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Zoologia (Curitiba)

Print version ISSN 1984-4670

Zoologia (Curitiba) vol.29 no.4 Curitiba Aug. 2012

https://doi.org/10.1590/S1984-46702012000400008 

TAXONOMY AND NOMENCLATURE

 

Species of Euglossa of the analis group in the Atlantic forest (Hymenoptera, Apidae)

 

 

Luiz R. R. FariaI,II; Gabriel A. R. MeloII,III

IDepartamento de Ciências Agrárias e Biológicas, Centro Universitário Norte do Espírito Santo, Universidade Federal do Espírito Santo. Rodovia BR-101 Norte, km 60, Litorâneo, 29932-540 São Mateus, ES, Brazil, E-mail: nunofariajr@gmail.com
IILaboratório de Biologia Comparada Hymenoptera, Departamento de Zoologia, Universidade Federal do Paraná. Caixa Postal 19020, 81531-980 Curitiba, Paraná, Brazil
IIICorresponding author, E-mail: garmelo@ufpr.br

 

 


ABSTRACT

The species of Euglossa Latreille, 1802 of the analis group inhabiting the Brazilian Atlantic forest are revised and identification keys for males and females are provided. Five species are recognized in the Atlantic forest: Euglossa cognata Moure, 1970, Euglossa marianae Nemésio, 2011, Euglossa roderici Nemésio, 2009 and two new species described here, Euglossa botocuda sp. nov. and Euglossa calycina sp. nov. These two new species have been misidentified by previous authors as Atlantic forest populations of, respectively, Euglossa iopyrrha Dressler, 1982 and Euglossa mixta Friese, 1899. Relevant morphological features are illustrated and distribution maps are also provided. Notes on the analis group are included and an additional available name, Euglossa aureiventris Friese, 1899, is placed in this species group.

Key words: Apinae; Euglossina; Euglossini; orchid bee; taxonomy.


 

 

Orchid bees (subtribe Euglossina) are a primarily Neotropical group of bees, occurring from the southern United States (Minckley & Reyes 1996, Skov & Wiley 2005) to the southern Brazil (Wittmann et al. 1988) and northern Argentina (Pearson & Dressler 1985), and constitute a key group in forest environments of the Neotropical region (Dodson et al. 1969). The development of a specific technique to attract males, based on the attractiveness of synthetic compounds analogous to substances collected by them in the environment (Dodson et al. 1969), advanced to a high level the knowledge of their biology, diversity and species distribution (Roubik & Hanson 2004).

The subtribe comprises approximately 200 species (Moure et al. 2007) described in five monophyletic genera (Michel-Salzat et al. 2004, Ramírez et al. 2010). Euglossa Latreille, 1802 is the most speciose genus, comprising, approximately, 100 species, divided into six subgenera (Moure et al. 2007). The taxonomy of this genus was subject of recent studies, dealing with the description of new species (e.g., Ramírez 2006, Hinojosa-Díaz & Engel 2007, Nemésio 2007, 2009), revalidation of synonymized names (e.g., Faria & Melo 2007, Nemésio 2009) and also the production of species checklists and catalog (Moure et al. 2007, Nemésio 2009, Nemésio & Rasmussen 2011).

The diversity of orchid bees found in the Atlantic forest is lower than in the Amazon forest and Central America (Nemésio & Silveira 2007). Regarding Euglossa, Moure et al. (2007) recognized 30 species inhabiting the Atlantic forest, 18 of them placed in the subgenus Euglossa s. str. Latreille [analis group: E. analis Westwood, 1840, E. cognata Moure, 1970 and E. mixta Friese, 1899; cordata group: E. chlorina Dressler, 1982, E. cordata (Linnaeus, 1758), E. fimbriata Moure, 1968, E. gaianii Dressler, 1982, E. leucotricha Rebêlo & Moure, 1996, E. melanotricha Moure, 1967, E. milenae Bembé, 2007, E. pictipennis Moure, 1943, E. securigera Dressler, 1982 and E. violaceifrons Rebêlo & Moure, 1996; purpurea group: E. anodorhynchi Nemésio, 2006, E. avicula Dressler, 1982, E. pleosticta Dressler, 1982, E. truncata Rebêlo & Moure, 1996 and E. townsendi Cockerell, 1904], four in the subgenus E. (Euglossella) Moure, 1967 [E. decorata, Smith, 1874, E. mandibularis Friese, 1899, E. perpulchra Moure & Schlindwein, 2002 and E. viridis (Perty, 1833)], four in the subgenus E. (Glossura) Cockerell, 1917 [E. annectans Dressler, 1982, E. ignita Smith, 1874, E. imperialis Cockerell, 1922 and E. iopoecila Dressler, 1982], three in E. (Glossurella) Dressler [E. crassipunctata Moure, 1968, E. sapphirina Moure, 1968 and E. stellfeldi Moure, 1947] and one species placed in E. (Glossuropoda) Moure, 1989 [E. cyanochlora Moure, 1996]. The number of species presented by Moure et al. (2007) is slightly different from the proposal of Nemesio (2009), in which 35 species are assumed to be inhabiting this biome. Distinct status given to a few species, mainly in the subgenera E. (Glossura) and E. (Glossurella), are the reason for this difference in species number.

Despite the lower diverstiy in the Atlantic forest, new species continue to be described (e.g., Rebêlo & Moure 1996, Moure & Schlindwein 2002, Nemésio 2006, 2009, 2010a, 2011b, Faria & Melo 2011), which suggests that orchid bee richness may be underestimated. Considering the outstanding biodiversity of this biome _ one of the 25 world's biodiversity hotspots (Myers et al. 2000) _ coupled with rapid habitat loss and anthropogenic pressure [the Atlantic forest has lost more than 93% of its original area (Tabarelli et al. 2005)], taxonomic studies focusing on species inhabiting this biome are imperative. Besides the intrinsic value of knowing the biodiversity (see Bengtsson et al. 1997), information on the taxonomy of a group is fundamental to the formulation of effective programs for species conservation (Lowry 2001, Brito 2004).

