu, Peru, with descriptions of seven new species (ColeopteraCarabidae, Lebiini). Part 1

Abstract A monographic contribution is presented on the species of the genus Calleida Dejean, 1825 at the BIOLAT Biological Station, Rio Manu, Pakitza, Peru, sampled by Terry Erwin and his co-workers. The following seven new species are described: C. solitariasp. nov., C. manuensissp. nov., C. anomalasp. nov., C. demathanisp. nov. (type locality: Peru, Tarapoto, but sampled also at Rio Manu), C. erwinisp. nov., C. marginithoraxsp. nov., and C. maximasp. nov. Relationships of each species are discussed, and a preliminary survey is presented of the genus Calleida in Peru.


Introduction
As recalled by Erwin (1983a, b), tropical rain forest canopies are "the last biotic frontier". The upper Amazon basin in South America, forming an arc included in the Eastern territories of Ecuador, Peru, Bolivia, and western Brazil, is currently cited as one of the A more recent, important contribution by Erwin et al. (2015) listed 721 species of carabids described so far from Peru, with eight Calleida species included. In recent years, I was able to examine several new Calleida species from different areas of Peru, but the present contribution dedicated to the Rio Manu area is particularly significant, showing how many close or sibling species can coexist in the same, reduced, surface area in tropical rainforests, as an example of adaptive radiation in macro-and micro-habitats (see Erwin and Pogue 1988). Furthermore, the sampling by Erwin is particularly exceptional for other reasons: first, it arises from a long-term program of research in a geographically defined surface (many species described so far from Peru having the mere type locality "Peru"); second, the area has been described in detail in all aspects from the geographical and ecological points of view; third, every sampled individual bears precise data and methods of collection on its label (arboreal species, coordinates).

Materials and methods
The following data come from a series of more than 200 examined specimens from the BIOLAT Biological Station, Rio Manu, Pakitza, Peru, at the southern edge of Manu National Park, in the upper Amazon basin of Peru. Additional Calleida specimens were examined, including type series important for the correct identification of the Peruvian material.
Male and female genitalia were dissected, dehydrated in ethanol, cleared in cold KOH, examined, and illustrated using standard techniques before their definitive inclusion on microscope slides. Line drawings were made using a camera lucida attached to microscopes Wild M-5 and M-3, and a microscope Leitz Orthoplan. For female specimens, in this preliminary contribution only features of the ovipositor (stylomeres) are illustrated for two species, a new species included known from one only individual. A key for identification of Peruvian Calleida species will be provided in a next contribution.

Taxonomic treatment and morphological terms
The genus Calleida is here treated in its widest sense, as a member (following the rules of ICZN) of the family Carabidae, Subfamily Lebiinae, Tribe Agrini, Subtribe Calleidina. This group, in the narrow sense of Casale (1998Casale ( , 2007, Casale and Shi (2018), Ball and Hilchie (1983), Ball and 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) are a monophyletic unit, markedly distinct for both adult and larval features, and the morphological characters of male and female genitalia. This fact was also confirmed by a phylogeny of Carabidae Harpalinae recently presented by Ober and Maddison (2008). The genus Calleida is treated in the wider sense, characterized by a set of morphological characters which distinguish it from other Neotropical genera of Lebiini (see Erwin 2004). The median lobe of the aedeagus is synonymous with the phallus of some authors. The proximal gonocoxite 1 and the more distal gonocoxite 2 (in the sense of Liebherr and Will 1998) are synonyms of stylomere 1 and stylomere 2 of authors, respectively.
Acronyms EL/EW ratio Length of Elytra, as linear distance from the basal ridge to the apex, measured along the suture/maximum Width of Elytra; L overall Length, from apex of mandibles to apex of elytra, measured along the suture; PL/PW ratio Length of Pronotum, as linear distance from the anterior to the basal margin, measured along the midline/maximum Width of Pronotum, as greatest transverse distance; TL body Total Length, from the anterior margin of clypeus to the apex of elytra, measured along the suture.

