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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.
Neotropics, Bolivia, Brazil, Ecuador, Perú, rainforest, canopy
The 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 (
These 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.
The 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 identified. The 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 identification and classification.
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
(
Methods and species concepts follow those previously described (
Included in this study are a total of 22 specimens from the National Museum of Natural History, Washington, DC (NMNH); H. Perrin, J. Menier, Museu National d’Historie Naturelle, Paris (MNHNP ); A. Kuska, Institut Zoologique, Warsaw, Poland (WAR); S. Fragoso, M. Monné, Brazil National Collection, Museu Nacional, Rio de Janeiro, Brazil (BNCRio); M. Baccis, British Museum (Natural History), London, England (BMNH); O.V. Ferreira and J. Jurberg, Ozwaldo Cruz Institute, Rio de Janeiro, Brazil, (ZIKAN); R. Woodruff, Florida Department of Agriculture, Gainesville (FSCA). Note that the curators named were on duty at the time of the loan; many of these have since retired.
The 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. The habitus images of the adults were made with a Visionary DigitalTM high resolution imaging system. Figure captions include an ADP number, which is a unique identification 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.
The species list below, as well as arrangement of descriptions that follows is ordered alphabetically.
Elegant Canopy Beetles
During evolution toward a canopy domain and away from a likely under canopy sister group, Agra
adults acquired numerous generic-level autapotypic features as follows:
head elongate with prognathate mandibles, securiform labial ultimate
palpomeres (
For descriptions of species groups previously recognized, refer to
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: The known composite range of the pusilla group extends from Amazonian Ecuador and Perú across southeastern Brazil and into the Mata Atlântica.
Agra cruciaria Erwin, sp. n. Brazil
Agra grace Erwin, sp. n. Ecuador, Perú
Agra max Erwin, sp. n. Brazil
Agra minasianus Erwin, sp. n. Brazil
Agra notpusilla Erwin, sp. n. Brazil
Agra perforata Liebke, 1938 Brazil
Agra pseudopusilla Erwin, sp. n. Brazil
Agra pusilla Chaudoir, 1847 Brazil
1 | Elytra bright metallic blue | 2 |
1’ | Elytra dull matte smoky black or smoky black-blue, not bright | 3 |
2(1) | Elytral interneurs each with a single row of cribriform punctures | Agra grace, sp. n. |
2’ | Elytral interneurs each with double rows of cribriform punctures at least in part in apical half | Agra max, sp. n. |
3(1’) | Antennal scape unicolorous testaceous, or testaceous with slight dorso-apical infuscation | 4 |
3’ | Antennal scape bicolored, ventrally testaceous, dorsally infuscated | 6 |
4(3) | Head posteriorly and pronotum with surface brassy | Agra cruciaria, sp. n. |
4’ | Head and pronotum matte smoky black , shiny or not | 5 |
5(4’) | Elytra constricted at apical third, side margin moderately arcuate, apex moderately lobed medially | Agra notpusilla, sp. n. |
5’ | Elytra not constricted at apical third, side margin straight, apex markedly lobed medially | Agra pusilla Chaudoir |
6(3’) | Elytral apex shallowly lobed medially, lobe rounded | Agra pseudopusilla, sp. n. |
6’ | Elytral apex moderately lobed medially, lobe obtusely dentate | 7 |
7(6’) | Frons laterad, anterior to eye markedly rugose; larger species (ABL = 8.2 mm) | Agra perforata Liebke |
7’ | Frons laterad, anterior to eye, unicarinate and smooth, smaller species (ABL = 7.2 mm) | Agra minasianus, sp. n. |
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Figs 3, 10Brazil: Rio de Janeiro, Rio de Janeiro, Corcovado, 585m, 22.9517°S, 43.2116°W, 5 May 1958 (C.A. Campos Seabra & M. Alvarenga)(NMNH: ADP 070044, male).
The epithet “cruciaria" is a Latin adjective meaning “of/pertaining to the cross/torture" and is based upon the large cross on Corcovado flooded each night with high powered lights that attract insects by the millions and is the type locality of this species.
