Zootaxa 2292: 15–24 (2009) ISSN 1175-5326 (print edition)

www.mapress.com / zootaxa/ Article ZOOTAXA

Copyright © 2009 · Magnolia Press ISSN 1175-5334 (online edition)

A synopsis of Actenosigynes Moure, Graf & Urban, 1999 (Hymenoptera:

Colletidae)—new species, possible oligolecty and biogeographic comments

FERNANDO A. SILVEIRA
Departamento de Zoologia, Instituto de Ciências Biológicas, Universidade Federal de Minas Gerais. Cx. Postal 486, 30123-970 Belo
Horizonte, MG, Brazil. E-mail: fernando@icb.ufmg.br.

Abstract

A second species of Actenosigynes from the Mantiqueira mountain range, in the southeastern Brazilian state of Minas
Gerais, is described. This new taxon can be readily distinguished from the type species of the genus, A. fulvoniger
(Michener, 1989), by its entirely black-pubescent mesosoma and by the light-yellow pilosity on its terga, among other
characters. Females of the two known species are illustrated for the first time and a key is presented for their
identification. Analysis of the pollen extracted from the scopa of the holotype of the new species and field observations
of A. fulvoniger indicate that both species of Actenosigynes are oligolectic on flowers of Loasaceae as pollen sources.
This presumed couple of sister species is the first example of a speciation event related to the isolation of populations of
temperate and subtropical bee species on the top of southeastern-Brazilian mountains. It suggests that complexes of
sibling species may exist among the other bee species with similar disjunct distribution.

Key words: Paracolletinae, Leioproctus, Atlantic Forest, Loasaceae, Neotropics, Pleistocene, biogeography, taxonomy

Introduction

The generic classification of Paracolletinae has been considered unatural (Michener 1989; Alexander &
Michener 1995; Almeida & Danforth 2009) but attempts to change it have been hindered by the lack of
knowledge of the phylogenetic relationships among its supraspecific groups. Its largest genus, Leioproctus
(sensu Michener 2007), groups the taxa that presumably retain the largest set of plesiomorphies in the
subfamily (Michener 1989) and that could not be attributed to any other genus (Michener 2007). This genus
has been shown to be paraphyletic by Almeida and Danforth (2009) and, in conformance with their
phylogenetic analyses, Almeida (2008) adopted the same position previously adopted by South American
taxonomists (e.g., Urban 1995; Moure et al. 1999, 2008; Silveira et al. 2002), who considered Leioproctus
subgenera in the Neotropics as genera. Moreover, to make the classification of Leioproctus coherent across
the world, he (Almeida 2008) also elevated all subgenera of Leioproctus in the Australian region to genus
level. Since this reduces the number of paraphyletic taxa in the classification, this is the arrangement adopted
here.
Among the many subgenera recognized by Michener (1989, 2000) in Leioproctus, L. (Leioproctus) was
composed of more than 140 species in the Australian Region (Almeida 2008) and, oddly, of a single one from
southern Brazil (Leioproctus fulvoniger Michener, 1989). When describing this species, Michener did not find
it to be very similar to any of the Australian species in the genus, but also could not find any putative
apomorphies for it. To accommodate this species, Moure et al. (1999) erected the genus Actenosigynes, which
was later considered as a subgenus of Leioproctus by Michener (2007).
In this paper, a second species, described as new, is assigned to Actenosigynes, a key to distinguish its two
known species is presented and a brief discussion is made on their biogeography and possible reliance on
flower resources of Loasaceae.