The area
The area and methods of sampling and mapping have been described in detail by Erwin (1991), after a survey performed from September 1987 to November 1990: the Reserved Zone of Manu (south-eastern Peru, at the frontier with Bolivia) is in the broad Manu River Valley between Manu National Park and the Rio Madre de Dios and includes, in the northern part, a narrow strip of upland ridgetop forest on both sides of the valley (Figs 1-3). The Vigilante Post, Pakitza, and the associated BIOLAT Biological Station (356 m) are approximately half way up the Rio Manu from its mouth at the Rio Madre de Dios, ca. 65 km (river distance), and ca. 32 km from the Andes, at 11°56'47"S, 071°17'00"W. The high plant diversity in the area includes more than 1850 species (Erwin 1991). Erwin (1991) illustrated both the habitats at the BIOLAT Station, which includes non-flooded forest, seasonally-flooded forests, open areas, and other vegetative habitats and microhabitats for Carabidae within or across habitats at Pakitza (canopy layer, subcanopy layer, undercanopy layer, ground cover layer, soil and gravel layer), with a list of sampling methods for each habitat. Furthermore Erwin (1991Erwin ( , 1996 provided a checklist and keys to all tribes and genera of carabids found at this site, with excellent drawings to aid future workers. Diagnosis. With the character states of the genus Calleida in the narrow sense, but characterized by the peculiar combination of the following morphological features: smallmedium sized (TL: mm 7.1); body brown, darkened on head and disc of pronotum; antennae, tibiae, tarsi, and abdominal segments rufous. Pronotum cordiform; elytra elongate, depressed, beaded at apex, with deep and deeply punctate striae. Abdominal sternum VII with one seta on each side in males. Male genitalia as in Fig. 8. Female unknown.

Descriptions of new
Description. General features as in Fig. 4. Small-medium sized: L: 7.6 mm; TL: 7.1 mm. Color: head dark brown, without reddish spots on frons; antennomeres dark rufous. Prothorax and pterothorax brown; pronotum dark reddish at sides. Elytra brown, with slight metallic bronze reflection; tibiae, tarsi, and abdominal segments dark rufous.
Luster and microsculpture: head and pronotum moderately glossy, with generally effaced cuticular microlines, hardly visible as reticulate meshes; elytra moderately glossy, with cuticular microlines evident in form of transverse pattern.
Head: wide, transverse, with evident, deep neck constriction; genae very swollen, contiguous with the posterior margin of eyes; frontal furrows deep, connected posteriorly with the supra-orbital keels; eyes very large and prominent; two supraorbital setae on each side. Prothorax: transverse-cordiform (ratio PL/PW: 8.82), with lateral margins moderately sinuate in the posterior third and constricted to the base. Lateral furrows narrow, punctate; lateral reflection slight. Disc convex, with superficial transverse wrinkles. Median furrow deep, reaching the posterior margin. Anterior angles rounded, not prominent; basal angles obtuse but evident. Base almost straight, beaded. One paramedial seta and one basolateral seta on each side present.
Abdominal sterna: sternum VII with one seta on each side in males (unknown, but probably two setae on each side in females).
Male genitalia: median lobe of aedeagus ( Fig. 8) very elongate and slender, with basal lobe markedly differentiated. Endophallus with a long flagellum, twisted at base.
Female genitalia: unknown. Geographical distribution and habitat. Known so far from the type locality only. It is probably a nocturnal forest-dwelling species at lower altitudes. The single male individual of this taxon was obtained in October, at the lights of the Biolab in Pakitza. Relationships. For its basic morphological features of shape, size, and characters of the median lobe of aedeagus and endophallus, this species seems to be close to the species of the cordicollis (= decora) species group in the sense of Casale in Desender et al (2002), from which it is markedly distinct by the different pattern of color on the dorsal side (mostly metallic in the cordicollis group), and by the different number of setae on the abdominal sternum VII (two in male in C. solitaria, three in species of the cordicollis group). Specific epithet. Geographical epithet, from the type locality of this species, the Rio Manu in Peru.

Calleida manuensis
Diagnosis. With the character states of the genus Calleida, but from the closest Neotropical species markedly characterized by the peculiar combination of the following morphological features: small-medium sized (TL: mm 7.5-8.0); body very elongate, slender and narrow; head ovate-elongate, with genae swollen and abruptly constricted at the neck; pronotum cordiform, with lateral margins deeply sinuate in the posterior third, disc transversely wrinkled and basolateral seta absent; elytra parallel sided, deeply striate, beaded at apex. Head and pronotum dark brown reddish (yellow-rufous in some individuals), contrasting in color with the dark metallic green or cupreous green elytra; antennae reddish yellow; legs rufous, with darkened, blackish brown femora. Abdominal sternum VII with two setae on each side in males (one only, exceptionally, in one examined individual), three (exceptionally four, on one side) setae in females. Male genitalia as in Fig. 9. Description. General features as in Fig. 5. Small-medium sized: L: 8.0-8.5 mm; TL: 7.5-8.0 mm.
