Agra, arboreal beetles of Neotropical forests: pusilla group and piranha group systematics and notes on their ways of life (Coleoptera, Carabidae, Lebiini, Agrina)

Abstract Revisions of two new species groups of the genus Agra Fabricius are presented with the following species described as new: pusilla group - Agra cruciaria sp. n. (Brazil), Agra grace sp. n. (Ecuador,Perú), Agra max sp. n. (Brazil), Agra minasianus sp. n. (Brazil), Agra notpusilla sp. n. (Brazil), Agra pseudopusilla sp. n. (Brazil); piranha group - Agra ce sp. n. (Perú), Agra risseri sp. n. (Bolivia,Brazil), Agra maia sp. n. (Bolivia), Agra piranha sp. n. (Ecuador); Agra tiputini sp. n. (Ecuador). Species of these two groups have adults that are the smallest in the entire genus, although this does not indicate they are closely related based on other attributes. All species are Amazonian in distribution.


Introduction
Th e purpose of this 17th contribution in my series of papers with diagnoses of new taxa and redescriptions of known taxa in the beetle genus Agra Fabricius, 1801 (Carabidae) is to present a revision of the species in the pusilla and piranha groups. For the previous papers, see Erwin (1978Erwin ( , 1982aErwin ( , 1982bErwin ( , 1983Erwin ( , 1984Erwin ( , 1986Erwin ( , 1993Erwin ( , 1998Erwin ( , 2000aErwin ( , 2000bErwin ( , 2000cErwin ( , 2002, Erwin and Pogue (1988), and Arndt et al. (2001). New species descriptions provided here will not be the end of the story. New species of this incredibly speciose and diverse genus continue to be discovered each year throughout the Neotropics and subtropical México /Texas and northern Argentina.
Th ese beetles belong to the Tribe Lebiini, Subtribe Agrina, of which the genus Agra Fabricius is by far the largest lineage in number of species. Subtribe Agrina consists of those species formerly included in the Subtribe Calleidina (cf. Lorenz 1998Lorenz , 2005. More than 2000 neotropical/subtropical species of Agra are known from museum collections; however, only some 592 of these have been described since Fabricius erected the genus in 1801, plus three species described in the very late 1700's by Olivier and Fabricius, but originally placed in the Linnaean genus Carabus. Th e sister group was hypothesized to be an African/Madagascan group near Callidiola Jeannel 1949, however Casale (1998 argues against that. His analysis suggested that Agra is more closely aligned with Physoderina, as the adelphotaxon. Certainly, the female stylomere 2 fi ts that hypothesis in a general way and the male median lobe is not Calleida or Callidiola like. However, there is little in the overall "gestalt" that might lead one to conclude these lineages are at all related. It is time for comparative DNA analysis, I suggest, to determine where these higher lebiine lineages fall out on an updated classifi catory scheme. Unfortunately, in the fi ne contribution of Ober and Maddison (2008), the Physoderina were not included, even though they saw possible monophyly amongst Calleidina (sensu Casale 1998), Agrina, Metallicina etc. Th is will mean that specimens of the pusilla group need to be in the DNA mix of samples because of their clearly, based on structural attributes, less derived features amongst all the other species groups of this very diverse genus. And, members of Physoderina will need to be in the mix, as well.
Th e pusilla group contains what appear to be the least derived lineage of Agra and obtaining DNA from them will be crucial in locating the sister lineage -an additional reason for this present revision is making the species and their localities more broadly available, so that fresh specimens can be appropriately acquired and identifi ed. Th e piranha group apparently is more derived than the pusilla group, but both groups are composed in part of little blue beetles and I thought it better to treat them together for purposes of easier identifi cation and classifi cation.
Likely, adult Agra are predaceous on other arthropods; one specimen of another species group dissected had fragments of termites in its gut contents. Adults also have been observed drinking exudates from young new shoots and young leaves on a variety of tree species, as well as feeding on pollen (Arndt et al. 2001). Adu lts are active on tree surfaces in the canopy and along forest edges at night, as well as in suspended dry leaves in the understory; in addition, at least one species group is often found on savannah grasses. Adult tarsi are adapted for running on the surfaces of leaves with pads similar to those found in adult Chrysomelidae and Cerambycidae. Adults rest under these leaves "concealed" with legs and antennae tucked close to the body, the beetle aligned perfectly with the midrib of the leaf; in the case of grass-dwelling species they align with the culms. Agra adults are nocturnal and commonly fl y to lights at night and into Malaise traps, however, none are known so far from ground level fl ight-intercept traps. Th ese beetles have a potent defensive secretion from paired glands in their abdomen that, in fi eld tests, bats defi nitely do not like (Erwin 1978). And, males of many species of this genus have special ventral patches of setae or pubescence that suggest they may waft pheromones to attract females. If correct, these chemicals have not been explored for their composition or possible uses. Known larvae are thought to occur normally under bark of standing trees, probably in burrows of other insects, and are thought to be predatory (Arndt et al. 2001). A few larvae have been obtained by rearing from eggs of known females and with insecticidal fogging techniques before dawn (Arndt et al. 2001), the latter indicating the larvae may wander on the tree surfaces at night.

