Scientific Papers
Natural History Museum
The University of Kansas
10 April 2002 Number 25:1-8
Two New Subgenera of Bees in the Genus Centris
(Hymenoptera: Apidae)
By
RicARDo Ayala Library
Estncion dc Biologia Chnnwln, biftitiito dc Biolo^in, UNAM,
Apartado Postal 21, Sail Pntricio,]alisco4S9S0, Mexico JD]_ no 2008
bnrnjas@sen'idor.unam.mx
I-
CONTENTS Ur.,vL.-,oi i Y
ABSTRACT 1
RESUMEN 1
INTRODUCTION 2
Acknowledgments 2
METHODS 2
SYSTEMATICS 2
Aplwriiisia, new subgenus 2
ScJiistheiJiisia, new subgenus 5
LITERATURE CITED 8
ABSTRACT Two new subgenera, Apheniisin and Schistlieniisia, of Coitns (Apinae: Centridini) are de-
scribed and figured. The species of these two new subgenera occur only in South America and were
previously included in the subgenus Mclaccntris.
Key Words: Bees; Apidae; Centridini; Centris; Aplieiuisia; Schistlieiiiisia; Melaeeiitris; neotropics.
RESUMEN Dos nuevos subgeneros Apiiemisia y Schisttieuiisia de Centris (Apinae: Centridini) son
descritas e ilustradas en este trabajo. Las especies de estos dos nuevos subgeneros son sudamericanas
y fueron anteriormente incluias en el subgenero Melaeeiitris.
Palabras Claves: Abejas; Apidae; Centridini; Centris; Apheniisia; Schistheniisia; Melaeeiitris; neotropico.
© Natural History Museum, Tlie University of Kansas ISSN No. 1044-0782
^1
Scientific Papers
Natural History Museum
The University of Kansas
10 April 2002 Number 25:1-8
Two New Subgenera of Bees in the Genus Centris
(Hymenoptera: Apidae)
By
RicardoAyala
Estacion dc Biologia Cliameln, histitiito dc Biologia, LINAM,
Apnrtndo Postal 21, San Patricio, Jalisco 48980, Mexico ^'; " - o j/ggg
baraias@sen'idor.uiMm.inx
CONTENTS L' y
ABSTRACT 1
RESUMEN 1
INTRODUCTION 2
Acknowledgments 2
METHODS 2
SYSTEMATICS 2
Apheiiiisia, new subgenus 2
Schisthemisia, new subgenus 5
LITERATURE CITED 8
ABSTRACT Two new subgenera, Aplictiiisia and Schtstliciiiisia, of Cciitris ( Apinae: Centridini) are de-
scribed and figured. The species of these two new subgenera occur only in South America and were
previously included in the subgenus Mclacciitris.
Kcii Words: Bees; Apidae; Centridini; Centris; Aphcuiisia; Sclnstlicinisia; Mclaccutris; nootropics.
RESUMEN Dos nuevos subgeneros Aphcniisia y Schisthemisia de Centris (Apinae: Centridini) son
descritas e ilustradas en este trabajo. Las especies de estos dos nuevos subgeneros son sudamericanas
y fueron anteriormente incluias en el subgenero Melacentris.
Palabras Chwes: Abejas; Apidae; Centridini; Centris; Apiiemisia; Schisthemisia; Mehiceutris; neotropico.
© Natural Hibtory Museum, The University of Kansas ISSN No. 1094-0782
Scientific Papers, Natural History Museum. The University of Kansas
INTRODUCTION
The tribe Centridini contains nuimTctus large and
robust to moderate-sized mainly neotropical species that
collect oil from flowers, principally Malpighiaceae and
Krameriaceae. The oil is used to provision cells and
sometimes in the construction of the cells in the nest (Neff
and Simpson, 1981). The members of this tribe can be
separated from the remainder of the Apinae by the
presence of a jugal lobe as large as one-third of the vannal
lobe and the absence of an arolium (Michener, 1944;
Stephen cl al., 1969).
The subgeneric classification of the Centridini was
proposed by Michener (1951, 1954) and Moure (1945), and
was later modified by Snelling (1966, 1974, 1984), who
recognized 12 subgenera in the genus Cciitris, nine in the
genus Ep'icliaris, in addition to the genus Ptilotopiis. A
cladistic study by Ayala (1998) indicated that Ccutris is
divisible into three groups, with 12 subgenera. Two of these
subgenera are described as new below. These new
subgenera are included in the group with Melaceniris,
Ptilocenlris, Ptilotopus (no longer considered generically
distinct), and Wagcnkiwchtin.
A classification of all the genera and subgenera of
Centridini is provided by Ayala (1998); papers reviewing
the subgenera of Ccntris are by Michener (1951), Snelling
(1974, 1984), and Ayala (1998).'