Accepted by E. Almeida: 27 Oct. 2009; published: 18 Nov. 2009 15

Material and methods

The new species is described below, based on four females collected in a pasture (22º41’15”S; 45º54’01”W;
1,600 m above sea level) in the municipality of Gonçalves, between the town of Gonçalves and the village of
Cantagalo, in the Mantiqueira mountain range. This region is part of an environmental protection area, the
“Área de Proteção Ambiental Fernão Dias”. All specimens were captured while foraging on the flowers of a
thorny shrub, then unidentified.
The specimens were examined under a dissection microscope and compared to the description and to a
specimen of the type species of the genus. In the description, morphological terminology is basically that of
Michener (1944), except that the propodeal triangle is here called the metapostnotum; the flagellomeres are
referred to as F1, F2, F3, etc., and metasomal terga and sterna as T1, T2, T3, etc, and S1, S2, S3, etc,
respectively. Puncture diameter was described in five arbitrary classes—minute, fine, moderately fine,
moderately coarse and coarse. Distance between punctures was given in puncture diameters between
punctures (dp) and punctation described as very sparse (> 5 dp), sparse (1–5 dp), dense (≈ 1 dp), very dense (<
1 dp) and crowded (intervals between punctures carinate). An effort was made to include, in the description of
the new species, the characters employed by Michener (1989) in the description of A. fulvoniger to make
comparisons between them easier. In the description, characters in italics are those clearly differing from those
of the type species, according to the description of Michener (1989) and to the direct examination of a
specimen.
In the reproduction of specimen labels, items between quotation marks are in the same label and a semi-
colon separates each line in one label. Portions of words between brackets are supplements of words that
appear abbreviated in the labels. Acronyms for depository collections are: DZUP (Coleção Entomológica
“Padre Jesus Santiago Moure”, Universidade Federal do Paraná, Curitiba, Brazil); MCP (Coleção de Abelhas
do Museu de Ciências e Tecnologia da Pontifícia Universidade Católica do Rio Grande do Sul, Porto Alegre,
Brazil); UFMG (Coleção Entomológica das Coleções Taxonômicas da UFMG, Universidade Federal de
Minas Gerais, Belo Horizonte, Brazil); and UKANS (Division of Entomology, Natural History Museum, The
University of Kansas, Lawrence, USA).
Identity of the plant on which the specimens described below were collected was checked through the
microscopic analysis of the pollen carried by the holotype. Pollen extracted from its scopae was acetolized
and mounted on microscope slides and compared to the pollen extracted from a specimen of Blumenbachia
latifolia Cambess., St. Hilairie, collected in the same region as the bee and which resembled the plant the bees
were foraging on. This specimen is deposited under the reference number 75063, in the herbarium of the
Taxonomic Collections of the Universidade Federal de Minas Gerais (BHCB).

Actenosigynes Moure, Graf and Urban

Leioproctus (Leioproctus) (partim) Michener, 1989: 641; 2000: 150.

Actenosigynes Moure, Graf and Urban, 1999: 3; Silveira et al. 2002: 163; Moure et al. 2007: 723.
Leioproctus (Actenosigynes) Michener 2007: 150.

The main characters distinguishing Actenosigynes from other paracolletine genera occurring in Brazil
(according to the classification and key by Silveira et al. 2002), or from other South American subgenera of
Leioproctus (according to the classification and key by Michener 2007) are: body, especially head and
mesosoma, covered by dense, plumose pilosity; punctation ordinary, i.e., not particularly strong; mid portion
of frons, below the ocelli, convex and clearly elevated relative to the paraocular region; anterior wing with
three submarginal cells, the second one much shorter than the third; inner hind spur, in the females, finely
serrate, with more than 25 teeth; outer surface of hind basitarsus covered by branched hairs; terga without
conspicuous marginal bands of hairs or enamel-like marginal pigmentation.

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Actenosigynes fulvoniger (Michener, 1989)
(Figs. 1a–d, 3)

Leioproctus (Leioproctus) fulvoniger Michener, 1989: 691.

Actenosigynes fulvoniger; Moure et al. 1999: 3; Silveira et al. 2002: 163; Moure et al. 2007: 723.
Leioproctus (Actenosigynes) fulvoniger; Michener 2007: 150.