Specimens and methods
Methods and species concepts follow those previously described (Erwin and Kavanaugh 1981;Kavanaugh and Erwin 1991). Th e species validation and diagnosis format follows as closely as possible that suggested in Erwin and Johnson (2000) and as used in Erwin (2000aErwin ( , 2004. Measurements of length (ABL, SBL) and width (TW) follow those of Ball (1972) and Kavanaugh (1979): ABL (apparent body length), measured from apex of labrum to apex of longer elytron; SBL (standardized body length), equals the sum of the lengths of the head (measured from apex of clypeus to a point on midline at level of the posterior edge of compound eyes), pronotum (measured from apical to basal margin along midline), and elytron length (measured from apex of scutellum to apex of the longer elytron); and TW, (total width), measured across both elytra at their widest point.
Th e habitus images of the adult beetles portray most of the character states referred to in the keys provided. Illustrations of male genitalia are standard for descriptive taxonomy of carabid beetles. Th e habitus images of the adults were made with a Visionary Digital TM high resolution imaging system. Figure captions include an ADP number, which is a unique identifi cation number for the specimen that was illustrated or imaged and links the specimen and associated illustrations and/or image to additional information in electronic databases at the NMNH. All scale lines are 0.5mm.
Geographical data are presented for species based on all known specimens available at the time of manuscript preparation. Georeferences have been determined from locality information provided on specimen labels; only those exact Georeferences that are provided on the label are placed in quotes, otherwise I have estimated the Georeferences as closely as possible from places, mileage, etc. listed on the label and searched with Google Earth Pro. Latitude and longitude are reported in decimal degrees. Distribution maps are provided for the species (Figs 10, 11). Here, English vernacular names are proposed, as common names are becoming increasingly needed in conservation and/or agricultural and forestry applications.