The objective of this paper is to publish two new
subgenera of Centridini that I recognized during a study
of the systematics and phylogeny of this tribe, so that the
names can be used before the publication of that work
(Ayala, 1998).
Acknowledgments
I am thankful to the following institutions and
curators, who graciously pro\ ided material used in this
study: Snow Entomological Collection, Division of
Entomology, Natural History Museum and Biodiversity
Research Center, Uni\'ersity of Kansas, Lawrence,
Kansas, M. S. Engel and R. W. Brooks (SEMC); Cornell
University Insect Collection, Ithaca, New York, J. K.
Liebherr and E. R. Hoebeke (CUIC); and American
Museum of Natural History, New York, New York, J.
G. Rozen, Jr. and E. Quinter (AMNH). Charles D.
Michener and Michael S. Engel kindly examined an
early version of the manuscript and provided valuable
criticisms.
Padre j. S. Moure of the Universidade Federal do
Parana in Curitiba, Brazil, independently recognized
the subgenera described below over 40 years ago and
at some point provided unpublished names for them
on labels and more recently in conversations. 1 am
indebted to him for permission (1998) to use and publish
his names, and 1 do so with pleasure to further, with
Padre Moure, the development of a useful classification
of the Centridini.
METHODS
Morphological terminology follows that of Michener
(1944, 2000), Michener and Brooks (1984), and Winston
(1979). In the descriptions the following abbreviations are
used: F, flagellomere; S, metasomal sternum; T, metasomal
tergum. Illustrations were prepared with the aid of a
camera lucida mounted on an Olympus SZH microscope.
Photographic images were produced by Michael S. Engel
using a Microptics ML-1000 Digital Imaging System.
Specimens for study were borrowed from the museums
cited in the Acknowledgments.
SYSTEMATICS
Aphemisia, new subgenus
(Figs. 3-14)
Type species. — Ceiitris pliiiiiipc^' \\ Smith, 1854.
Diagnosis. — Female (Figs. 3, 7, 13, 14): Maxillary
palpus with three free segments. Mandible curved in distal
half, with fourth sharp teeth, two from the rastellum and
two from the pollex; trimma with median denticular
projection. Labrum yellow, wider than long, gently convex.
Clypeus with longitudinal median black mark, its margins
converging apically. Clypeocular distance about as wide
as minimum width of Fl. Lower paraocular area with
yellow mark. Ocellocular space 1.5x diameter of lateral
ocellus. Scutellum with two strong lobes (two strong
elevations) lightly projected backwards (Fig. 13). Basitibial
plate with sharp apex (Fig. 14); secondary plate elevated.
apex slightly projecting beyond attachment to principal
plate. Pygidial plate with pointed apex (Fig. 7); lateral
margin concave; secondary plate triangular with margins
elevated and projected above principal plate; surface of
principal plate concave.
Mali; (Figs. 4-6, 8-12): Similar to female except as
follows: Mandible strongly curved in the apical third (Fig.
6), with three sharp teeth, two from rastellum and one from
pollex. Scutellum with two rounded lobes. Hind tibia and
basitarsus with scopa-shaped pubescence (Fig. 8), large on
the anterior and posterior margins. Hind femur swollen.
Hind basitarsus with single spine on distal, inner margin.
Terga with dark iridescent integument (variable among
specimens). Pvgidial plate absent but tergal margin with
two lobes delimiting a notch.
New Subgenera of Bees
Figs. 1-4. Faces of Ceiitris species. 1. Facial view of female of Cattris (Scliisllifiiiiiu^) flavilnbrii Mocsary. 2. Facial view of male of C. (S.)
flavilabris. 3. Facial view of female of C. (Aphcmisia) pliimipLV Smith. 4. Facial view of male of C. {A.) plumipcs.
Description. — As in the diagnosis with the following
additions: Fem.ale: Maxillary palpus with sparse
pubescence, if present no longer than width of segment,
first free segment longer than second. Mandible without
subapical, internal tooth; acetabular carina reaching base
of fourth tooth; trimmal extension with tooth-like
projection at two-thirds or three-fourths of distance
between internal tooth and mandibular base; rastellum
with two teeth; pollex with tooth (usually very sharp).
Labrum wider than long, with sparse pubescence. Clypeus
little projected, in lateral view gently rounded above, with
discal area gently convex to weakly conca\'e; lower margin
and submargin with weak sulcus, defined by hyaline area;
clypeus with longitudinal yellow line. Fl at least a little
longer than scape. Paraocular area with alveolus-orbital
space longer than diameter of alveolus. Lateral ocellus
SciENTiiic Papers, Natural History Museum, The University oe Kansas
Figs. 5-12. Ceiitris {Aplwiiiisia) phiinipcs Smith, male. 5. Lateral view of galeal base, maxillary palpus, and apex of stipes. 6. Mandible.