Cross Elegant Canopy Beetle.
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 reflections. Frons laterad unicarinate and rugose. Occiput with sparse punctures, some punctures with short setae.
Elytra with moderately lobed apex.
(Fig. 3). Size: Small, ABL = 6.4 – 7.7 mm, SBL = 5.75 – 6.48 mm, TW = 1.74 –1.8 mm. Color: Head black with brassy reflection posteriorly, pronotum with brassy reflections; antennae blackish blue with scape testaceous and with slight metallic blue reflections , mouthparts piceous, and legs and tarsi blackish blue, with mostly testaceous femur. Luster: Head, pronotum and legs shiny metallic, elytra matte smoky-blue. Head: Labrum moderately elongate and truncate apically, barely emarginate medially. Frons medially raised and smooth, laterally depressed and unicarinate and rugose. Gena with hind angles broadly rounded to constricted neck in males. Genae and occiput moderately densely and coarsely some punctures setiferous.
Prothorax: Slightly broader medially, flared 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.
These beetles are macropterous and are probably capable of flight; they are swift and agile runners.
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 first light. Members of Agra cruciaria occur at midland altitudes in the Mata Atlântica. Adults are active in May, the rainy season.
Brazil: Minas Gerais, (MNHNP: ADP 060040, male paratype).
(Fig. 10). This species is currently known from the type locality and an unknown location in the State of Minas Gerais.
Right antenna glued to card of holotype not of this specimen.
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Figs 1, 4, 10Perú: Madre de Dios, Pakitza, Tachigali Trail /47, 324m, “11.9352°S, 71.3039°W, " 6 October 1991 (T.L. Erwin & M.G. Pogue)(NMNH: BIOLAT 012952, female).
The epithet “grace" is an eponym, based on the given name of the Peruvian Ornithologist, Grace Servat, who has shared the bird-infested Amazon and Andes with me for many years, including two of the known localities of this species, including the type locality.
Grace’s Elegant Canopy Beetle.
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 reflections. Frons laterad unicarinate and very finely rugose. Occiput not punctate, some fine rugae present. Elytra with moderately lobed apex in female, more so in male. Hind coxae of male multisetiferous.
Agra grace Erwin, sp. n., dorsal aspect (BIOLAT 012952).
(Figs 1, 4). Size: Very small, ABL = 5.71 – 6.1 mm, SBL = 4.77 – 6.67 mm, TW = 1.44 – 1.66 mm. Color: Head and pronotum bright blue, body metallic blue; antennae and mouthparts piceous, scape and antennomeres 2 and 3 with slight metallic blue reflections. Luster: Shiny metallic, elytra somewhat matte. Head: (Fig. 1) Labrum moderately elongate and truncate apically. Frons medially raised and smooth, laterally depressed, unicarinate, not rugose. Gena slightly tapered with broadly rounded corners to constricted neck in both sexes. Occiput not punctate, some fine rugae present. Prothorax: Slightly broader medially, flared basally; surface with dense coarse punctures, some setiferous; lateral elongate callous with single row of punctures, along middle. Pterothorax: Elytron (Fig. 1) markedly convex, intervals slightly costate, interneurs of rows of somewhat laterally ovate punctures, apex truncate, barely oblique, apical dentation asymmetric, lateral tooth broad, obtuse, sutural apex not produced. Metasternum sparsely setiferous in both sexes. Legs: Normal. Abdomen: Abdominal sterna III to VII of both sexes moderately and bilaterally setiferous; sternum VII of both sexes barely emarginate, corners rounded. Male genitalia: Phallus (Fig. 4) elongate and narrow with ostium not elongate, extended to about 1/3 the length of phallus, apex a small rounded lobe. Parameres small, left twice the size of the right, both broadly rounded. Female ovipositor: Stylomere 2 as in Agra notpusilla (Fig. 7).
These beetles are macropterous and are probably capable of flight; they are swift and agile runners.