According to Moure et al. (2007), three species of the analis group inhabit the Brazilian Atlantic forest: E. analis, E. cognata and E. mixta. Nemésio (2009), on the other hand, considered that five species can be found in this domain, the above three considered by Moure et al. (2007), besides E. iopyrrha Dressler, 1982 and E. roderici Nemésio, 2009. It must be pointed out that when Moure et al. (2007) published their contribution, the description of E. roderici had not yet been published. The identity of the species in the analis group in the Atlantic forest is far from being resolved. The statement of Nemésio (2009: 143) in his monograph illustrates this scenario: "Further studies are needed for the entire Eg. analis group from the Atlantic Forest, particularly the specimens treated as Eg. mixta and Eg. iopyrrha, to be conclusive concerning their actual identity. It is possible that the specimens treated as Eg. mixta and Eg. iopyrrha from the Atlantic Forest belong to new, undescribed species". Indeed, the form occurring in the Atlantic forest, and previously identified as E. analis, has been recently described by Nemésio (2011b) as a new species _ Euglossa marianae Nemésio, 2011.

Another important issue regarding the identity of the species in the analis group found in the Atlantic forest concerns their geographic range. Three of the five species assumed to inhabit this biome supposedly have wide distributions in the Neotropical region: E. iopyrrha would be found in the Atlantic and Amazon forests (South American species sensu Nemésio 2009), while E. cognata and E. mixta are assumed to be pan-neotropical species, being both found in South and Central America (Moure et al. 2007, Nemésio 2009). The correct definition and identification of widely distributed species is one of the main challenges in taxonomy (e.g., Gill & Kemp 2002) and, particularly regarding the species placed in the analis group, the type locality of the four aforementioned species is not in the Atlantic forest domain. Therefore, it is possible that the names used in the identification of the forms inhabiting the Atlantic forest may constitute, in fact, a misapplication of names of species found elsewhere in the neotropics. The accurate identification of species of the analis group may be even more important in this scenario, since some of them (e.g., E. marianae) have been considered bioindicators, due to their close association to well-preserved forest fragments (Tonhasca et al. 2002, Nemésio & Silveira 2006, Ramalho et al. 2009 as E. analis, see Nemésio 2011b for details).

This contribution represents a taxonomic study of the species belonging to the analis group inhabiting the Brazilian Atlantic forest, including description of new species, new sex associations, identification key based both on males and females, and distribution maps.

 

MATERIAL AND METHODS

Depository institutions and their acronyms are as follows: Departamento de Zoologia, Universidade Federal de Minas Gerais, Belo Horizonte, Brazil (DZMG); Coleção de Entomologia Pe. J.S. Moure, Departamento de Zoologia, Universidade Federal do Paraná, Curitiba, Brazil (DZUP); Museu de Zoologia da Universidade de São Paulo, São Paulo, Brazil (MZSP); Laboratório de Ciências Ambientais, Universidade Estadual do Norte Fluminense, Campos dos Goytacazes, Brazil (UENF); Hope Entomological Collections, Oxford University Museum of Natural History, Oxford, England (OUM); and Zweite Zoologische Abteilung, Naturhistorisches Museum Wien, Vienna, Austria (NMW).

The morphological terminology adopted here follows Dressler (1982), Michener (2000), and Roubik (2004). Metasomal terga and sterna are indicated, respectively, as T1 to T7, and S1 to S8. The density of punctation and intervals between the punctures were based on relative puncture diameter, pd (e.g., <1pd: less than 1x the puncture diameter between the punctures). Also, the term beveled puncture follows the definition presented in Cure (1989) and translated by Almeida (2008) from the Portuguese "pontuação biselada". All measurements are in millimeters and integumental color was described under a white fluorescent lamp. The color images were obtained with a Leica DFC 500 camera, associated to a stereomicroscope MZ 16, and processed by the software Automontage (Syncroscopy), a facility provided by the Taxonline Project (Rede Paranaense de Coleções Biológicas, Departamento de Zoologia, UFPR).

The labels of examined specimens were transcribed in the section "Material examined", where one inverted bar symbol (\) indicates the different lines in the label, two inverted bars (\\) indicate information on the back side of the label, and the double quotation marks indicate different labels associated with one specimen. In the labels, the signs of male and female were transcribed as M and F, respectively. When a label was similar to other associated to a specimen, only the different information between the labels is presented between single quotation marks. In the Appendices 1 and 2, when the depositary institution is not indicated, the material belongs to the DZUP.

Distribution maps for the studied species were generated in DIVA-GIS 7.3 (Hijmans et al. 2001), based on locality data from examined specimens. Records outside the study area were not included in the maps for E. cognata, a species widely distributed outside eastern Brazil. In the maps, the Atlantic forest is represented in grey. The shapefile, derived from the ecoregions presented in Olson & Dinerstein (2002), was obtained at http://worldwildlife.org/science.

 

TAXONOMY

The analis group

In an important paper for the taxonomy of Euglossa, Dressler (1978) dealt with the infrageneric classification of this genus. Besides the description of a new subgenus and delimitation of other subgenera, Dressler proposed the delimitation of some species groups: "Then I will present a revised subgeneric classification and delineate a number of species groups that have proven useful in the identification of Euglossa"(Dressler 1978: 188). According to his proposition, the subgenus Euglossa sensu stricto was divided into five species groups: analis, viridissima, cybelia, purpurea and cordata.

The analis group was delimited by Dressler (1978) based on the following characters found in males: posterior tuft of mid tibia usually reduced or lacking, anterior tuft shallowly notched or attenuate; hind tibia usually rhomboid; mandible with two or three teeth; parocular white markings present; integument predominantly dark blue-violet with the apical terga usually green or bronze.

The species in this group are also characterized by the coarse and dense punctation on the mesoscutum and scutellum, and by the preponderance of beveled punctures on the terga, particularly on T2. Also, in this group the lateral corners of the pronotal collar are conspicuously pointed, both in males and females. This feature had been already pointed out by Moure (1968: 22, 1970: 157) in the original descriptions of Euglossa villosiventris Moure, 1968 and E. cognata, respectively, as well as by Dressler (1982: 122). Males of most species of the analis group, as stated by Faria & Melo (2007: 276), also possess a characteristic depression on the inner surface of their hind tibia. Only males of E. analis and E. villosiventris have unmodified hind tibiae, lacking the depression.