Diagnosis. Interocellar distance nearly two ocellar diameters; ocellocular distance about two ocellar
diameters; pubescence on dorsum of mesosoma, bright fulvous to ferruginous, plainly black on metasoma;
wings, including veins and pterostigma, dark brown; integument on S3–S6 dull, microreticulate.
Examined specimen. A female labeled “Brasil, Paraná.; Tunas do Paraná. Parque; Estadual de
Campinhos; 25º02’S 49º05’W, Alt.; 860m. 27.x.2007. L; Zanette, & J. Almeida” (deposited at UFMG).
Distribution. BRAZIL (Paraná, Rio Grande do Sul, Santa Catarina).
Flight-activity. According to data in the literature and in the labels of specimens in MCP, UFMG and
UKANS, A. fulvoniger is active during the rainy season, between October and December.
Comments. Actenosigynes fulvoniger is only known from the southernmost, temperate Brazilian states
(Moure et al. 1999, 2008). References to the biology of A. fulvoniger in the literature are meager. Wittmann
and Schlindwein (1995) recorded its females collecting pollen and nectar from the flowers of Blumenbachia
eichleri Urb. (Loasaceae) (referred to in their paper as Cajophora eichleri), and observed that these flowers
were patrolled and used as sleeping shelters by its males. Although most of the specimens for which
information is available were collected on flowers of this plant, which is considered its only pollen source,
one female, collected in Porto Alegre, state of Rio Grande do Sul, was caught on flowers of Desmodium
icanum (Gmel.) Schinz et Thell (Leguminosae, Papilionoidea) (information made available by B. Blochtein,
based on a specimen label data in MCP).

Actenosigynes mantiqueirensis sp. nov.

(Figs. 2a–d, 3)

Diagnosis. Actenosigynes mantiqueirensis can be readily distinguished from the only other species in the
genus by its entirely black-pubescent mesosoma (Fig. 2a,d) (ferruginous dorsally in A. fulvoniger—Fig. 1a,d)
and by the light-yellow pilosity on the terga (Fig. 2a,d) (entirely black in A. fulvoniger—Fig. 1a,d). Other
characters distinguishing these species are given in the identification key and (in italics) in the description,
below.
Description. Female (holotype):
Measurements (mm): Approximate body length: 10.0; maximum head width: 4.25; forewing length: 8.0;
Length of 1st, 2nd and 3rd submarginal cells measured on their posterior margins: 1.24, 0.46 and 0.86; F1 (length
× width) = 0.18×0.21; F2 = 0.12×0.20; F3 = 0.12×0.20; F4 = 0.18×0.20; F8 = 0.20×0.22; F9 = 0.20×0.22; F10
= 0.33×0.21.
Structure: mandible bidentate; basal area of labrum triangular with a central, elliptical depression; clypeus
gently convex, not protuberant, with a weak, longitudinal, central depression; clypeoantennal distance slightly
larger than diameter of antennal socket; inner eye margin slightly concave, eyes slightly converging bellow,
except for upper end, converging above; ocelloccipital distance slightly larger than ocellar diameter;
interocellar and ocellocular distances about 1.7 ocellar diameters; vertex gently convex in facial view, not
raised behing ocelli; malar area linear; scape surpassing lower margin of mid ocellus; surface of
metapostnotum depressed in relation to lateral areas of propodeum; pygidial plate rounded apically, lateral
margins diverging approximately in a 60º angle.
Body color: body black except apical half of mandible, dark ferruginous (interrupted by blackish area);
and distitarsi light ferruginous. Veins and pterostigma light ferruginous, except Sc+R, brown; wing membrane
hyaline with yellowish hue especially basally and on marginal cell.

SYNOPSIS OF THE BEE GENUS ACTENOSIGYNES Zootaxa 2292 © 2009 Magnolia Press · 17
FIGURE 1. Female of Actenosigynes fulvoniger (Michener, 1989)—a) habitus, dorsal view; b) head, frontal view; c)
metasoma, ventral view (notice the dull integument of sternal discs); d) habitus, lateral view.