Adults are found in the canopy of rainforest trees; known larvae of this genus (
Ecuador: Orellana, 1 km S Onkone Gare Camp, Entomology Transect, 216m, “0.6569°S, 76.4527°W, " 7 October 1994 (T.L. Erwin, et al.)(NMNH: ADP 087438, male paratype). Perú: Madre de Dios, Pakitza, 324m, “11.9352°S, 71.3039°W, " 9 September 1988 (T.L. Erwin & B.D. Farrell)(NMNH: BIOLAT 008430, male paratype).
(Fig. 10). This species is currently known from Perú and Ecuador.
Males are smaller than females.
Brazil: Santa Catarina, Nova Teutonia, 823m, 27.047°S, 52.394°W, 4 February 1938 (F. Plaumann)(WAR: ADP 004374, female).
The epithet “max" is an eponym, based on the given name of Max Liebke, an early pioneer in the taxonomy of the genus Agra.
Max’s Elegant Canopy Beetle.
With the attributes of the genus and species-group as described above and frons laterally unicarinate and rugose; occiput finely punctate; all elytral interneurs in apical half with double rows of cribriform punctures.
Size: Small, ABL = 7.0 mm, SBL = 5.95 mm, TW = 1.5 mm. Color: Head and pronotum black with bluish reflections, venter metallic blue, elytra matte blue, antennae and mouthparts piceous with slightly bluish reflections, scape rufous with shiny infuscated apex. Luster: Shiny metallic, elytra matte metallic. Head: Labrum moderately elongate and truncate apically. Frons medially raised and smooth, laterally depressed, slightly rugose. Gena slightly markedly rounded to constricted neck in female. Genae and occiput with sparse coarse punctures, some setiferous.
Prothorax: Short, slightly broader medially, flared basally; surface with dense and coarse punctures, some setiferous; lateral elongate callous with single row of setiferous puncture along middle. Pterothorax: Elytron markedly convex, intervals slightly costate, interneurs of rows of somewhat laterally ovate punctures, doubled in some places, apex moderately oblique and lobed, apical dentation asymmetric, lateral tooth small, broad, obtuse, sutural apex not produced. Metasternum sparsely setiferous in females. Legs: Normal in female. 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 Agra notpusilla (Fig. 7).
These beetles are macropterous and are probably capable of flight; they are swift and agile runners.
Adults of other Agra species are found in the canopy of rainforest trees; known larvae of this genus (
None.
(Fig. 10). This species is currently known only from the type locality.
Brazil: Minas Gerais, (Laferte)(MNHNP: ADP 060090, female).
The epithet “minasianus" is a Latinized adjective meaning “derived from, or pertaining to" Minas Gerais, a State in Brazil, and the type area.
Minas Elegant Canopy Beetle.
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.
Size: Very small, ABL = 7.2 mm, SBL = 6.24 mm, TW = 1.5 mm. Color: Head and prothorax black, legs and scape bicolored; flagellar antennomeres and mouthparts piceous. Luster: Matte. Head: Labrum moderately elongate and truncate apically, anterior margin slightly emarginate medially. Frons laterad, anterior to eye, unicarinate and smooth. Gena rounded to constricted neck in females. Genae and occiput with sparse coarse punctures, some setiferous.
Prothorax: Slightly broader at basal third, constricted near base and flared 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 Agra notpusilla (Fig. 7).
These beetles are macropterous and are probably capable of flight; they are swift and agile runners.
Adults of other Agra species are found in the canopy of rainforest trees; known larvae of this genus (
None.
(Fig. 10). This species is currently known from Minas Gerais, Brazil.
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Figs 7, 10Brazil: (MNHNP: ADP 058647, male).
The epithet “pusilla" is a Latin adjective meaning very little, small, pretty. Although this species resembles Agra pusilla it is “not" that species.
Brazilian Elegant Canopy Beetle.
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.