Head: elongate, ovate, with deep neck constriction, evident also on the dorsal side; genae swollen, markedly curved and abruptly constricted at the neck, not contiguous with the posterior margin of eyes which are longer than genae and very prominent; frons smooth; frontal furrows superficial, deeply and densely punctate in front; two supraorbital setae on each side.
Prothorax: transverse-cordiform (ratio PL/PW: 0.78), with lateral margins deeply sinuate in the posterior third and constricted to the base. Lateral grooves wide, flattened, punctate; lateral reflection moderate, evident in the basal fourth. Disc convex, with dense, deep transverse wrinkles and sparse, small punctuations. Anterior angles rounded, fully effaced; basal angles obtusely marked, slightly prominent outside. Base beaded, oblique at the extreme lateral margins. One paramedial seta on each side present; basolateral seta absent.
Hind wings: fully developed. Legs: short, robust. Tarsomeres of stout form. Metatarsomeres 1-3 flattened, with shallow traces of dorsal grooves; metatarsomere 4 deeply bilobed, its lobes slightly widened and rounded at apex. Tarsal claws denticulate, each with five long teeth on the inner side.
Abdominal sterna: sternum VII with two setae on each side in males (exceptionally one, in one examined individual), three setae in females (exceptionally and asymmetrically four on one side in one examined individual).
Female genitalia: not examined. Geographical distribution and habitat. Calleida manuensis is known so far from the type locality only: Peru, Madre de Dios, Rio Manu, BIOLAT Biological Station, Pakitza, 356 m a.s.l. The specimens of the type series were obtained by insecticidal fog of bamboo, Astrocaryum and Guadua in September and October.
Relationships. Calleida manuensis, in both external features (general shape of body, disc of pronotum transversely wrinkled, and abdominal sternum VII with two setae on each side in males and three in females) and the characters state of male genitalia (endophallus with a long, bent flagellum and a small basal copulatory piece), seems to be related to C. gounellei Liebke, 1935 from southern Brazil and Paraguay. From the latter, it is easily distinguishable for the different pattern of color (the pronotum is mostly metallic golden-green in C. gounellei), ovate head (much more transverse in C. gounellei), and the markedly cordate pronotum (subquadrate, with lateral margins not or slightly sinuate basally in C. gounellei). cality as holotype, 23 June 1993, T.L. Erwin & F. Pfuno, Insecticidal fogging of mediumsized dry leaves at 6 m in dry vines of tree, 3 m, some light bamboo above but not tree leaves Tr. Tachigali / 22 Lot 498, BIOLAT/COLE 000018460, 18461, 18462, 18463, 18464; 1 ♀, same locality, date and collectors, Insecticidal fogging of medium-sized dry leaves at 7 m in dry vines of tree, 2 m,, some light broad leaf above but not bamboo, Tr. Tachigali / 22 Lot 500, BIOLAT/COLE 000018513; 1♂, 2♀♀, same locality, date and collectors, Insecticidal fogging of complex canopy at 20 m top, W / 9 vines, crown 7 m dia, with some dry leaves in vines at 6 m Tr. Tachigali / 20 Lot 505.

Calleida anomala
Specific epithet. The specific name, anomala, indicates the unique features of this new species into the genus, i.e., the absence of the paramedial (anterior) seta of the pronotum, and the peculiar shape of head and pronotum.