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FIGURE 2. Female of Actenosigynes mantiqueirensis sp. nov., holotype—a) habitus, dorsal view; b) head, frontal view;
c) metasoma, ventral view (notice the shiny integument of sternal discs); d) habitus, lateral view.

SYNOPSIS OF THE BEE GENUS ACTENOSIGYNES Zootaxa 2292 © 2009 Magnolia Press · 19
 Pubescence: black, except brownish gray on face, light yellow on T2–T6, marginal fringes of S4 and S5,
on disc of S6 and laterally on margin of S3. On lateral surface of mandible, labrum and submarginal groove of
clypeus long, simple, stiff, semi-erect to erect; on face, genae, and vertex, fine and plumose, on disc of
clypeus intermixed with fine, simple hairs, finer and shorter on vertex; on mesosoma, similar to face, but
longer on anterolateral angles of mesoscutum, posterior fringe of scutellum, mesepisternum (shorter, very fine
and simple ventrally), and propodeum, relatively dense but not entirely hiding body surface; on front leg,
long, shortly branched on coxa, forming a fringe of long branched hairs on posterior edge of trochanter and
femur, formed by shorter, simple, stiff, semi-erect bristles on inner and outer surfaces of tibia and on whole
basitarsus, of shortly branched bristles on posterior surface of tibia; on mid leg, with finer, shorter, simple,
erect hairs on ventral surface of trochanter, otherwise, similar to front leg; on hind leg, femoral and tibial
scopae formed of long, stiff, gently curved hairs, short-branched on basal half, unbranched on apical half; on
terga, very fine and appressed, short and very sparse on T1, shorter and denser on disc of T2, longer and
denser on margin of T2 and on disc and margin of other terga; on S2–S5 forming a brush of relatively long,
semi-erect, stiff bristles restricted to the apical half of the sterna, less developed on S5, which is also fringed
by long, densely plumose hairs; on S6 forming a discal tuft of fine, poorly branched hairs.
Body surface: apical margin of clypeus impunctate, separate from disc by a narrow, very finely and very
densely punctate groove; disc of clypeus, supraclypeal area and frons coarsely and densely punctate, interval
between punctures less than their diameters, crowded on the longitudinal, central depression of the clypeus ;
on paraocular area moderately coarse and densely punctate; on vertex, including interocellar space, finely and
densely punctate, except for dull, irregular, impunctate area between eye and lateral ocellus, which narrowly
extends behind ocellus; on mesoscutum, finely and densely punctate; on scutellum, moderately coarsely and
very densely punctate; on mesepisternum, anterior to mesepisternal sulcus, moderately coarse and very
densely punctate, distance between punctures less than a puncture diameter; posteriorly, moderately coarsely
and moderately sparsely punctate; on metepisternum, minutely, somewhat inconspicuously punctate,
punctures microtuberculate; on lateral areas of propodeum, somewhat irregular and dull, punctation similar to
that of mesepisternum, slightly denser, with finer punctures scattered among the moderately coarse ones;
basal area of metapostnotum transversely rugose, posterior surface with central longitudinal microreticulate
area; metapostnotum separated from lateral surface of propodeum by a row of wide, deep pits, wider on upper
lateral area, much narrower laterally; on terga, minutely punctate, very sparse (≥ 10 dp) on T1, sparse (≥ 3 dp)
on T2, dense (≈ 1 dp) on T2–T5; on basal area of sterna, impunctate, densely microreticulate, on S2 almost
microrugulose, on S3–S6 shiny microreticulate; on wide apical area of S2–S6, finely but densely punctate.
Surface between punctures, except where otherwise noted above, polished and shiny.
Variation: the irregular impunctate area between the superior eye margin and the lateral ocellus varies in
size among the paratypes and its surface can be relatively smooth but dull (as in the holotype), shiny
microreticulate or smooth and shiny.
Male unknown.
Holotype. Female: “Faz[enda]. Tião Loreano; APA Fernão Dias; S[erra]. Mantiqueira; 13441-39731”
“Gonçalves MG; BRASIL 25/10/2006; F. A. Silveira” and “Actenosigynes mantiqueirensis; Silveira sp. nov.;
HOLOTYPUS” (deposited at UFMG).
Paratypes. Three females, collected with the holotype. Their labels are identical to the ones with the
holotype, except that their top labels, carry one of the following record numbers: 13441-39732, 13441-39733
and 13441-39734 (deposited at UFMG, DZUP and UKANS, respectively).
Etymology. The species is named after the mountain chain, “Serra da Mantiqueira,” located on the
borders of the Brazilian states of Minas Gerais, Rio de Janeiro and São Paulo, and where the type series was
collected.
Distribution. Known only from its type locality, in the municipality of Gonçalves, Minas Gerais state,
Brazil. Probably occurs all along the Serra da Mantiqueira, including also areas in the states of São Paulo and
Rio de Janeiro and, maybe, in other nearby mountain ranges.
Flight activity. The type series was collected in October, in the beginning of the rainy season.