In Dressler's (1978) original proposition, the analis group contained four species, E. analis, E. cognata, E. mixta and E. villosiventris. After his work, five additional species have been proposed: Euglossa bidentata Dressler, 1982, E. iopyrrha, Euglossa retroviridis Dressler, 1982, E. roderici and E. marianae.

Another available name, Euglossa aureiventris Friese, 1899, also belongs in the analis group. The female lectotype (NMW), examined through photographs as well as based on Moure's notes, shows unequivocal features associated with this species group: pattern of body color with a predominantly greenish-blue head and mesosoma, including legs, and basal terga mostly coppery red; lateral corners of the pronotal collar pointed; scutellar tuft relatively small, its length occupying a little less than one-half of scutellum length; pubescence on T2 dense and composed of relatively long dark setae. It is possible that E. aureiventris might turn out to be the female of E. analis or of E. retroviridis.

Besides the ten species mentioned above and the two new ones proposed here, study of the DZUP collection indicates that additional undescribed species should be recognized in the analis group (G.A.R. Melo, unpub. data). However, a revision of the entire group is beyond the scope of the present contribution, which focuses on the species found in the Brazilian Atlantic forest, considering as separate species some forms previously regarded as belonging to taxa with a wider geographic distribution.

Five species are recognized based on males for this region: E. cognata, E. marianae, E. roderici and the two newly proposed species, E. botocuda sp. nov. and E. calycina sp. nov. Except for E. botocuda sp. nov., it was possible to confidently associate female specimens for the other four species. Although the current taxonomy of orchid bees is based almost exclusively on males, description of females is also an important step to improve our precarious knowledge of sex association for the species of Euglossa.

Distributional patterns of two species, E. botocuda sp. nov., and E. roderici should be highlighted, since they seem to be restricted to particular areas in the Atlantic forest. Euglossa botocuda sp. nov. is restricted to the "Hileia Baiana", a pattern that also appears in the distribution of other euglossine species, such as Eufriesea brasilianorum (Friese, 1899) and E. cyanochlora (see Nemésio 2009). The "Hileia Baiana"region, encompassing southern Bahia and northern Espírito Santo, is recognized by its high levels of endemism (e.g., Thomas et al. 1998) and the discovery of E. botocuda sp. nov. reinforces this pattern and the importance of this particular region in the context of the biogeography of the Atlantic forest. Euglossa roderici, on the other hand, is restricted to the coastal forests from northeastern São Paulo to northern Santa Catarina. Endemicity in this region of the Atlantic forest has also been previously detected (e.g., Pinto-da-Rocha et al. 2005), a pattern supported by the presence of E. roderici. It must be pointed out, however, that the southern portion of the Atlantic forest, in southern São Paulo and Paraná states, has a low diversity of euglossine bees, with few endemic elements (Mattozo et al. 2011).

Key to the species of the analis group in the Atlantic forest (geographic distribution indicated by acronyms of Brazilian states)

Males

1. Mandible bidentate (Fig. 26). T5 varying from coppery to golden green, T6-T7 bright green (Fig. 27). Posterior mesotibial tuft conspicuous, approximately as large as half ocellar diameter (Fig. 47). Depression on inner surface of hind tibia restricted to its apical half (Fig. 54). BA, ES, MG, RJ, SP................... Euglossa marianae

1'. Mandible tridentate (Figs 3, 9, 18). T5-T7 coppery red, bright red or magenta (Figs 4, 10, 19, 35). Posterior mesotibial tuft absent or vestigial (Figs 43-45, 49). Depression on inner surface of hind tibia extending over part of its basal half (Figs 51-53, 55)................... 2

2. Sligthly larger bees, maximum head width usually over 5 mm. Punctation on mesoscutum very dense, punctures almost contiguous; pilosity on disc relatively short, longest setae conspicuously shorter than length of first flagellomere. Anterior mesotibial tuft comma-shaped, its apical portion conspicuously narrowed and directed anterad (Fig. 45). Hind tibia more swollen (best seen in posterior view); postglandular fringe relatively short, longest setae distinctly shorter than maximum width of glandular opening in outer view (Fig. 20). BA, ES, MG................... Euglossa cognata

2'. Smaller bees, maximum head width usually less than 5 mm. Punctation on mesoscutum less dense, punctures in general not contiguous and separated from one another by one-third to one-half puncture diameter; pilosity on disc longer, longest setae about as long as length of first flagellomere. Anterior mesotibial tuft oblong, its apical portion about as wide as basal portion (Figs 43, 44, 49). Hind tibia less swollen; postglandular fringe longer, longest setae at least as long as maximum width of glandular opening in outer view (Figs 51, 52, 55)................... 3

3. Head and mesosoma, including lateral portion of mesepisternum, dark blue to violet. Parocular ivory stripe short and slender, widely separated from clypeus (Fig. 34). Pilosity on mesepisternum and fore femur almost entirely black (Fig. 33). SP, PR, SC................... Euglossa roderici

3'. Head and mesosoma mostly blue with some bluish-green areas, especially on head; lateral portion of mesepisternum mostly bluish-green to greenish-blue. Parocular ivory stripe usually long and almost reaching clypeus (Figs 2, 8), only rarely short and slender. Pilosity on mesepisternum and fore femur mostly white, at most with some dark hairs intermixed (Figs 1, 7)................... 4

4. Anterior mesotibial tuft varying from oblong to quadrangular, its anterior margin almost straight, notch between basal and apical lobe very inconspicuous (Fig. 44), lobes when noticeable subequal in length. Pit around vestigial posterior mesotibial tuft conspicuous, extending over margin of tibia (Fig. 11). Mandibular teeth subequal in length (Fig. 9). Postglandular fringe relatively short (Fig. 12), longest setae ca. 0.45 mm in length. Protuberance on inner surface of apex of hind tibia less pronounced than projection over base of inner spur. T5 dark magenta to purple, T7 dark red. PE, BA, ES, MG................... Euglossa calycina sp. nov.

4'. Anterior mesotibial tuft distinctly oblong, its anterior margin with a discrete notch, basal lobe distinctly longer than apical lobe (Fig. 43). Posterior mesotibial tuft absent, pit very shallow and inconspicuous. Apical mandibular tooth distinctly longer than subapical teeth (Fig. 3). Postglandular fringe longer (Fig. 5), longest setae ca. 0.6 mm in length. Protuberance on inner surface of apex of hind tibia more developed than projection over base of inner spur. T5 magenta laterally and dark red in the middle, T7 bright coppery red. BA, ES................... Euglossa botocuda sp. nov.