Diagnosis. With the character states of the genus Calleida (in the wider sense: see Materials and methods), but from the closest Neotropical species markedly characterized by the peculiar combination of the following morphological features: small-medium sized (TL: mm 7.1-8.5); body moderately elongate, depressed; head subquadrate, with genae very swollen and abruptly constricted at the neck; pronotum transversecordiform, with lateral margins markedly sinuate in the posterior third and paramedial seta absent; elytra slightly widened at the apical third, deeply striate, beaded at apex. Head, pronotum, underside and legs dark brown to blackish, moderately contrasting in color with the dark metallic green, blue green, blue violet, or violet elytra. Abdominal sternum VII with one seta on each side in males, two setae in females. Abdominal sternum IX as in Fig. 10C. Male genitalia as in Fig. 10A, B. nov., male holotype 9 C. manuensis sp. nov., male holotype 10 C. anomala sp. nov., male holotype A left lateral aspect B dorsal aspect C abdominal segment IX. Scale bar: 1 mm.
Color: Head, prothorax and pterothorax concolorous dark brown to blackish; mandibles and palpomeres at apex, and antennomeres 1 and 2 dark reddish; following antennomeres markedly darkened at apex and on the dorsal side; abdominal sterna 4-7 blackish, each with a dark reddish spot at sides; elytra dark metallic green, blue green, blue violet or cupreous violet.
Head: subquadrate, with deep neck constriction; genae swollen, markedly curved and abruptly constricted at the neck, not contiguous with the posterior margin of eyes which are longer than genae and prominent; frons smooth; frontal furrows elongate, with deep longitudinal wrinkles and sparse punctuations; two supraorbital setae on each side.
Prothorax: transverse-cordiform, slightly wider than long (ratio PL/PW:0.82), with lateral margins markedly sinuate in the posterior third and constricted to the base. Lateral grooves wide, punctate, widened and flattened in the posterior two/thirds; lateral reflection moderate, evident in the basal fourth. Disc moderately convex, with dense, deep transverse wrinkles. Basal foveae deep, punctate. Anterior angles rounded, fully effaced; basal angles right. Base straight, completely beaded. Paramedial seta at sides absent; basolateral seta on each side present.
Hind wings: fully developed. Legs: elongate. Meso-and metatibiae markedly curved; tarsomeres of stout form. Metatarsomeres 1 and 2 flattened, with evident dorsal grooves; metatarsomere 4 deeply bilobed, its lobes markedly widened and rounded at apex. Tarsal claws denticulate, each with five long teeth on the inner side.
Abdominal sterna: abdominal sternum VII with one seta on each side in males, two setae in females. Abdominal sternum IX as in Fig. 10C.
Male genitalia: median lobe of aedeagus (Fig. 10A, B) very small, stout, moderately curved; apex short, rounded distally. Endophallus with two elongate, bent and basally connected pieces; dorsal piece longer than the ventral one, and more sclerotized.
Female genitalia: not examined. Geographical distribution and habitat. Calleida anomala is known so far from the type locality only: Peru, Madre de Dios, Rio Manu, BIOLAT Biological Station, Pakitza, 356 m a.s.l. The specimens of the type series were obtained mostly by insecticidal fog of bamboo, in September and October.
Relationships. Calleida anomala is very isolated into the genus owing to some unique and peculiar morphological characters, i.e., the absence of paramedial (anteri-or) seta at sides of pronotum and the general features of head and prothorax. However, for some features such as the small-medium sized body, and the abdominal sternum VII with one seta on each side in males, two setae in females, it should be related to a large group of species widely distributed mostly in Venezuela and Brazil (many of them undescribed), including C. similis Reiche, 1842, C. subaenea Mannerheim, 1837, and C. purpuripennis Chaudoir, 1872. Calleida demathani sp. nov. http://zoobank.org/ED3D93C1-092E-4321-BDD8-559872A87E29 Figures 11, 12, 14 Type locality. Peru, Tarapoto. Specific epithet. I am very pleased to dedicate this handsome new species to the excellent French explorer and collector of carabid beetles, Marc de Mathan (1876-1908) (for a biographical note, see Moret 1993). He collected in Brazil, Ecuador, Peru, Bolivia, and Panama. More than 130 years ago, he collected a series of specimens of this new species (preserved at the MNHN) in Peru. Several specimens that I attribute to this species were more recently collected at Pakitza, Rio Manu, by T.L. Erwin and co-workers(see below, in Description: notes about the variability).