(Fig. 7). Size: Very small, ABL = 7.74 mm, SBL = 6.58 mm, TW = 1.88 mm. Color: Piceous, antennae and legs bicolored, scape and basal 5/6th of femur testaceous; antennal flagellum and mouthparts piceous. Luster: Head and abdomen shiny, elytral disc somewhat matte. Head: Labrum moderately elongate and truncate apically, slightly emarginate medially. Frons medially raised and smooth, laterally depressed, multicarinate. Gena rounded in female to constricted neck. Genae and with occiput sparse and moderately coarse punctures, some of which likely setiferous in undamaged specimens (no setae present in holotype), and with two larger punctures, . Prothorax: Slightly broader in basal third, constricted, and flared basally; surface densely punctate, laterally and ventrally setiferous. Pterothorax: Elytron markedly convex, intervals not costate, interneurs of double rows (in part) of somewhat irregularly shaped punctures, apex truncate, moderately lobate, apical dentation asymmetric, lateral tooth small, acute, sutural apex slightly produced, rounded. Metasternum sparsely setiferous in both females. Legs: Legs normal. Abdomen: Abdominal sterna III to VII of female moderately and bilaterally setiferous; sternum VII of females barely emarginate, corners rounded. Male genitalia: Unknown. Female ovipositor: Stylomere 2 (Fig. 7).
Adults of other Agra species are found in the canopy of rainforest trees; known larvae of this genus (
None.
(Fig. 10). This species is currently known from Brazil.
Brazil: Rio de Janeiro, Rio de Janeiro, 585m, 22.9517°S, 43.2116°W (WAR: ADP 060035, female).
The epithet “perforata" is a Latin adjective referring to the cribriform punctures of the elytral interneurs.
Perforated Elegant Canopy Beetle.
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.
Size: Small, ABL = 8.22 mm, SBL = 6.58 mm, TW = 1.88 mm. Color: All black with mostly testaceous femur. Luster: Head slightly shiny, pronotum and elytra matte black. Head: Labrum moderately elongate and rounded apically. Frons medially raised and smooth, laterally depressed, moderately rugose. Gena markedly rounded to constricted neck in female. Genae and occiput with sparse and coarse punctures, some setiferous. Prothorax: Short, slightly broader medially, flared basally; surface with dense and coarse punctures, some setiferous; lateral elongate callous with single row of setiferous punctures along middle. Pterothorax: Elytron markedly convex, intervals slightly costate, interneurs of double rows of somewhat irregularly-shaped punctures, apex moderately oblique and lobed, apical dentation asymmetric, lateral tooth small, broad, obtuse, sutural apex not produced. Metasternum sparsely setiferous in females. Legs: Normal in female. 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 Agra notpusilla (Fig. 7).
These beetles are macropterous and are probably capable of flight; they are swift and agile runners.
Adults of other Agra species are found in the canopy of rainforest trees; known larvae of this genus (
None.
(Fig. 10). This species is currently known only from the type locality.
Additional character state information can be found in
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Figs 5, 10Brazil: (Comte G. de Mniszech)(MNHNP: ADP 060088, female).
The epithet “pseudopusilla" refers to the similarity between adults of this species and those of Agra pusilla, treated below.
Mniszech’s Elegant Canopy Beetle.
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.
(Fig. 5). Size: Small, ABL = 6.57 – 8.63 mm, SBL = 5.67 – 7.15 mm, TW = 1.34 – 2.08 mm. Color: Head and pronotum black, elytra smoky black, legs bicolored, antennae and mouthparts piceous, scape with testaceous venter, piceous dorsum . Luster: Shiny forebody, matte elytra. Head: Labrum moderately elongate and truncate apically, anterior corners rounded. Frons medially raised and smooth, laterally depressed, multicarinate. Gena slightly tapered with broadly rounded corners to constricted neck in both male. Occiput coarsely bi-punctate, with several smaller punctures. Prothorax: Slightly broader medially, flared basally; surface with dense and coarse punctures, some setiferous; lateral elongate callous with single row of setigerous punctures along middle. Pterothorax: Elytron markedly convex, intervals slightly costate, interneurs of rows of somewhat laterally ovate punctures, doubled in some places, apex oblique, slightly lobed at middle, apical dentation asymmetric, lateral tooth short, acute, sutural apex not produced. Metasternum sparsely setiferous in males. Legs: Normal. Abdomen: Abdominal sterna III to VII of male moderately and bilaterally setiferous; sternum VII of males barely emarginate, corners rounded. Male genitalia: Phallus (Fig. 5) elongate and narrow with ostium not elongate, extended to about 1/2 the length of phallus, apex a small rounded lobe. Parameres small, left twice the size of the right, both moderately rounded. Female ovipositor: Female unknown
These beetles are macropterous and are probably capable of flight; they are swift and agile runners.