Females (the female of E. botocuda sp. nov. is not known)

1. Head and mesosoma, including legs, mostly dark blue to violet (Figs 37-40), except for greenish-blue lower parocular area and lower gena. Mesepisternum and propodeum with mostly dark pubescence. SP, PR, SC................... Euglossa roderici

1'. Head predominantly green (Figs 14, 22, 30), at most with reddish-yellow reflexes on clypeus and frons; when blue portions present, restricted to upper frons and vertex; mesosoma variable, at least mesepisternum, metepisternum and propodeum mostly green. Mesepisternum and propodeum with mostly white pubescence or at most with some dark hairs intermixed................... 2

2. Clypeus, lower parocular area and mid frons yellowish-green, with some reddish and bluish reflexes (Fig. 22); mid longitudinal carina on clypeus reddish-purple; upper frons and vertex mostly dark blue (Fig. 23); mesoscutum and scutellum mostly greenish-blue to blue (Fig. 24). BA, ES, MG................... Euglossa cognata

2'. Head and mesosoma predominantly green, at most with some bluish reflexes or small blue portions on tegulae and metapostnotum (Figs 14-16, 30-32)................... 3

3. Vertex and mesoscutum entirely green (Figs 31, 32); metapostnotum green; T5-T6 (Fig. 29) and metallic portions of sterna bright green. BA, ES, MG, RJ, SP................... Euglossa marianae

3'. Vertex and mesoscutum with abundant bluish reflexes (Figs 15, 16); metapostnotum greenish-blue; T5-T6 (Fig. 13) and metallic portions of sterna coppery to dark red. PE, BA, ES, MG................... Euglossa calycina sp. nov.

Euglossa botocuda sp. nov.

Figs 1-6, 43, 51, 56

Diagnosis. This species can be recognized by its tridentate mandible; posterior mesotibial tuft absent, and surrounding pit very shallow and inconspicuous; anterior mesotibial tuft distinctly oblong, its anterior margin with a discrete notch, basal lobe longer than apical lobe; apical lobe of anterior tuft distinctly wider than basal lobe; depression on the inner surface of hind tibia extending over its basal half (Fig. 51); a relatively long postglandular fringe; T5 magenta laterally and dark red in the middle, T6-T7 bright coppery red.

Description. Holotype male. Body length: 10.9; head width: 4.7; maximum interorbital distance: 3.0; Color: ivory parocular markings well-developed, almost reaching clypeus and extending to tangent to lower rim of antennal sockets; clypeus blue to turquoise blue; upper frons, vertex and upper gena blue; anterior surface of scape black; mesoscutum and scutellum dark blue with some purple reflexes; mesepisternum mostly blue laterally and becoming golden green ventrally; hind tibia blue becoming turquoise blue toward basitarsus; T1-T4 purple, with some dark blue reflexes, T3-T4 with stronger reddish reflexes laterally; T5 red with some purple reflexes; T6 and T7 coppery orange red.

Pubescence: predominantly white; gena densely covered with long plumose white hairs (ca. 0.98); transverse band on upper frons with mostly black setae (longest ones ca. 0.43); long black setae on vertex ca. 0.82; mesoscutum covered with mostly simple black setae (ca. 0.35-0.38); longest black setae along posterior margin of scutellum ca. 0.43; upper portion and posterior one-half of lateral portion of mesepisternum with abundant plumose black setae intermingled; upper quarter of metepisternum with mostly finely plumose black setae; anterior mesotibial tuft as in Fig. 43; posterior tuft vestigial, practically absent; longest setae of postglandular fringe with ca. 0.55-0.6 mm in length; T2-T4 covered mostly with short black setae, about as long as puncture diameter; T5-T7 mostly with long white setae, longest ones on T7 ca. 0.6 mm in length.

Integumental surface: mesoscutum and scutellum densely and coarsely punctured, distance between punctures about one-third pd on most of mesoscutum, punctures on scutellum slightly coarser, especially along its posterior margin; central portion of disc of T2-T3 with relatively coarse and strongly beveled punctures, puncture diameter about as large as those on lower parocular area; T4 with slightly coarser and more beveled punctures; on T5-T7, punctures much coarser and more strongly beveled.

Strucuture (measurements in mm): mandible tridentate, apical mandibular tooth distinctly longer than the subapical teeth; head about 1.75x wider than long (4.8:2.7); ocelo-orbital distance, in dorsal view, about 1.7x the distance between posterior ocelli (0.59:0.35); scape, excluding radicle, about 3.3x longer than its maximum width (0.90:0.27); clypeus 1.03x longer than wide (1.38:1.33); scutellum about 2.3x wider than long (3.03:1.33); pit around vestigial posterior mesotibial tuft very shallow and inconspicuous, hind tibia about 1.03x longer than the maximum width (3.27:3.15), depression on its inner surface extending over part of its basal half.

Female. Unknown.

Type material. Holotype male, "Brasil, Bahia,\Itamaraju,\1-10.iii.1971, C. Elias"(DZUP). Paratypes: 1 male (DZUP), "BRAZIL: BAHIA\Res. Mte. Paschoal\8/XI/1968\R. L. Dressler\1251""\\methyl\salicylate""mixta"; 1 male (DZUP), "BRAZIL: E. Santo\No. Linhares\12/XI/1968\R.L.Dressler\ 1252""\\methyl cinnamate"; 1 male (DZUP), "BRAZIL: E. Santo\CONCEIÇÃO DA BARRA\13 XI/1968 6\R. L. Dressler 1285""\\Methyl\salicylate"; 1 male (DZUP), "BRAZIL: E. Santo\CONCEIÇÃO DA BARRA\15 XI/1968 6\R. L. Dressler 1285""\\Methyl\salicylate"; 1 male (DZUP), "BRAZIL: E. Santo\CONCEIÇÃO DA BARRA\15 XI/1968 6\R. L. Dressler 1285""\\Methyl\salicylate""Euglossa\iopyrrha Moure\det. R. L. Dressler 1968\\E. cognata?".