Diagnosis. With the character states of the genus Calleida in the wider sense, but from the closest Neotropical species markedly characterized by the peculiar combination of the following morphological features: medium sized (TL: mm 8.7-10.5); pronotum transverse-cordiform, with lateral furrows very wide, flattened, and lateral margins reflexed and sinuate in the basal fourth; lateral grooves very wide and flat-tened, deeply punctate and wrinkled; lateral reflection evident. Elytra elongate-ovate, not beaded at apex and with outer apical angle thickened and obtusely rounded; elytral striae deep, punctate, intervals convex. Body markedly bicolored: head blackish, pronotum brown with evident metallic green reflection on disc and lateral margins widely reddish, underside and legs dark brown, contrasting with the metallic purple or purple greenish or green elytra. Abdominal sternum VII with median deep excision and two setae on each side in males (three on one side, exceptionally, in one examined individual), three setae in females. Male genitalia as in Fig. 14. Description. General features as in Figs 11, 12. Medium sized: L: 9.5-11.2 mm; TL: 8.7-10.5 mm.
Color: head black, with two slightly distinct or vanished reddish spots on frons; antennomere 1 reddish, antennomeres 2-4 blackish, following antennomeres reddish, darkened at apex. Prothorax, pterothorax, legs, abdomen, basal margin and epipleura of elytra concolorous brown with metallic reflection. Lateral margins of pronotum widely reddish. Elytra on disc bright metallic purple red or red greenish or green with purple reflection.
Luster and microsculpture: head, pronotum and elytra glossy, with generally effaced microlines on head and pronotum, more evident on the elytral intervals, in form of isodiametric mesh pattern.
Head: wide, flattened, with evident neck constriction; frontal furrows short, wide; genae short, very swollen, not contiguous with the posterior margin of eyes; eyes very large and prominent; two supraorbital setae on each side.
Prothorax: transverse-cordiform (ratio PL/PW: 0.91), with lateral margins widely rounded in the anterior half, in some individuals constricted in front, markedly sinuate in the basal fourth. Lateral grooves very wide and flattened, deeply punctate and wrinkled; lateral reflection evident. Disc depressed, with shallow transverse wrinkles. Median furrow deep, complete, reaching both the anterior and posterior margins; basal foveae deep, wide, almost smooth. Anterior angles rounded, fully effaced; basal angles obtuse. Base oblique at sides, beaded. One paramedial seta and one basolateral seta on each side present.
Elytra: moderately elongate (ratio EL/EW: 1.60), slightly widened at the apical third; striae deeply impressed, punctate; intervals convex, with small but evident punctuations. Interval 3 with two discal and one apical setiferous punctures; umbilicate series of 15 punctures along stria 8, interrupted in the middle. Post-humeral sinuation shallow but evident; pre-apical callosity small, distinct on interval 8. Apex not beaded, with outer apical angle thickened and obtusely rounded. Interval 3 with two discal and one apical setiferous punctures; umbilicate series of 14 punctures along stria 8, interrupted in the middle.
Hind wings: fully developed. Legs: robust, tarsomeres wide, depressed. Metatarsomeres 1 and 2 shallowly grooved on the dorsal side; metatarsomere 4 deeply bilobed, its lobes widened at apex. Tarsal claws denticulate, each with six long teeth on the inner side.
Abdominal sterna: abdominal sternum VII with deep median excision and two setae on each side in males (three on one side, exceptionally, in one examined individual), three setae in females.
Male genitalia: median lobe of aedeagus (Fig. 14) relatively large-sized, depressed at sides, widened and ventrally prominent at the basal third; apical lamina obtusely rounded at apex. Endophallus with a long, basally twisted flagellum, and two subapical spines.
Female genitalia: not examined. Notes about the variability of the species. Despite the long distance between Tarapoto (type locality) and Pakitza, there are no evident morphological characters that differentiate specimens collected at Rio Manu from those from Tarapoto. Specific epithet. I am very pleased to dedicate this new, attractive species to my old friend and colleague Terry Erwin, for his huge contribution to the knowledge of the New World carabids, tropical rainforests and Neotropical biodiversity, and its conservation.
Diagnosis. With the character states of the genus Calleida (in the wider sense: see Materials and methods), but from all Neotropical species markedly characterized by the peculiar combination of the following morphological features: medium sized (TL: mm 10.7-11.7); pronotum subquadrate-transverse, widened at base, with lateral margins slightly constricted in front; elytra elongate and narrow, not beaded at apex and with outer apical angle acutely prominent; elytral striae deep, shallowly punctate. Body markedly bicolored: head, pronotum, underside and appendages concolorous brown to blackish, contrasting with the metallic purple or purple red or red greenish elytra. Abdominal sternum VII with one seta on each side in males, two setae in females. Male genitalia as in Fig. 15. Description. General features as in Fig. 13. Medium sized: L: 11.0-12.0 mm; TL: 10.7-11.7 mm.