Adults of other Agra species are found in the canopy of rainforest trees; known larvae of this genus (
Brazil: (Ehrenreiche)(BNCRio: ADP 070045, male paratype).
(Fig. 10). This species is currently known only from Brazil, without specific location.
Brazil: (MNHNP: ADP 060087, male).
The epithet “pusilla" is a Latin adjective meaning very little, small, pretty.
Small Elegant Canopy Beetle.
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.
(Fig. 6). Size: Very small, ABL = 6.28 – 6.45 mm, SBL = 5.27 – 5.37 mm, TW = 1.32 – 1.46 mm. Color: Piceous, antennae and legs bicolored, scape and basal 5/6th of femur testaceous; antennal flagellum and mouthparts piceous. Luster: Shiny, elytral disc somewhat matte. Head: Labrum moderately elongate and rounded apically, slightly emarginate medially. Frons medially raised and smooth, laterally depressed, unicarinate, and shallowly rugose. Gena slightly tapered-rounded in both sexes to constricted neck. Genae and occiput moderately finely punctate and wrinkled, and with two larger punctures each of which is setiferous. Prothorax: Slightly broader medially, flared basally; surface with densely punctures, some setiferous; lateral elongate callous with single row of setiferous punctures along middle. Pterothorax: Elytron markedly convex, intervals not costate, interneurs of rows of single somewhat irregularly shaped punctures, apex truncate, markedly lobate, apical dentation asymmetric, lateral tooth small, acute, sutural apex slightly produced, rounded. Metasternum sparsely setiferous in both sexes. Legs: Legs normal. Abdomen: Abdominal sterna III to VII of male moderately and bilaterally setiferous; sternum VII of males barely emarginate, corners rounded. Male genitalia: Phallus (Fig. 6) with ostium elongate, about half the length of the phallus, apex a small rounded lobe with basal corners. Parameres small, left twice the size of the right, both moderately rounded. Female ovipositor: Stylomere 2 as in Agra notpusilla (Fig. 7).
These beetles are macropterous and are probably capable of flight; they are swift and agile runners.
Adults of other Agra species are found in the canopy of rainforest trees; known larvae of this genus (
Brazil: Espirito Santo, nr. Itapemirim, Rio Itapemirim, 3m, 21.005°S, 40.834°W, 15 October 1906 (J.F. Zikan)(ZIKAN: ADP 070043, male).
(Fig. 10). This species is currently known from eastern Brazil.
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.
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: The known composite range of the piranha group extends from Amazonian Ecuador to Bolivia across into south-central Brazil (Goiás).
Included SpeciesAgra ce Erwin, sp. n. Perú
Agra risseri Erwin, sp. n. Bolivia, Brazil
Agra maia Erwin, sp. n. Bolivia
Agra piranha Erwin, sp. n. Ecuador
Agra tiputini Erwin, sp. n. Ecuador
1 | Prothorax markedly setiferous both laterally and ventrally, pronotum with 3 long setae on each side | 2 |
1’ | Prothorax not setiferous laterally, pronotum with 4 long setae on each side | 4 |
2(1) | Head very broad across occiput, wider than pronotum at its widest | Agra risseri, sp. n. |
2’ | Head not broad across occiput, narrower than pronotum at its widest | 3 |
3(2’) | Elytra constricted at apical third, side margin markedly arcuate | Agra maia, sp. n. |
3’ | Elytra barely constricted at apical third, side margin slightly arcuate | Agra ce, sp. n. |
4(1’) | Elytra constricted at apical third, side margin markedly arcuate; head markedly narrowed basally | Agra tiputini, sp. n. |
4’ | Elytra barely constricted at apical third, side margin slightly arcuate; head slightly tapered yet broad basally | Agra piranha, sp. n. |
Perú: Madre de Dios, 30 air km SW Puerto Maldonado, 205m, “12.8368°S, 69.2933°W, " 10 September 1984 (T.L. Erwin, et al.)(NMNH: ADP 093837, female).