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 Comments. The arrangement of sternal pilosity on both species of Actenosigynes, as dense, wide,
marginal bands, and the way the pollen is held in the holotype of A. mantiqueirensis (Fig. 2c) suggests that
females may actively use them to collect and/or carry pollen from flowers, despite the fact that the scopae in
the hind leg appear to be normal, comparatively to other Paracolletinae, and fully functional.

FIGURE 3. Occurrence records for Actenosigynes fulvoniger Michener (dots) and A. mantiqueirensis sp. nov. (triangle).
Shaded areas are above 1,000 m above sea level. Acronyms for Brazilian states are, from South to North: RS = Rio
Grande do Sul; SC = Santa Catarina; PR = Paraná; SP = São Paulo; MS = Mato Grosso do Sul; MG = Minas Gerais; RJ
= Rio de Janeiro; and GO = Goiás.

Key to the species of Actenosigynes

(Male of A. mantiqueirensis unknown)

1 Mesosoma ferruginous dorsally (Fig. 1a,d); terga entirely black pubescent, tegula ferruginous; pterostigma dark
brown, obscured by black pilosity; basal glabrous area of S2–S5 dull microreticulate (Fig. 1c) ..................................
.................................................................................................................................... A. fulvoniger (Michener, 1989)
- Mesosoma entirely black (Fig. 2a,d); T2–T6 light-yellow pubescent (Fig. 1a,d); tegula blackish brown to black;
pterostigma ferruginous; basal glabrous area of S2–S5 shiny microreticulate (Fig. 2c) ... A. mantiqueirensis sp. nov.

Discussion

Floral relationships of Actenosigynes

Microscopic analysis of the pollen carried by the holotype of A. mantiqueirensis confirmed that this bee
was foraging on flowers of Loasaceae, possibly, Blumenbachia latifolia. The information available on pollen

SYNOPSIS OF THE BEE GENUS ACTENOSIGYNES Zootaxa 2292 © 2009 Magnolia Press · 21
foraging by A. fulvoniger (Wittmann & Schlindwein 1995; Alves-dos-Santos 1999b) plus personal
information supplied by Clemens Schlindwein, Gabriel A. R. Melo and Eduardo A. B. Almeida suggest that
this species is oligolectic, depending on pollen of species of Blumenbachia such as B. eichleri Urb. and B.
catharinensis Urban & Gilg. The discovery of A. mantiqueirensis foraging for pollen on flowers of a species
of this genus suggests that the association between Actenosigynes and Loasaceae is inherited from common
ancestors, including the closest common ancestor from which both known species of the genus evolved. Other
paracolletine bees known to depend on flowers of Loasaceae are Edwyniana sp. and Perditomorpha
rufiventris (which depend on pollen of Loasa), and Perditomorpha frankii and P. pampeana (dependent on
Blumenbachia, including species generally included in Cajophora) (reviewed by Almeida 2006). The lack of
knowledge of the phylogenetic relationships of Actenosigynes prevents the discussion on whether or not
dependence on loasaceous pollen is a consequence of a single host-shift event associated with the common
ancestor of a single clade containing all these paracolletine taxa or whether more than one shift to Loasaceae
as pollen source occurred among these bees