Color: head dark brown or blackish, with two reddish spots on frons; apex of mouth parts and antennomere 1 reddish; following antennomeres brown reddish, darkened in the apical half. Prothorax, pterothorax (except the lateral margins), legs, abdomen, basal margin and epipleura of elytra concolorous brown to dark brown. Lateral margins of pronotum widely reddish. Elytra on disc metallic purple red or red greenish, with sutural interval, and lateral and apical margins dark metallic green.
Luster and microsculpture: head, pronotum and elytra glossy, with generally effaced microlines on head and pronotum, more evident on the elytral intervals, in form of isodiametric mesh pattern.
Head: wide, flattened, with evident neck constriction; frontal furrows short, with some deep transverse wrinkles; genae short, markedly swollen, almost contiguous with the posterior margin of eyes; eyes very large and prominent; two supraorbital setae on each side.
Prothorax: subquadrate-transverse (ratio PL/PW: 0.9), widened at base, with lateral margins slightly sinuate in the basal half and constricted in front. Lateral grooves wide and flattened, deeply punctate; lateral reflection moderate, evident in the basal fourth. Disc depressed, with shallow transverse wrinkles. Median furrow very deep; basal foveae deep, elongate, smooth. Anterior angles rounded, fully effaced; basal angles obtuse. Base sinuate at sides, beaded. One paramedial seta and one basolateral seta on each side present.
Abdominal sterna: sternum VII with one seta on each side in males, two setae in females.
Male genitalia: median lobe of aedeagus (Fig. 15) relatively big-sized, elongate, with wrinkled and markedly lobate ventral margin at the basal third. Endophallus with a short, curved, spine like flagellum in the middle.
Female genitalia: not examined. Geographical distribution and habitat. Known so far from the Rio Manu area. Individuals were obtained in September and October, on leaves or from insecticidal fogging on different plants.
Relationships. Based on several features (general shape of pronotum and elytra, the number and position of setae on abdominal sternum VII and the peculiar structure of the median lobe of aedeagus), C. erwini seems to be related to C. rufocuprea Chaudoir, 1872, from Eastern Brazil. It is markedly distinct from it by the larger size, the different color pattern (elytra are metallic cupreous in C. rufocuprea) and the apical margins of elytra which in C. erwini have both the outer and sutural angles obtusely but markedly prominent. Specific epithet. The specific name stresses the wide, deep lateral furrows of pronotum, which are reddish, markedly contrasting in color with the blackish disc.
Diagnosis. With the character states of the genus Calleida, but from the closest Neotropical species markedly characterized by the peculiar combination of the following morphological features: small-medium sized (TL: mm 7.5-8.0); pronotum subquadrate, widened at base, with lateral margins slightly constricted in front, sinuate in the posterior third, and lateral furrows markedly widened and deep; elytra elongate, beaded at apex, with outer apical angle rounded but thickened; elytral striae deep, punctate; intervals convex, finely punctate. Body markedly bicolored: head, pronotum, underside and appendages brown to blackish, contrasting with the metallic purple red or green elytra. Abdominal sternum VII with two setae on each side in males, three to six setae in females. Male genitalia as in Fig. 16. Fig. 17. Small-medium sized: L: 8.0-8.5 mm; TL: 7.5-8.0 mm.

Description. General features as in
Color: head blackish, without reddish spots on frons; antennomere 1 light reddish; following antennomeres brown reddish. Prothorax, pterothorax (except the lateral margins), legs, abdominal sterna, basal margin and epipleura of elytra brown to dark brown, the latter with marked metallic green reflection. Lateral margins of pronotum and sides of abdominal sterna reddish. Elytra on disc metallic purple red or red greenish, with lateral margins dark metallic green, or fully metallic green.
Luster and microsculpture: head, pronotum and elytra very glossy, with generally effaced transverse microlines on head and pronotum, more evident on the elytral intervals, in form of isodiametric mesh pattern.