The 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.
Graceful Elegant Canopy Beetle.
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.
Size: Small, ABL = 8.04 – 9.21 mm, SBL = 7.09 – 8.07 mm, TW = 2.36 – 2.78 mm. Color: Head black with bluish reflection posteriorly, body and legs metallic blue; antennae and mouthparts piceous, scape with slight metallic blue reflections. Luster: Shiny metallic. Head: Labrum moderately elongate and rounded apically. Frons medially raised and smooth, laterally depressed and rugose. Gena almost squared to constricted neck in females. Genae and occiput moderately densely punctate, each puncture setiferous. Prothorax: Slightly broader medially, flared basally; surface with dense and coarse setiferous punctures; lateral elongate callous with single row of setiferous punctures along middle. Pterothorax: Elytron markedly convex, intervals moderately costate, interneurs of rows of somewhat laterally ovate punctures, apex truncate, barely lobate, apical dentation asymmetric, lateral tooth small, acute, sutural apex slightly produced, narrowly pointed. Metasternum sparsely setiferous in female. Legs: Normal in female. Abdomen: Abdominal sterna III to VII of female moderately and bilaterally setiferous; sternum VII of female barely emarginated, corners rounded. Male genitalia: Unknown. Female ovipositor: Stylomere 2 as in Agra piranha (Fig. 9).
These beetles are macropterous and are capable of flight; they are swift and agile runners.
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 first light. Members
of Agra ce
occur at lowland altitudes in the Amazon Basin. Adults are active in
September, the late dry season. The 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 (
Perú: Madre de Dios, Pakitza, Trocha Uno /14, 324m, “11.9352°S, 71.3039°W, " 8 September 1989 (R.A. Faitoute, et al.)(NMNH: BIOLAT 017465, female paratype).
(Fig. 11). This species is currently known only from two localities in southeastern Perú.
Bolivia: Santa Cruz, 4–6 km SSE Buena Vista, Hotel Flora & Fauna, 400–500m, “17.479°S, 63.631°W, " 1–10 November 2002 (S.W. Lingafelter)(NMNH: ADP 116043, female).
The epithet “maia" is a Latinized genitive eponym, based on the given name of Maia Samuel, Executive Producer of the Smithsonian Spotlight program, The Bug House, in recognition of the hours of dedication she put into developing the program.
Maia’s Elegant Canopy Beetle.
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.
Size: Small, ABL = 8.57 mm, SBL = 7.25 mm, TW = 2.2 mm. Color: Head black with faint bluish reflection posteriorly, pronotum brassy black legs and venter black, elytra metallic blue; antennae and mouthparts piceous. Luster: Shiny, pronotum brassy and elytra shiny metallic. Head: Labrum moderately elongate and moderately rounded apically. Frons medially raised and smooth, laterally depressed and smooth. Gena almost squared to constricted neck in females. Genae and occiput with moderately dense disc each side with four long setae, punctures, most setiferous. Prothorax: Slightly broader medially, flared basally; surface densely punctuate, disc each side with four long setae; lateral elongate callous with single row of setiferous punctures along middle. Pterothorax: Elytron moderately convex, broadly flared at apical third, intervals moderately costate, interneurs of rows of somewhat laterally ovate punctures, apex truncate, barely lobate, apical dentation asymmetric, lateral tooth small, acute, sutural apex slightly produced, narrowly pointed. Metasternum sparsely setiferous in female. Legs: Normal in female. Abdomen: Abdominal sterna III to VII of female moderately and bilaterally setiferous; sternum VII of female barely emarginated, corners rounded. Male genitalia: Unknown. Female ovipositor: Stylomere 2 as in Agra piranha (Fig. 9).