Th e pusilla species-group
Members of the pusilla group are the smallest of the genus Agra and in part, of a beautiful blue color and matte luster with highly contrasting bicolored femora.
Diagnosis: Femur bicolored. Back of head rounded and sparsely punctate. Adult males with extensive ventral tarsomere pads on both front and middle legs. Prothorax markedly punctate. Elytral interneurs of uni-and/or biseriate rows of cribriform punctures. Aedeagus with narrow spatulate apex. Female stylomere 2 short and arcuate, glabrous, and apically armed with two ensiform setae, as in Fig. 7.
Note: Th e known composite range of the pusilla group extends from Amazonian Ecuador and Perú across southeastern Brazil and into the Mata Atlântica.
Brazil Agra pusilla Chaudoir, 1847 Brazil Key to the species of the pusilla group of Agra Fabricius, 1801 Derivation of specifi c epithet. Th e epithet "cruciaria" is a Latin adjective meaning "of/pertaining to the cross/torture" and is based upon the large cross on Corcovado fl ooded each night with high powered lights that attract insects by the millions and is the type locality of this species.
Proposed English vernacular name. Cross Elegant Canopy Beetle. Diagnosis. With the attributes of the genus and species-group as described above and medium sized for the pusilla group. Adults with black integument; head behind eyes and prothorax with slight brassy refl ections. Frons laterad unicarinate and rugose. Occiput with sparse punctures, some punctures with short setae.
Prothorax: Slightly broader medially, fl ared basally; surface with dense punctures, many setiferous; lateral elongate callous with single row of setiferous punctures along middle. Pterothorax: Elytron markedly convex, intervals slightly costate, interneurs of rows of somewhat irregularly shaped punctures that are double in some places, apex slightly oblique and moderately lobate, apical dentation asymmetric, lateral tooth small, acute, sutural apex slightly produced, narrowly pointed. Metasternum sparsely setiferous in male. Legs: Normal in male. Abdomen: Abdominal sterna III to VII of male moderately and bilaterally setiferous; sternum VII of male barely emarginated, corners rounded. Male genitalia: Phallus (Fig. 3) elongate and narrow with ostium elongate, nearly half the length of the phallus, apex a narrowly lobate expansion of distal end, this slipper-shaped in lateral aspect. Parameres small, left twice the size of the right, both broadly rounded. Female ovipositor: Female unknown.
Dispersal potential. Th ese beetles are macropterous and are probably capable of fl ight; they are swift and agile runners.
Way of life. Adults of other Agra species are found in the canopy of rainforest trees; larvae of this genus are found under the bark of these trees, however they must also roam on the surface, as they have been collected by insecticidal fogging techniques in the very early morning before fi rst light. Members of A. cruciaria occur at midland altitudes in the Mata Atlântica. Adults are active in May, the rainy season.
Geographic distribution. (Fig. 10). Th is species is currently known from the type locality and an unknown location in the State of Minas Gerais.
Notes. Right antenna glued to card of holotype not of this specimen. Derivation of specifi c epithet. Th e epithet "grace" is an eponym, based on the given name of the Peruvian Ornithologist, Grace Servat, who has shared the birdinfested Amazon and Andes with me for many years, including two of the known localities of this species, including the type locality.

Agra grace
Proposed English vernacular name. Grace's Elegant Canopy Beetle. Diagnosis. With the attributes of the genus and species-group as described above and small sized for the pusilla group. Adults with blue integument; head behind eyes and prothorax with metallic refl ections. Frons laterad unicarinate and very fi nely rugose. Occiput not punctate, some fi ne rugae present. Elytra with moderately lobed apex in female, more so in male. Hind coxae of male multisetiferous.
Dispersal potential. Th ese beetles are macropterous and are probably capable of fl ight; they are swift and agile runners.
Way of life. Adults are found in the canopy of rainforest trees; known larvae of this genus (Arndt et al. 2001) are found under the bark of these trees, however they must also roam on the surface, as they have been collected by insecticidal fogging techniques in the very early morning before fi rst light. Members of A. grace occur at lowland altitudes in the Amazon Basin. Adults are active in September -October, the transition season between dry and wet seasons. Th e holotype was fogged from a medium-sized tree with lianas and suspended dead leaves; the area fogged was from 2 meters up to 15 meters in the tree. Th e Ecuadorian paratype was fogged from a mixed canopy consisting of the trees Geographic distribution. (Fig. 10). Th is species is currently known from Perú and Ecuador.
Notes. Males are smaller than females. Derivation of specifi c epithet. Th e epithet "max" is an eponym, based on the given name of Max Liebke, an early pioneer in the taxonomy of the genus Agra.