Head: wide, flattened, with deep neck constriction; frontal furrows deep, short, reaching the anterior supraorbital pore, with some punctuations or longitudinal wrinkles; genae long, markedly swollen, regularly bent, almost contiguous with the posterior margin of eyes; eyes very large, hemispherical, prominent; two supraorbital setae on each side.
Prothorax: subquadrate (ratio PL/PW: 0.93), widened at base, with lateral margins slightly sinuate in the basal third and slightly constricted in front. Lateral grooves wide, widened and flattened in the posterior third, punctate; lateral reflection moderate, evident in the basal fourth. Disc sub-convex, smooth or with shallow transverse wrinkles. Median furrow deep, reaching the basal margin; basal foveae deep, obliquely impressed, smooth. Anterior angles rounded, fully effaced; basal angles obtuse. Base oblique at sides, beaded. One paramedial seta and one basolateral seta on each side present.
Elytra: moderately elongate (ratio EL/EW: 1.50), sub-convex but with a shallow depression in the basal third, slightly widened at the apical third; striae deep, punctate; intervals convex. Post-humeral sinuation shallow but evident; pre-apical callosity small, slightly distinct on interval 8, or fully vanished. Apex beaded, obliquely truncate, with outer angle rounded, but thickened. Interval 3 with two discal and one apical setiferous punctures; umbilicate series of 13 or 14 large foveate punctures along stria 8, interrupted in the middle.
Abdominal sterna: sternum VII with two setae on each side in males, three to six setae in females.
Male genitalia: median lobe of aedeagus (Fig. 16) small-sized, moderately elongate, rounded at apex. Endophallus with two long, bent and spine-like copulatory pieces connected at base.
Female genitalia: not examined. Geographical distribution and habitat. Known so far only from the type locality. Individuals were obtained mostly in September and October by insecticidal fog from different species plants. Diagnosis. With the character states of the genus Calleida (in the wider sense: see Materials and methods), but from the closest Neotropical species markedly characterized by the peculiar combination of the following morphological features: larger in size (TL: mm 14.6), elongate; pronotum moderately transverse, slightly constricted in front; elytra deeply striate, not beaded at apex, with apical external angle acutely prominent, spine-like. Body and appendages uniformly brown blackish, contrasting in color with the metallic green elytra. Abdominal sternum VII with five or six setae on each side in the female (holotype). Female genitalia (ovipositor) as in Fig. 20. Male unknown.
Luster and microsculpture: head and pronotum glossy, with highly effaced microsculpture, hardly visible as transverse mesh pattern; elytra glossy, with fine, hardly visible microlines in form of isodiametric mesh pattern.
Head: transverse, dorsally smooth, with moderate neck constriction; genae markedly convex, not contiguous with the posterior margin of eyes; eyes moderately large but very prominent; two supraorbital setae on each side.
Prothorax: moderately transverse (female) (ratio PL/PW:0.8); pronotum with lateral margins regularly arcuate, slightly constricted in front, very slightly sinuate anteriorly to the basal angles. Lateral reflection moderate, more evident anteriorly to the basal margin. Disc depressed, with very deep median furrow and very shallow transverse wrinkles. Basal foveae very elongate and deep. Anterior angles rounded, slightly prominent; basal angles obtuse. Basal margin beaded. One paramedial seta and one basolateral seta on each side present.
Abdominal sterna: sternum VII with five or six setae on each side in the female (holotype).
Female genitalia: (gonocoxites 1 and 2 of ovipositor) as in Fig. 20. Male unknown. Geographical distribution and habitat. Known so far only from the type locality. The only female individual was obtained by insecticidal fog canopy, in September.
Relationships. Based on its general features, large size, structure of female genitalia, and the high number of setae on the 7 th abdominal sternum, C. maxima sp. nov. is close to C. gigantea Casale, 2008 from southern Ecuador (Macas) and C. jeanneli Liebke, 1935 from Peru. From the latter, sympatric at the Rio Manu river (see Figs 19, 23), it is markedly distinct for its larger size, much wider head, pronotum, and elytra, different color of elytra (which are concolorous metallic purple-red in C. jeanneli), the apical outer angle of elytra acutely prominent (Figs 22, 23), and the more elongate gonocoxite 2 of ovipositor (Figs 20, 21). These taxa form a very homogeneous group of highly derived and rare, specialized species, apparently localized to a reduced area between southern Ecuador and Peru, on the Amazon side of the Andes (Casale 2008).