Distribution (Fig. 56). From southern Bahia to northern Espírito Santo, restricted to the lowland coastal forests. Nemésio's (2009: 132) records of E. iopyrrha from mountainous sites in Espírito Santo (Santa Teresa and Domingos Martins) mostly likely refers to E. calycina sp. nov. and not to E. botocuda sp. nov.

Etymology. The specific epithet honors the "botocudos"(also known as Aimorés or Tapuias), the foreign name for an ethnic group of natives famous for their bravery. They inhabited large areas of the Brazilian Atlantic forest, probably from the "Vale do Salitre"(Bahia) in the north to the "Rio Doce"(Espírito Santo) in the south (Paraíso 1992, Duarte, 1998) encompassing most of the known distribution of this new species.

Remarks. Euglossa botocuda sp. nov. is very similar to E. iopyrrha, both having the apical mandibular tooth distinctly longer than the subapical teeth and lacking the posterior tuft of the mid tibia. It differs from E. iopyrrha by its predominantly greenish-blue head (mostly violet blue in E. iopyrrha); in the details of the anterior midtibial tuft, shorter and with basal and apical lobes very weakly differentiated from one another, anterior margin of apical lobe aligned with that of basal lobe (in E. iopyrrha, midtibial tuft longer and with basal and apical lobes distinct from one another, anterior margin of basal lobe distinctly separate from anterior margin of mid tibia compared to apical lobe; Fig. 46); depression on inner surface of hind tibia relatively long, extending over part of its basal half (in E. iopyrrha, depression restricted to apical half of hind tibia). Previous papers dealing with the Atlantic forest forms have misinterpreted E. botocuda sp. nov. as populations of E. iopyrrha from eastern Brazil (Moure 1970: 155, Silveira et al. 2002: 77, Nemésio & Silveira 2007: 887, Neves & Viana 2003: 225, 226); Nemésio 2009: 4, 13, 17, 74, 102, 114, 132, 141, 143, 149, 208, 209, 210, 211).

Attractive compounds. Cineole, methyl cinnamate and methyl salicylate.

Additional examined material. 107 males (see Appendix 1).

Euglossa calycina sp. nov.

Figs 7-16, 44, 52, 57

Diagnosis. Males of this species can be recognized by their tridentate mandible, the teeth subequal in length; posterior mesotibial tuft vestigial; anterior midtibial tuft varying from oblong to quadrangular, its anterior margin almost straight, notch between basal and apical lobe very inconspicuous, lobes when noticeable subequal in length; pit around vestigial posterior mesotibial tuft very conspicuous, extending over margin of tibia; depression on the inner surface of hind tibia extending over its basal half (Fig. 52); T5-T6 dark magenta to purple, T7 dark red.

Description. Holotype male. Body length: 10.8; head width: 4.7; maximum interorbital distance: 2.9. Colour: ivory parocular markings well-developed, lower portion wider and almost in contact with clypeus, upper portion extending to lower rim of antennal sockets; disc of clypeus blue, with a lower transverse green band adjacent to the labrum; lateral portion of clypeus and lower parocular area mostly greenish-blue; frons green to turquoise blue; anterior surface of scape black; mesoscutum and scutellum mostly dark blue, with some violet reflexes; mesepisternum mostly green, except for turquoise blue upper portion; hind tibia mainly dark blue, with turquoise blue reflexes toward hind basitarsus; T1-T3 mainly purple; T4 violet basally and magenta along marginal zone; T5 dark magenta; T6 and T7 dark red with magenta reflexes.

Pubescence: predominantly white; gena densely covered with long plumose white hairs (ca. 0.90); transverse band on upper frons with mostly black setae (longest ones ca. 0.47); long black setae on vertex ca. 0.78; mesoscutum covered with mostly simple black setae (ca. 0.35-0.40); longest black setae along posterior margin of scutellum ca. 0.39; lateral portion of mesepisternum with thick plumose black setae intermixed with finely plumose white hairs; upper half of metepisternum densely covered with finely plumose black setae; upper portion of propodeum laterally, anteriorly to spiracle, with abundant dark plumose setae; anterior mesotibial tuft as in Fig. 44; posterior tuft absent; postglandular fringe with ca. 0.400.45 mm in length; T2-T4 covered mostly with short black setae, about as long as puncture diameter; T5 with a mixture of short black setae and sparse long white setae; T6-T7 mostly with long white setae, longest ones on T7 ca. 0.74 mm in length.

Integumental surface: mesoscutum densely and coarsely punctured, distance between punctures about one-third pd on most of mesoscutum; punctures on disc of scutellum sparser; central portion of disc of T2-T3 with relatively coarse and strongly beveled punctures, puncture diameter about as large as those on lower parocular area; T4 with slightly coarser and more beveled punctures; on T5-T7, punctures much coarser and more strongly beveled.

Structure (measurements in mm): mandible tridentate, teeth subequal in length; head about 1.6x wider than long (4.7:2.9); ocelo-orbital distance, in dorsal view, about 1.6x the distance between posterior ocelli (0.63:0.39); scape, excluding radicle, about 3.4x longer than its maximum width (0.86:0.25); clypeus 1.03x longer than wide (1.30:1.26); scutellum about 2.2x wider than long (2.87:1.30); pit around vestigial posterior mesotibial tuft conspicuous (Fig. 11), extending over margin of tibia; hind tibia about 1.02x longer than the maximum width (3.11:3.03), depression on its inner surface extending over part of its basal half.