Agra max
Proposed English vernacular name. Max's Elegant Canopy Beetle. Diagnosis. With the attributes of the genus and species-group as described above and frons laterally unicarinate and rugose; occiput fi nely punctate; all elytral interneurs in apical half with double rows of cribriform punctures.
Dispersal potential. Th ese beetles are macropterous and are probably capable of fl ight; they are swift and agile runners.
Way of life. Adults of other Agra species are found in the canopy of rainforest trees; known larvae of this genus (Arndt et al. 2001) are found under the bark of these trees, however they must also roam on the surface, as they have been collected by insecticidal fogging techniques in the very early morning before fi rst light. Members of A. max occur at midland altitudes in the Mata Atlântica. Adults are active in February, the dry season.
Other specimens examined. None. Geographic distribution. (Fig. 10). Th is species is currently known only from the type locality. Derivation of specifi c epithet. Th e epithet "minasianus" is a Latinized adjective meaning "derived from, or pertaining to" Minas Gerais, a State in Brazil, and the type area.

Agra minasianus
Proposed English vernacular name. Minas Elegant Canopy Beetle. Diagnosis. With the attributes of the genus and species-group as described above and scape and legs bicolored, frons laterad, anterior to eye, unicarinate and smooth, prothorax moderately setiferous both laterally and ventrally, and elytra barely constricted at apical third, side margin slightly arcuate, apex moderately lobed medially, lobe obtusely dentate, smaller species.
Prothorax: Slightly broader at basal third, constricted near base and fl ared basally; surface densely punctate, laterally and ventrally setiferous, pronotum apparently with six lateral setae as in other species of the group, but not present on holotype. Pterothorax: Elytron markedly convex, intervals not costate, interneurs of rows of somewhat laterally ovate punctures, some in doubles, apex lobate, lobe well developed obtuse projection, apical dentation asymmetric, lateral and sutural apices slightly produced, each obtuse. Metasternum sparsely setiferous in female. Legs: Normal. Abdomen: Abdominal sterna III to VII of female moderately and bilaterally setiferous; sternum VII of female barely emarginate, corners rounded. Male genitalia: Unknown. Female ovipositor: Stylomere 2 as in A. notpusilla (Fig. 7).
Dispersal potential. Th ese beetles are macropterous and are probably capable of fl ight; they are swift and agile runners.
Way of life. Adults of other Agra species are found in the canopy of rainforest trees; known larvae of this genus (Arndt et al. 2001) are found under the bark of these trees, however they must also roam on the surface, as they have been collected by insecticidal fogging techniques in the very early morning before fi rst light. Th e single specimen of A. minasianus has no associated data on its labels.
Other specimens examined. None. Geographic distribution. (Fig. 10). Th is species is currently known from Minas Gerais, Brazil. Derivation of specifi c epithet. Th e epithet "pusilla" is a Latin adjective meaning very little, small, pretty. Although this species resembles A. pusilla it is "not" that species.

Agra notpusilla
Proposed English vernacular name. Brazilian Elegant Canopy Beetle. Diagnosis. With the attributes of the genus and species-group as described above and frons laterally multicarinate; elytra with interneurs composed of double rows (in part) of coarse irregularly shaped punctures, apex moderately lobed medially.
Way of life. Adults of other Agra species are found in the canopy of rainforest trees; known larvae of this genus (Arndt et al. 2001) are found under the bark of these trees, however they must also roam on the surface, as they have been collected by insecticidal fogging techniques in the very early morning before fi rst light. Members of A. notpusilla have no recorded information.
Other specimens examined. None. Geographic distribution. (Fig. 10). Th is species is currently known from Brazil. Diagnosis. With the attributes of the genus and species-group as described above and frons laterally multicarinate; occiput coarsely punctate; all elytral interneurs throughout with double rows of cribriform punctures.
Dispersal potential. Th ese beetles are macropterous and are probably capable of fl ight; they are swift and agile runners.
Way of life. Adults of other Agra species are found in the canopy of rainforest trees; known larvae of this genus (Arndt et al. 2001) are found under the bark of these trees, however they must also roam on the surface, as they have been collected by insecticidal fogging techniques in the very early morning before fi rst light. Members of A. perforata occur at midland altitudes in the Mata Atlântica.
Other specimens examined. None. Geographic distribution. (Fig. 10). Th is species is currently known only from the type locality.
Notes. Additional character state information can be found in Liebke, 1938:60. Derivation of specifi c epithet. Th e epithet "pseudopusilla" refers to the similarity between adults of this species and those of A. pusilla, treated below.