Concluding remarks
In his contribution to the knowledge of the Rio Manu Area, Erwin (1991) estimated the occurrence of 118 different genera of Carabidae, and for each genus furnished a hypothetical number of species. As a rule in tropical areas, Lebiini are the most numerous representatives: Calleida was estimated at 23 species and Agra was confirmed as the most diverse and speciose genus, with 44 species. Examination of his material showed that this estimation was almost exact: 22 Calleida species of the same genus coexist in sympatry in this area. Some of them are undescribed (seven described in this contribution), and a few, such as C. jeanneli Liebke (see Fig. 23) and C. demathani sp. nov., also occur in other areas of the Amazon basin. Erwin (1991), again concerning Calleida, added that "many new ones are in need of revision, and the numerous species ... will make it a troublesome group to study".
From a biogeographical standpoint, as in Casale (1998Casale ( , 2008, Calleidina, and related subtribes of Lebiini, are phyletic lineages of carabids mostly having originated and differentiated in tropical Gondwanan areas with few immigrants (or scarce extant elements) in the Holarctic Region. What about the representatives of the genus Calleida in Peru, and the Pakitza region in particular? In spite of the inadequate knowledge of this genus in the country, the available data show the presence of some phyletic lineages perfectly in agreement with the geographical position of the country, and its exceptional landscape mentioned in the Introduction. At present, some species groups seem to be absent from Peru, such as the sumptuosa species group, mostly represented in Central America. Almost all other groups of species are represented in Peru by at least one endemic (precinctive) species.
This datum once again makes evident that this area is one of the main hotspots of biodiversity in the Neotropics, in which several lineages have differentiated and speciated in the Amazon basin, or in the Pacific forests on the western side of the Andes, subsequently overlapping in the country, with some examples of adaptive radiation in sympatric conditions.
The pre-zygotic isolation amongst sibling species in the same area was favored by features of the male and female genitalia, which in Calleidina are highly complex and differentiated, compared to other Lebiini, and very informative from the phylogenetic standpoint. In this lineage, it is possible to show the modification of the median lobe of aedeagus from an elongate, sub-cylindrical, generalized shape, to a depressed, ovate or complex, ventrally lobate shape, and the transformation of the copulatory piece into a bilobed piece, or into a long or very long and twisted flagellum (see Figs 8-10, 14-16). These states have been verified in hundreds of examined and dissected individuals of Calleida species, both described and not yet described and demonstrate the occurrence, in this genus, of very homogeneous, markedly distinct to each other species groups, associated with external features, like the number of setiferous punctures in the anal abdominal segment, and other sexually dimorphic characters.
Female genitalia in Calleidina are also very informative (Casale 1998), but have not yet been carefully examined in a sufficiently high number of species. The reproductive tract, in particular, presents markedly modified structures in both spermatheca and spermathecal gland (Casale 2008;Casale and Shi 2018;Shi and Casale 2018). Concerning the ovipositor, it is well known that morphology of gonocoxites in all carabids is tied to different functions (for instance, the telescopic ovipositor of Agra may be an adaptation to lay eggs deep in existing burrows in woods or other fissures: Erwin and Pogue 1988). Highly modified and slightly sclerotized gonocoxae are typical of the most derived Lebiini, such as the parasitoid Lebiina, the arboreal Dromiina and Calleidina, and the myrmecophilous Pseudotrechina. Changes in gonocoxite 2 are surely tied to the loss of the digging function of ovipositor. In all Calleidina, the loss of ensiform setae and preapical furrow is assumed as the main synapomorphic feature.
In Calleida, the modifications of gonocoxite 2 (see Figs 20, 21) are evident, and should indicate different ways of life, and different adaptations of laying eggs on trees and shrubs. We know that Calleida species, both as adults and larvae, are actively predatory beetles on caterpillars of small-sized Lepidoptera of different families , and Erwin (1991) reported that some of them are specialized canopy dwellers; however, various species are adapted to tree crowns, dry leaf clumps suspended in the undercanopy, understory shrubs, and even tall grasses; most species are diurnal, but a few are nocturnal (e.g., C. solitaria) and often fly to lights.