Geographic distributions and biogeography

The occurrence of Actenosigynes fulvoniger seems to be mainly associated with the Araucarian Forest, a
subtropical rain forest dominated by Araucaria angustifolia (Bertol) Kuntze (Araucariaceae) and considered
to be part of the Brazilian Atlantic Rain Forest domain (e.g., Rizzini 1981). The exception is its southernmost
record, a bee collected in the “Morro de Santana,” a 311-m-high hill in the city of Porto Alegre, Rio Grande
do Sul, which is deposited in the MCP. The local environment includes natural dry fields on the hilltop, and
Atlantic Forest, with no representation of the Araucarian Forest. Natural vegetation in humid environments is
now restricted to small spots, due to anthropogenic impact (Penter et al. 2008). All other available records for
A. fulvoniger (Fig. 3), however, are in transitional areas between the Araucarian Forest and the Atlantic Forest
sensu stricto. This includes sites on the “Serra Geral” and the “Serra do Mar,” near the coasts of Rio Grande
do Sul, Santa Catarina and Paraná, as well as in northwestern Rio Grande do Sul and southwestern Santa
Catarina, in the Uruguay river basin.
In northeastern Rio Grande do Sul, A. fulvoniger was said to occur only above 400 m elevation (Alves-
dos-Santos 1999a), in the transition between the Atlantic Rain Forest s.s. and the Araucarian Forest (C.
Schlindwein, pers. comm.), the same environment where the female A. fulvoniger examined here was
collected in the state of Paraná (E.A.B. Almeida, pers. communication). However, a few specimens were
collected below 200 m, in the foothills of the Serra Geral (B. Blochtein, personal information on specimens
deposited in the bee collection of the MCP). Across its range, it is probably found only in or near areas with
humid soil, where its presumed sole pollen source, B. eichleri, grows (Blochtein & Harter 2003). It is not
known, however, if the absence of A. fulvoniger in the core area of the Araucarian Forest is fact or an artifact
due to lack of sampling throughout that region.
The core of the Araucarian Forest spreads through parts of the Brazilian states of Rio Grande do Sul,
Santa Catarina and, especially, Paraná, reaching also the province of Missiones, in northeastern Argentina,
between elevations of 500 m and 1,200 m above sea level, under humid subtropical climate (Castella & Britez
2004). Outside of this core area, the Araucarian Forest occurs north of the Tropic of Capricorn, only on top of
high mountain chains as the Serra da Mantiqueira (where A. mantiqueirensis was found—Fig. 3), surrounded
by a matrix of the montane Atlantic Rain Forest. The Araucarian Forests, on this mountain chain and on
others in the same region, are said to be remnants of Pleistocene communities that expanded northward from
southern Brazil during glaciation events (e.g., Brown & Ab’Saber 1979).
Silveira and Cure (1993) presented several examples of bee species common both at high altitudes and
lowlands in southern Brazil and neighboring countries, and which reappear only on the tops of mountains
north of the Tropic of Capricorn, in southeastern Brazil. Those authors suggested that the geographic range of
those species would have expanded during cool, dry periods, following the northward expansion of the
Araucarian Forest. According to them, those northern, isolated bee populations might be remnants, which
were trapped at the cool mountaintops by climate warming after the last glacial events in the Pleistocene. For