Female (specimen from Santa Teresa, ES). Body length: 10.3; head width: 4.5; maximum interorbital distance: 2.9; tongue in repose reaching base of S1; labrum 1.32x wider than long (1.14:0.86); scutellum 2.3x wider than long (2.83:1.22); navicular tuft 0.41x the length of scutellum (0.51:1.22); hind tibia 1.4x longer than wide (2.83:2.0); posterior lobe of corbicula 0.51x the hind tibia's width (1.02:2.0); labrum ivory white between pair of oval spots; clypeus, lower parocular area and frons mostly green, with light yellowish reflexes; vertex mainly green with strong blue hues in front of ocelli; mesoscutum green laterally, with turquoise blue hues medially in the region of the medial line; scutellum green/turquoise blue; mesepisternum green with some bluish reflexes; metapostnotum mostly blue with greenish reflexes; hind tibia green with turquoise blue reflexes; T1 purple; T2 purple medially and magenta posteriorly, its lateral ventral portions dark red; disc of T3 purple basally and mainly magenta posteriorly, its lateral portions dark red; T4-T6 light magenta to dark red; metallic portions of sterna predominantly red; pubescence of mesepisternum mostly white and finely plumose, with abundant black setae intermixed; pubescence on dorsal surface of hind femur predominantly pale yellow, with a few dark hairs intermixed; longer thick setae on borders of hind tibia predominantly dark brown to black, with a few pale yellow setae along anterior border; short black setae on disc of T2 about as long as 1.5 pd; mesoscutum and scutellum densely and coarsely punctured, punctures separated by about one-third pd; T2 densely punctured, punctures strongly beveled and smaller than those on lower parocular area.

Type material. Holotype male, "BRAZIL: E. Santo\CONCEIÇÃO DA BARRA\10 XI/1968\R.L. Dressler 1271""\\skatole"(DZUP). Paratypes: 3 males (DZUP), "BRAZIL: E. Santo\CONCEIÇÃO DA BARRA\10 XI/1968\R.L. Dressler 1271""\\skatole"; 1 male (DZUP), "BRAZIL: E. Santo\CONCEIÇÃO DA BARRA\10 XI/1968\R.L. Dressler 1271""\\vanillin"; 1 male (DZUP), BRAZIL: E. Santo\No. Linhares\12/XI/1968\R.L. Dressler\1292""Methyl\salicylate".

Additional examined material. 421 males and 2 females (see Appendix 1).

Distribution (Fig. 57). From Pernambuco to Espírito Santo, mainly in lowland forests along the coast. It has been found also in more inland sites along the Mucuri river basin, in Minas Gerais, and the Pardo river basin, in Bahia, as well as in submontane forests in Espírito Santo.

Attractive compounds: cineole, methyl salicylate, skatole and vanillin.

Etymology. The specific epithet refers to the Latin calyx (cup), in reference to the well-developed pit around the vestigial posterior midtibial tuft of the males.

Variation. There is some variation in the color of the plumose pubescence on the lateral portion of the propodeum, with some males having abundant dark hairs among the white ones, as in the holotype, while others having only white hairs. Also, differently from the holotype, some specimens have a vestigial posterior mesotibial tuft.

Remarks. Euglossa calycina sp. nov. is most similar to E. mixta and E. roderici. Its male differs from that of E. mixta by its turquoise blue head and mesepisternum (violet blue in E. mixta); anterior midtibial tuft shorter, only slightly longer than wide (in E. mixta distinctly longer than wide); posterior tuft absent or completely vestigial, pit around it very conspicuous, extending over margin of tibia (in E. mixta posterior tuft small, but visible, pit around it shallow, not pronounced) (Figs 44 and 48); T6-T7 predominantely dark red, with coppery reflections (T6 and especially T7 coppery red in E. mixta). From E. roderici, the new species differs mostly by the characters given in couplet 3 of the identification key and by the details of the midtibial tufts (see under E. roderici). A female of E. calycina sp. nov., identified as E. mixta, has been illustrated and briefly described by Nemésio (2009: 142, Fig. 136). Previous papers dealing with the Atlantic forest forms have misinterpreted E. calycina sp. nov. as populations of E. mixta from eastern Brazil (Friese 1899: 135, 1930: 134, Schrottky 1902: 590, Lutz & Cockerell 1920: 548, Silveira et al. 2002: 78, Neves & Viana 2003: 225, 226, Nemésio & Silveira 2007: 888, Moure et al. 2007: 233, Nemésio 2009: 4, 13, 17, 74, 102, 114, 132, 141, 142, 143, 149, 208, 211, 2010b: 58, 59, 60, 2011a: 49, 50, 2011b: 62) or as populations of E. iopyrrha (Milet-Pinheiro & Schlindwein 2005: 855, 856, Darrault et al. 2006: 244, 245).

Euglossa cognata Moure, 1970

Figs 17-24, 45, 53, 58

Euglossa cognata Moure, 1970: 156. Holotype male, Brazil, Pará, Mocajuba, 'Mangabeira' (DZUP), examined.

Description. Male (specimen from Porto Seguro, BA). Body length: 12.9; head width: 5.1; maximum interorbital distance: 3.1; mandible tridentate, apical teeth longer than subequal teeth; tongue in repose slightly surpassing apex of hind coxae; disc of clypeus dark blue and turquoise blue laterally; frons dark blue-violet, becoming turquoise blue towards compound eyes; vertex blue-violet; lower gena green; ivory parocular markings well-developed, almost in contact with clypeus and reaching tangent to lower rim of antennal sockets; anterior surface of scape black; mesoscutum and scutellum mostly dark blue-violet; mesepisternum blue to violet blue laterally and bluish-green ventrally; hind tibia mostly blue, with small violet and green reflexes; T1-T4 mainly purple; T5 dark red to magenta; T6 dark red; T7 coppery red; mesoscutum with mostly simple black setae (ca. 0.3-0.33); longest black setae along posterior margin of scutellum ca. 0.4; upper portion and posterior half of mesepisternum with sparse black setae intermixed with finely plumose white setae; upper half of metepisternum mostly with finely plumose black setae; upper portion of propodeum laterally, anteriorly to spiracle, with abundant dark plumose setae; anterior mesotibial tuft comma-shaped (Fig. 45); posterior tuft vestigial; postglandular fringe short (ca. 0.35); depression on inner surface of hind tibia extending over part of its basal half (Fig. 53); mesoscutum and scutellum densely and coarsely punctured, punctures on mesoscutum mostly contiguous; central portion of disc of T2 densely punctured (<0.5pd), punctures strongly beveled.