These beetles are macropterous and are probably capable of flight; they are swift and agile runners.
Adults of other Agra species are found in the canopy of rainforest trees; known larvae of this genus (
None.
(Fig. 11). This species is currently known only from the type locality.
urn:lsid:zoobank.org:act:0B2DB88C-89F6-485D-B2FB-C2B54813D0DC
Figs 8, 9, 11Ecuador: Orellana, 1 km S Onkone Gare Camp, Entomology Transect, 216m, “0.6569°S, 76.4527°W, " 2 July 1995 (T.L. Erwin, et al.)(NMNH: ADP 087440, male).
The epithet “piranha" or piraña, is a translation of the Huaorani word, Onkone Gare, the name of the camp near which the holotype was discovered.
Piraña Elegant Canopy Beetle.
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.
(Fig. 8, 9). Size: Very small, ABL = 7.18 – 7.29 mm, SBL = 5.98 – 6.13 mm, TW = 1.76 – 1.86 mm. Color: Head black with bluish reflection posteriorly, body and legs with metallic blue reflections, elytra metallic cobalt blue; antennae and mouthparts piceous, scape with slight metallic blue reflections. Luster: Shiny metallic, elytra matte. Head: Labrum moderately elongate and rounded at corners, slightly emarginate medially. Frons medially raised and smooth, laterally depressed and uni-carinate. Gena slightly tapered, hind angle obtuse to constricted neck in both sexes. Genae and occiput with moderately dense and coarse punctures, some setiferous. Prothorax: Slightly broader medially, flared basally; surface densely punctuate, disc each side with three long setae; lateral elongate callous with single row of non-setiferous punctures along middle. Pterothorax: Elytron markedly convex, intervals moderately costate, interneurs of rows of somewhat laterally ovate punctures, apex truncate, barely lobate, apical dentation asymmetric, lateral tooth small, acute, sutural apex not produced, rounded. Metasternum sparsely setiferous in both sexes. Legs: Middle tibia of male with dense brush of short setae on medial margin. Abdomen: Abdominal sterna III, IV, and V of both sexes sparsely setiferous bilaterally; sternum VII of both sexes very slightly emarginate. Male genitalia: Phallus (Fig. 8) elongate, narrow, with moderately broad arrow-shaped apex; ostium elongate. Parameres small, left twice the size of the right, both broadly rounded. Female ovipositor: Stylomere 2 (Fig. 9).
These beetles are macropterous and are probably capable of flight; they are swift and agile runners.
Adults are found in the canopy of terre firme rainforest trees; known larvae of this genus (
Ecuador: Rio Tiputini, Erwin Transect, 232m, “0.63173°S, 76.14420°W, " 23 October 1998 (T.L. Erwin, et al.)(NMNH: ADP 117227, female paratype).
(Fig. 11). This species is currently known from the Ecuadorian Amazon Basin.
Bolivia: Santa Cruz, 4–6 km SSE Buena Vista, Hotel Flora & Fauna, 383m, “17.479°S, 63.631°W, " 27–29 October 2000 (J.E. Wappes & R. Morris)(FSCA: ADP 115786, female).
The 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.
Risser’s Elegant Canopy Beetle.
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.
Agra risseri Erwin, sp. n., dorsal aspect (ADP 115786).
3 Agra cruciaria Erwin, sp. n., male genitalia (dorsal, ventral, left lateral aspects) (ADP 070044) 4 Agra grace Erwin, sp. n., male genitalia (dorsal, ventral, left lateral aspects) (ADP 087638) 5 Agra pseuopusilla Erwin, sp. n., male genitalia (dorsal, ventral, left lateral aspects) (ADP 070045) 6 Agra pusilla Chaudoir, sp. n., male genitalia (dorsal, ventral, left lateral aspects) (ADP 060087).