Agra pseudopusilla
Proposed English vernacular name. Mniszech's Elegant Canopy Beetle. Diagnosis. With the attributes of the genus and species-group as described above and frons laterally multicarinate; occiput coarsely bi-punctate, with several smaller punctures; elytral interneurs with mostly uni-serial rows of cribriform punctures, doubled apico-laterally.
Way of life. Adults of other Agra species are found in the canopy of rainforest trees; known larvae of this genus (Arndt et al. 2001) are found under the bark of these trees, however they must also roam on the surface, as they have been collected by insecticidal fogging techniques in the very early morning before fi rst light. Members of A. pseudopusilla are labeled Brazil without further information.
Geographic distribution. (Fig. 10). Th is species is currently known only from Brazil, without specifi c location.
Proposed English vernacular name. Small Elegant Canopy Beetle. Diagnosis. With the attributes of the genus and species-group as described above and frons laterally unicarinate and rugose; elytra with interneurs composed of a single row of coarse irregularly shaped punctures, apex markedly lobed medially.
Dispersal potential. Th ese beetles are macropterous and are probably capable of fl ight; they are swift and agile runners.
Way of life. Adults of other Agra species are found in the canopy of rainforest trees; known larvae of this genus (Arndt et al. 2001) are found under the bark of these trees, however they must also roam on the surface, as they have been collected by insecticidal fogging techniques in the very early morning before fi rst light. Members of A. pusilla occur at lowland altitudes in the Mata Atlântica. Adults are active in October, the rainy season.
Geographic distribution. (Fig. 10). Th is species is currently known from eastern Brazil.
Notes. Because of the severe deforestation over the last 100 years in the area where this species was found, it is likely it is now extinct or at least with a much smaller range.

Th e piranha species-group
Members of this group are of very small size for the genus and range in color from midnight metallic blue to smoky-black with a somewhat matte luster. Male adults have a much reduced version of the expansive ventral tarsomere pads found in all other species in the genus. Femur unicolored. Occiput and prothorax markedly punctuate. Elytral interneurs of uniseriate rows of cribriform punctures. Aedeagus with typical arrowhead shape. Female stylomere 2 short and arcuate, setiferous, and armed with two ensiform setae. Female stylomere as in Fig. 9.
Notes: Th e known composite range of the piranha group extends from Amazonian Ecuador to Bolivia across into south-central Brazil (Goiás).

Included Species
Agra ce Erwin, sp. n  Derivation of specifi c epithet. Th e epithet "ce" is a combination of pronounceable letters that when joined with the last three letters of the genus name, Agra, spells "grace," for the Peruvian Ornithologist, Grace Servat, who has shared the lowland Amazon and the high Andes with me for many years, including the known localities of this species.
Proposed English vernacular name. Graceful Elegant Canopy Beetle. Diagnosis. With the attributes of the genus and species-group as described above and elytra and prothorax metallic blue, legs unicolored, frons laterad slightly rugose, prothorax markedly setiferous both laterally and ventrally, and elytra barely constricted at apical third, side margin slightly arcuate.
Dispersal potential. Th ese beetles are macropterous and are capable of fl ight; they are swift and agile runners.
Way of life. Adults of other Agra species are found in the canopy of rainforest trees; larvae of this genus are found under the bark of the these trees, however they must also roam on the surface, as they have been collected by insecticidal fogging techniques in the very early morning before fi rst light. Members of A. ce occur at lowland altitudes in the Amazon Basin. Adults are active in September, the late dry season. Th e holotype was collected in an Erwin Plot at the type locality; the forest of this plot is designated as a Swamp Forest with internal drainage (Erwin 1985) and is dominated by the palm Mauritia fl exuosa L. and the hardwood tree Lueheopsis hoehnei Burret. Th e holotype was fogged from the later named species. Th e paratype was attracted to MV light.
Geographic distribution. (Fig. 11). Th is species is currently known only from two localities in southeastern Perú. Derivation of specifi c epithet. Th e epithet "maia" is a Latinized genitive eponym, based on the given name of Maia Samuel, Executive Producer of the Smithsonian Spotlight program, Th e Bug House, in recognition of the hours of dedication she put into developing the program.