Female (specimen from Conceição da Barra, ES). Body length: 11.7; head width: 5.0; maximum interorbital distance: 3.12; tongue in repose reaching apex of hind coxae; labrum 1.28x wider than long (1.26:0.98); scutellum relatively long, about only twice as wide as long (2.95:1.46); navicular tuft 0.41x the length of scutellum (0.6:1.46); hind tibia 1.35x longer than wide (3.15:2.32); posterior lobe of corbicula 0.43x the hind tibia's width (1.02:2.32); labrum ivory white between pair of oval spots; disc of clypeus yellowish-green with strong reddish reflexes, mid longitudinal carina reddish-purple; lateral portions of clypeus and supraclypeal area green; lower parocular area and mid frons yellowish-green with some coppery reflexes; upper frons and vertex mainly dark blue, except for bluish-green upper parocular area; mesoscutum mainly blue with two well-defined bluish-green stripes laterally to the medial line; scutellum mainly turquoise blue; mesepisternum green with weak bluish reflexes; metapostnotum blue; hind tibia bluish-green with strong purple hues; T1-T2 purple; T3 magenta with strong purple reflexes; T4 magenta, gradually becoming red laterally; T5-T6 bright red, with some magenta reflexes; metallic portions of sterna predominantly red, except for coppery yellow on S6; pubescence of mesepisternum mostly white and finely plumose, with abundant black setae intermixed; pubescence on hind femur entirely black; longer thick setae on borders of hind tibia dark brown to black; short black setae on disc of T2 at most as long as 1.5 pd; mesoscutum and scutellum densely and coarsely punctured, punctures separated by about one-quarter pd; T2 densely punctured, punctures strongly beveled and smaller than those on lower parocular area.

Examined material from eastern Brazil. 517 males and 6 females (see Appendix 2).

Distribution (Fig. 58). In eastern Brazil, mainly in the lowland coastal forests from southern Bahia to northern Espírito Santo. It has been found also in more inland sites along the Doce river in Minas Gerais (Nemésio 2009: 114).

Remarks. This species is interpreted here as originally proposed by Moure (1970), with a wide distribution, occurring from eastern Brazil to northern South America and reaching Costa Rica, in Central America (Roubik & Hanson 2004). Except for a slightly smaller body size, both males and females from eastern Brazil are identical to those from the Amazon basin. There is some variation in the size of the posterior tuft in the male mid tibia. In most specimens, the tuft is very small, its diameter comparable to that of a puncture of the mesoscutal disc. In a few specimens, the tuft is totally lacking, while in a few others, it is more conspicuous, being slightly wider than a mesoscutal puncture.

Attractive compounds. Benzyl acetate, cineole, eugenol, skatole, methyl salicylate and vanillin.

Euglossa marianae Nemésio, 2011

Figs 25-32, 47, 54, 59

Euglossa marianae Nemésio, 2011b: 63. Holotype male, Brazil, Minas Gerais, Marliéria, Parque Estadual do Rio Doce (DZMG), not examined.

Description. Male (specimen from Conceição da Barra, ES). Body length: 11.5; head width: 4.8; maximum interorbital distance: 3.05; mandible bidentate; tongue in repose reaching apex of hind coxae; disc of clypeus dark blue, with green areas laterally towards lower parocular area; frons mostly turquoise blue; vertex and upper gena greenish-blue; ivory parocular markings well-developed, almost reaching clypeus and barely reaching tangent to lower rim of antennal sockets; anterior surface of scape black; mesoscutum dark blue medially with turquoise blue/green areas toward tegulae and violet areas toward scutellum; scutellum violet; mesepisternum turquoise blue laterally and green ventrally; hind tibia dark blue/violet basally and dark green next to glandular opening; T1-T4 purple with violet blue reflexes; T5 coppery yellow with red reflexes; T6 and T7 bright yellowish-green; mesoscutum covered with mostly simple black setae (ca. 0.38), setae becoming shorter towards scutellum; longest black setae along posterior margin of scutellum ca. 0.4; mesepisternum with finely plumose white setae; upper quarter of metepisternum mostly with finely plumose black setae; anterior mesotibial tuft as in Fig. 47; posterior tuft conspicuous, approximately as large as half ocellar diameter; postglandular fringe relatively short (ca. 0.45-0.5); depression on inner surface of hind tibia restricted to its apical half; mesoscutum and scutellum densely and coarsely punctured, punctures near parapsidal line separated by one-fifth of puncture diameter to mostly contiguous; central portion of disc of T2 densely punctured (<0.5 pd), punctures weakly beveled.

Female (specimen from Linhares, ES). Body length: 11.0; head width: 4.72; maximum interorbital distance: 2.96; tongue in repose reaching S1; labrum 1.46x wider than long (1.1:0.75); scutellum 2.1x wider than long (2.75:1.30); navicular tuft 0.36x the length of scutellum (0.47:1.30); hind tibia 1.45x longer than wide (2.87:1.97); posterior lobe of corbicula 0.47x the hind tibia's width (0.94:1.97); labrum dark brown between pair of oval spots; clypeus bright green, except for thin reddish reflexes along mid longitudinal carina; frons and vertex green; mesoscutum, scutellum and hind tibia green; mesepisternum green, with weak yellowish reflexes; metapostnotum green; T1 purple; T2 dark purple anteriorly, becoming reddish posteriorly, its lateral ventral portions mostly yellowish-green; disc of T3 and T4 coppery red, gradually turning into bright yellowish-green laterally; T5 and T6 bright green; metallic portions of sterna predominantly green, laterally more yellowish-green; pubescence of mesepisternum predominantly white and finely plumose, a few black setae intermixed; pubescence on dorsal surface of hind femur predominantly pale yellow; longer thick setae on borders of hind tibia predominantly pale yellow; short black setae on disc of T2 about as long as 2 pd; mesoscutum and scutellum densely and coarsely punctured, punctures mostly contiguous; T2 densely punctured, punctures strongly beveled and smaller than those on lower parocular area.

Examined material. 764 males and 1 female (see Appendix 2)

Distribution (Fig. 59). From southern Bahia to northern Rio de Janeiro, mainly in lowland forests along the coast. It has been found also in more inland sites along the Doce river in Minas Gerais, and at the southern edge of the Chapada Diamantina, in Bahia. Nemésio (2009: 102) also provides a disjunct southern record, based on a single male collected in São Sebastião, northeastern São Paulo.