22 · Zootaxa 2292 © 2009 Magnolia Press SILVEIRA

other examples known to date, the time since these major climatic changes was insufficient for significant
morphological differentiation to occur between southern and northern populations, and they have been
considered as conspecific. The presumed sister species now included in Actenosigynes suggest that other
examples of vicariant populations on the mountaintops of southeastern Brazil may belong to sibling species,
different from their morphologically similar southern sisters. Careful morphological analysis of these taxa
and/or analysis of molecular markers, such as DNA barcodes (e.g., Gibs 2009), may help finding character
variation supporting this hypothesis.
It may not be a coincidence that the first speciation event recognized among such remnant bee populations
involves a specialist lineage. Results by Packer et al (2005) and Zayed et al. (2005) suggest that increased
genetic differentiation observed in specialist clades may result from rapid evolution in small isolated
populations limited to environment patches were their host plants occur. Given that all populations of all bee
species were presumably trapped on the mountaintops of southeastern Brazil and isolated from southern
conspecific populations at the same time, by the same vicariant event, it may appear that, in this case,
speciation opportunities should have been equal for both specialist and generalist species. However, it should
be noted that high level of genetic differentiation among populations, as found by Zayed et al. (2005) for
Perditomorpha rufiventris (Spinola), may have allowed speciation to occur in Actenosigynes, even before
climate warming occurred.

Conservation remarks
In Rio Grande do Sul, A. fulvoniger was included in the official state list of threatened species, as
vulnerable (Blochtein & Harter-Marques 2003). The main threat to its conservation was considered to be the
reduction of the populations of its sole pollen source, B. eichleri, due to environmental degradation.
Although no information is available on the conservation status of A. mantiqueirensis or even on its
distribution range, the fact that it is an oligolectic species, requires its situation to be evaluated, since it has
been shown that genetic variation of populations and effective population size of specialist bees tend to be
smaller than those of generalist ones (Packer et al. 2005; Zayed et al. 2005). The need for an evaluation of this
species’ conservation status is reinforced by the fact that the maximum potential range for the species is
probably relatively small and part of it will consist of altitudinal islands emerging from a matrix of forested
lowlands, now largely transformed in pastures and crop fields. Moreover, even in the largest continuous
altitudinal areas, environments suitable for A. mantiqueirensis are probably patchy. This makes these bees
more prone to extinction, due to genetic and demographic reasons. Moreover, dependence on a single or small
number of related pollen sources also increases the bees’ extinction risk, especially when these plants are not
abundant, mass-flowering species as it seems to be the case of Loasaceae in the southeastern mountain tops.

Acknowledgments

The author is indebted to Dr. Eduardo A. B. Almeida, for the donation of a female Actenosigynes fulvoniger
for comparative study; to Roderic B. Martines for the bee photographs appearing in the Figures 1 and 2; to
Diego Hoffmann, for drawing the map in Figure 3; to Dr. João R. Stehmann, for allowing extraction of pollen
from a specimen of Blumenbachia in the herbarium of the Coleções Taxonômicas da UFMG (BHCB); to Dr.
Karin E. B. Meyer and her team for help with pollen slides and pollen identification; to Dr. Clemens
Schlindwein, Dr. Isabel Alves dos Santos, Dr. Betina Blochtein, Dr. Gabriel A. R. Melo and Dr. Eduardo A. B.
Almeida for occurrence data for Actenosigynes fulvoniger, and information on the landscape where they were
recorded; to Dr. Laurence Packer and Dr. Eduardo A. B. Almeida for careful revisions of the text and usefull
suggestions. Thanks are also due to the “Minas Gerais state foundation for research support” (FAPEMIG) for
financial aid (grant APQ 00284-08).

SYNOPSIS OF THE BEE GENUS ACTENOSIGYNES Zootaxa 2292 © 2009 Magnolia Press · 23
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