Agra maia
Proposed English vernacular name. Maia's Elegant Canopy Beetle. Diagnosis. With the attributes of the genus and species-group as described above and prothorax brassy black, legs unicolored, frons laterad unicarinate, smooth, prothorax markedly setiferous both laterally and ventrally, and elytra markedly constricted at apical third, side margin markedly arcuate.
Dispersal potential. Th ese beetles are macropterous and are probably capable of fl ight; they are swift and agile runners.
Way of life. Adults of other Agra species are found in the canopy of rainforest trees; known larvae of this genus (Arndt et al. 2001) are found under the bark of these trees, however they must also roam on the surface, as they have been collected by insecticidal fogging techniques in the very early morning before fi rst light. Members of A. maia occur at lowland altitudes in the Amazon Basin. Adults are active in November, the rainy season.
Other specimens examined. None. Geographic distribution. (Fig. 11). Th is species is currently known only from the type locality.
Derivation of specifi c epithet. Th e epithet "piranha" or piraña, is a translation of the Huaorani word, Onkone Gare, the name of the camp near which the holotype was discovered.
Proposed English vernacular name. Piraña Elegant Canopy Beetle. Diagnosis. With the attributes of the genus and species-group as described above and brassy pronotum, legs unicolored, frons laterad unicarinate, smooth, prothorax not setiferous laterally, and elytra barely constricted at apical third, side margin slightly arcuate, apex truncate, barely lobed medially.
Dispersal potential. Th ese beetles are macropterous and are probably capable of fl ight; they are swift and agile runners.
Way of life. Adults are found in the canopy of terre fi rme rainforest trees; known larvae of this genus (Arndt et al. 2001) are found under the bark of these trees, however they must also roam on the surface, as they have been collected by insecticidal fogging techniques in the very early morning before fi rst light. Members of A. piranha occur at lowland altitudes in the Amazon Basin. Adults are active in July and October, in both the rainy and transition seasons. Th e holotype was fogged from the hardwood Eschweilera cf. laevicarpa in the family Lecythidaceae. Th e paratype was fogged from a mixed canopy consisting of the palms Iriartea deltoidea Ruiz. & Pav. and Wettinia maynensis Spruce, and the hardwood Macrolobium cf. ischnocalx, plus an unidentifi ed species of Apocynaceae.
Geographic distribution. (Fig. 11). Th is species is currently known from the Ecuadorian Amazon Basin.  Derivation of specifi c epithet. Th e epithet "risseri" is a Latinized genitive eponym, based on the surname of Dr. Paul G. Risser, outgoing Chairman of the National Board of the Smithsonian's National Museum of Natural History, in honoring his long and invaluable service to the Smithsonian Institution.

Diagnosis.
With the attributes of the genus and species-group as described above and legs unicolored, head very broad across occiput, wider than pronotum at its widest, frons laterad unicarinate, smooth, prothorax markedly setiferous both laterally and ventrally, and elytra slightly constricted at apical third, side margin barely arcuate.
Dispersal potential. Th ese beetles are macropterous and are probably capable of fl ight; they are swift and agile runners.
Way of life. Adults of other Agra species are found in the canopy of rainforest trees; known larvae of this genus (Arndt et al. 2001) are found under the bark of these trees, however they must also roam on the surface, as they have been collected by insecticidal fogging techniques in the very early morning before fi rst light. Members of A. risseri occur at lowland to midland altitudes in the Amazon Basin. Adults are active in October, the rainy season.
Geographic distribution. (Fig. 11). Th is species is currently known from Bolivia and Brazil. Derivation of specifi c epithet. Th e epithet "tiputini" is the name of the Research Station and the river near which the holotype was collected.