Remarks. This species can be recognized by its bidentate mandibles; posterior mesotibial tuft conspicuous, approximately as large as half ocellar diameter (Fig. 47); anterior mesotibial tuft longer than wide, noticeably notched near its mid portion (Fig. 47); depression on inner surface of hind tibia present, relatively short, restricted to its apical half (Fig. 54); T5 varying from coppery to golden green, T6-T7 bright green. Due to its green apical terga, E. marianae has been misidentified as E. analis in studies dealing with the Atlantic forest fauna (see Nemésio 2011b). We agree with Nemésio (2011b) that all records of E. analis from the Atlantic forest correspond, in fact, to E. marianae. Despite Nemésio's (2011b:63) interpretation of E. analis as "its most closely related species", these two species are only remotely related within the analis group. Euglossa marianae is most closely allied to E. bidentata, the two species being structurally almost identical, including the details of the midtibial tufts (Figs 42 and 47; E. analis, Fig. 41) and the morphology of the inner surface of the hind tibia. In E. bidentata, however, T5 is purple to coppery red, and T6-T7 are bright reddish-yellow. Future studies might indicate that E. marianae is just a color morph of E. bidentata inhabiting the Atlantic forest. There is variation in the color of T5 within E. marianae. Nemésio (2011b) states that "the three last terga are bright green", as shown in his Fig. 2C. However, many specimens studied by us, in particular those from Bahia, have a golden green to coppery yellow T5.

Attractive compounds. Cineole, eugenol, methyl salicylate, skatole and vanillin.

Euglossa roderici Nemésio, 2009

Figs 33-40, 49, 55, 56

Euglossa roderici Nemésio, 2009: 149. Holotype male, Brazil, São Paulo, São Sebastião, Parque Estadual de Ilha Bela (DZMG), not examined.

Description. Male (specimen from Paranaguá, PR). Body length ca. 10.9; head width ca. 4.7; mandible tridentate, teeth subequal in length; tongue in repose reaching apex of hind coxae; clypeus violet; frons dark blue to violet; vertex violet; parocular ivory stripe short and slender, widely separated from clypeus and barely reaching tangent to lower rim of antennal sockets; anterior surface of scape black; mesoscutum and scutellum dark blue to violet; mesepisternum dark blue to violet laterally and greenish-blue ventrally; hind tibia mostly violet; T1-T4 mostly purple, with violet reflexes; T5 dark reddish-purple; T6-T7 mostly dark red, with purple reflexes on T6; longest black setae on disc of mesoscutum ca. 0.35 and those along posterior margin of scutellum ca. 0.40-0.43; mesespisternum, metepisternum, metapostnotum and propodeum mostly with dark plumose pubescence, except for pale hairs on mesepisternum ventrally; anterior mesotibial tuft as in Fig. 49; posterior tuft vestigial; postglandular fringe relatively short (ca. 0.400.45); depression on inner surface of hind tibia extending over part of its basal half (Fig. 55); mesoscutum and scutellum densely and coarsely punctured, punctures on disc of mesoscutum separated by one-third pd; central portion of disc of T2 densely punctured (<0.5 pd), punctures weakly beveled.

Female (specimen from Morretes, PR). Body length: 11.0; head width: 4.8; maximum interorbital distance: 3.12; tongue in repose reaching hind coxae; labrum 1.33x wider than long (1.15:0.86); scutellum 2.2x wider than long (3.03:1.38); navicular tuft 0.43x the length of scutellum (0.60:1.38); hind tibia 1.56x longer than wide (3.20:2.05); posterior lobe of corbicula 0.52x the hind tibia's width (1.07:2.05); labrum pale brownish-yellow between pair of oval spots, except for narrow brown stripe uniting the spots basally; clypeus dark blue; lower parocular area and narrow stripe along orbits bluish-green; frons and vertex dark blue to violet; mesoscutum, scutellum, mesepisternum, metepisternum and metapostnotum dark blue to violet; hind tibia dark blue with violet reflexes; T1 purple; T2 dark purple basally, becoming dark magenta apically and laterally; T3-T5 dark red to magenta, with purple reflexes; metallic portions of sterna predominantly dark red to magenta, except for coppery red on S6; pubescence of mesepisternum mostly black, with a few finely plumose white setae intermixed; pubescence on hind femur entirely black; longer thick setae on borders of hind tibia dark brown to black; short black setae on disc of T2 only slightly longer than one pd; mesoscutum and scutellum densely and coarsely punctured, punctures mostly contiguous on mesoscutum; T2 densely punctured, punctures strongly beveled and smaller than those on lower parocular area.

Examined material. 66 males and 1 female (see Appendix 2).

Distribution (Fig. 56). In the coastal forests from northeastern São Paulo to southern Paraná, reaching more inland sites along the Ribeira river in São Paulo.

Remarks. The recognition of E. roderici should pose no difficulties, since it is only found in the southern portion of the Atlantic forest. The male is most similar structurally to that of E. calycina sp. nov. However, the distributions of these two species do not overlap. They differ in coloration of the integument and of the pubescence, as indicated in the identification key, as well as in details of the midtibial tufts. In E. roderici, the anterior tuft is slightly more elongate, with the apical lobe distinct and wider than the basal lobe (in E. calycina sp. nov., the anterior tuft is more compact and with weak differentiation between the basal and apical lobes). Also, the pit around the vestigial posterior tuft is well-developed in E. roderici, but to a lesser degree compared to E. calycina sp. nov.

Attractive compounds. b-ionone, methyl salicylate and skatole.

 

ACKNOWLEDGMENTS

We would like to thank the curators Carlos R.F. Brandão (MZUSP), Maria C. Gaglianone and Magali Hoffmann (UENF) for providing access to collections under their responsability; we also acknowledge Dominique Zimmermann (NWM) and James Hogan (OUM), who kindly made available photographs of the lectotype of E. aureiventris and of the holotype of E. analis, respectively; Rodrigo B. Gonçalves and Nicolle V. Sydney for their support during the visit to the MZUSP; Guilherme Silveira for his support during the visit to UENF; Vitor Nardino (Taxonline) for helping photographing the specimens; LRRF thanks the colleagues from the LBCH for their support; Luciane Marinoni for her efforts in order to ensure the implementation of the PROTAX fellowship to the first author; the authors acknowledge CNPq for funding (PROTAX and PQ). Contribution number 1869 of the Departamento de Zoologia, Universidade Federal do Paraná.

 

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Submitted: 19.III.2012; Accepted: 06.V.2012.
Editorial responsibility: Gabriel L.F. Mejdalani

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