Agra notpusilla Erwin, sp. n., female stylomeres (dorsal aspect) (ADP 058647).
8 Agra piranha Erwin, sp. n., male genitalia (dorsal, ventral, left lateral aspects) (ADP 087440) 9 Agra piranha Erwin, sp. n., female stylomeres (dorsal aspect) (ADP 117227).
(Fig. 2). Size: Small, ABL = 9.42 – 10.45 mm, SBL = 8.14 – 8.77 mm, TW = 2.4 – 3.08 mm. Color: Head, body, and appendages black, pronotum with brassy reflection elytra metallic blue. Luster: Shiny. Head: (Fig. 2) Labrum moderately elongate and slightly rounded at anterior corners. Frons medially raised and smooth, laterally depressed and unicarinate, smooth. Gena almost squared to constricted neck in females. Genae and occiput with moderately dense punctures, most setiferous. Prothorax: Slightly broader medially, flared basally; surface with dense punctures, some setiferous; lateral elongate callous with single row of setiferous punctures along middle. Pterothorax: Elytron (Fig. 2) with discal area flat, intervals moderately costate, interneurs of rows of somewhat laterally ovate punctures, apex truncate, barely lobate, apical dentation asymmetric, lateral tooth small, acute, sutural apex slightly produced, narrowly pointed. Metasternum sparsely setiferous in female. Legs: Normal in female. Abdomen: Abdominal sterna III to VII of female moderately and bilaterally setiferous; sternum VII of female shallowly emarginate, corners rounded. Male genitalia: Unknown. Female ovipositor: Stylomere 2 as in Agra piranha (Fig. 9).
These beetles are macropterous and are probably capable of flight; they are swift and agile runners.
Adults of other Agra species are found in the canopy of rainforest trees; known larvae of this genus (
Brazil: Goiás, Jataí (Jatahy), 736m, 17.880°S, 51.720°W (BMNH: ADP 004375, female paratype).
(Fig. 11). This species is currently known from Bolivia and Brazil.
Distribution map of the species of the pusilla group. Names marked with a “?" do not have precise localities on the specimen label(s).
Distribution map of the species of the piranha group.
Ecuador: Ecuador: Orellana, Tiputini Biodiversity Station, Rio Tiputini, Erwin Transect, 232m, “0.63173°S, 76.14420°W, " 23 October 1998 (T.L. Erwin, et al.)(NMNH: 117233, female).
The epithet “tiputini" is the name of the Research Station and the river near which the holotype was collected.
Tiputini Elegant Canopy Beetle.
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, flared at apical third, side markedly arcuate, intervals not costate.
Size: Small, ABL = 5.98 – 7.17 mm, SBL = 5.98 – 6.04 mm, TW = 2.04 mm. Color: Head black with bluish reflection posteriorly, body and legs with metallic blue reflections, elytra metallic cobalt blue; antennae and mouthparts piceous, scape with slight metallic blue reflections. 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 flared 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, flared at apical third, intervals not costate, interneurs of rows of somewhat laterally ovate cribriform punctures, apex truncate, 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 Agra piranha (Fig. 9).
These beetles are macropterous and are probably capable of flight; they are swift and agile runners.
Adults are found in the canopy of terre firme rainforest trees; known larvae of this genus (
Orellana, 1 km S Onkone Gare Camp, Entomology Transect, 216m, “0.6569°S, 76.4527°W, " 8 February 1995 (T.L. Erwin, et al.)(NMNH: ADP 087439, male paratype).
(Fig. 11). This species is currently known from Amazonian Ecuador.
The 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.
In addition to the Museum curators and technicians noted above, I also extend a hearty thanks to Charyn Micheli (the Resumen, measures, and friendly review of the manuscript), Warren Steiner (dissections), Karie Darrow (images and plates), and Vichai Malikul (genitalic illustrations) all of the Department of Entomology at the Smithsonian Institution.