Agra tiputini
Proposed English vernacular name. Tiputini Elegant Canopy Beetle. Diagnosis. With the attributes of the genus and species-group as described above and pronotum brassy, legs unicolorous, lateral depression of frons unicarinate, smooth, gena and occiput with sparse and moderately small setigerous punctures plus two larger ones; elytra constricted at basal third, fl ared at apical third, side markedly arcuate, intervals not costate. Description. Size: Small, ABL = 5.98 -7.17 mm, SBL = 5.98 -6.04 mm, TW = 2.04 mm. Color: Head black with bluish refl ection posteriorly, body and legs with metallic blue refl ections, elytra metallic cobalt blue; antennae and mouthparts piceous, scape with slight metallic blue refl ections. Luster: Shiny metallic, elytra matte. Head: Labrum moderately elongate and rounded at corners, slightly emarginate medially. Frons medially raised and smooth, laterally depressed and smooth. Gena slightly tapered, hind angle obtuse to constricted neck in both female. Genae and occiput with sparse setiferous punctures, some coarsely so. Prothorax: Slightly broader medially, slightly fl ared basally; surface with dense punctures, disc each side with four long setae, with short setae both basally and apically; lateral elongate callous with single row of non-setiferous puncture along middle. Pterothorax: Elytron moderately convex, fl ared at apical third, intervals not costate, interneurs of rows of somewhat laterally ovate cribriform punctures, apex trun- Figure 10. Distribution map of the species of the pusilla group. Names marked with a "?" do not have precise localities on the specimen label(s). cate, barely lobate, apical dentation asymmetric, lateral tooth small, acute, sutural apex not produced, rounded. Metasternum sparsely setiferous in females. Legs: Legs normal. Abdomen: Abdominal sterna III, IV, and V of females sparsely setiferous bilaterally; sternum VII of female very slightly emarginate. Male genitalia: Unknown. Female ovipositor: Stylomere 2 as in A. piranha (Fig. 9).
Dispersal potential. Th ese beetles are macropterous and are probably capable of fl ight; they are swift and agile runners.
Way of life. Adults are found in the canopy of terre fi rme rainforest trees; known larvae of this genus (Arndt et al. 2001) are found under the bark of these trees, however they must also roam on the surface, as they have been collected by insecticidal fogging techniques in the very early morning before fi rst light. Members of A. tiputini occur at lowland altitudes in the Amazon Basin. Adults are active in February and October, Figure 11. Distribution map of the species of the piranha group. the dry season and the transition season. Th e holotype was fogged from a mixed canopy consisting of the crowns of the palms Iriartea deltoidea Ruiz. & Pav. and Wettinia maynensis Spruce, and the hardwood Macrolobium cf. ischnocalyx. Th e paratype was fogged from the hardwood family Myrtaceae. Th e February specimen is very teneral suggesting that the dry season triggers pupation and emergence.
Geographic distribution. (Fig. 11). Th is species is currently known from Amazonian Ecuador.

Discussion
Th e relatively narrow tarsi, spatulate apex of the median lobe of the male genitalia, and very small size of its adults suggest that the pusilla species group represents the most basal lineage in the evolution of the hyper diverse genus Agra, given that more highly derived states of tarsal width, male genitalic form, and size are found in the platyscelis, famula, and formicaria species groups, as well as the piranhna group, treated herein? In the "Introduction" above, I suggested that fresh specimens should be collected to provide DNA that can be sequenced and analyzed concomitant with those of other adult Agrina genera from Africa to determine the adelphotaxon. Subsequent to such an analysis, a better understanding of the evolution of structural attributes among Agra lineages and species will be possible than it is at present.