7. Pygidial plate. 8. Hind leg. 9. Genital capsule, dorsal aspect. 10. Genital capsule, ventral aspect. 11. Metasomal sternum 7. 12. Metasomal
sternum 8.
below upper tangent of compound eyes. Vertex, in frontal tibia longer than outer spur Scopa of hind tibia consisting
view, at same level as upper ends of eyes. Occipital area of plumose hairs with branches arising from a robust
with plumose pubescence. Scutellum with abundant central rachis. Basitibial plate with secondarv plate
pubescence but not dense, with two tubercles (Fig. 13).
Length of marginal cell less than distance from apex of
cell to wing tip; wing membrane dark brown, translucent.
Fore basitarsus with elaiospathe on anterior margin and
long setae making other comb on anterior margin. Mid
tibia with elaiospathe. Spur of the mid tibia flat and short,
usually as long as Fl and F2 together. Inner spur of hind
ele\ated (Fig. 14), with the margin well defined and apex
insiefe principal plate. Hind basitarsus with posterior
margin of distal internal area with simple robust hairs.
Metasomal terga with metallic iridescence, integument of
T2 punctate but not strongly, slightly shiny; pubescence of
the T2 and T3 very short, not dense; T2-4 without fringes
at distal margins; T4-5 without dense pubescence, T5 with
New Subgenera of Bees
apical fringe of appressed strong dense hairs. Pygidial
plate narrowing to attenuate acute apex (Fig. 7). Secondary
pygidal plate formed with a basal triangle. S4 and S5 with
plumose hairs only on distal margins.
1VI.'\le: Labrum much wider than long. Clypeus with
black macula narrowing from superior margin to inferior
Malar area much longer than clypeus-ocular distance.
Scape shorter than Fl, Fl longer than combined lengths of
F2 and F3. Compound eyes converging dorsally, or slightly
so. Vertex in frontal view below level of superior margins
of eyes. Scutellum with two tubercles. Femora swollen
(Fig. 8). Pygidial plate with posterior margin bilobulate;
S7 with medial projection (Fig. 11); S8 with medial
projection on distal margin elongate with medial part
broad (Fig. 12). Genitalia as in figures 9 and 10.
Comments. — The species of this subgenus were
previously included in the subgenus Mclacentris ( =
Melanocentris sensu §nelling, 1984). Aphemisia was named
by Pe. J. S. Moure on the labels of specimens in the insect
collections of Cornell Uni\'ersity (CUIC) and of the Natural
History Museum, University of Kansas (SEMC). It seems
appropriate to use the name proposed by Pe. Moure for
this taxon, as recommended by Pe. Moure himself (pers.
comm., 1998). Aplicmisia presents many apomorphic
characters. Tlie phylogenetic analysis ( Ayala, 1998) shows
that this new subgenus is sister to Siiiiftlwiiiisin and is part
of the larger group including Mclaccntiis.
Distribution. — The known specimens are from
Amazonas in Venezuela (Territorio del Amazonas),
Ecuador (Guapore; Limoncocha), Brazil (Distrito Federal;
Para), Peru (Azupizu), Bolivia ("Chiapare" [=Chapare]),
French Guiana (Montagne de Pere, Kourou), and Panama
(Canal Zone).
Included species. — Two species are presently
included, Centris (Aphciiiisia) plumipcs Smith, 1854 and
Centris (Aphemisia) n. sp. (SEMC).
Schisthemisia, new subgenus
(Fig. 1-2, 13-23)
Type species. — Centris flavilahris Mocsary, 1899.
Diagnosis. — Female: (Figs. 1-2, 15, 18): Mandible with
strong angle at level of distal third, distal margin broad,
with five teeth, three on rastellum and two on pollex;
trimma with tooth-like projection on distal area. Clypeus
yellow in lower two-thirds or with two light spots.
Paraocular area with lower part yellow. Clypeocular area
as wide as minimum width of Fl. Vertex above level of
upper margin of eyes. Scutellum rounded with two
tubercles; posterior margin vertical, continuous with
metanotum and propodeum. Basitibial plate with
secondary plate ele\'ated, with the margins well projected
along anterior and distal margins, along posterior margin
only slightly projected over surface of principal plate,
distal margin rounded (Fig. 15). Margins of pygidial
plate converging posteriorly, but apex broad, notched
(Fig. 18); secondary pygidial plate rounded, with a small
point (Fig. 18).
Male: Maxillary palpus with three free segments (Fig.
16). Mandible with four teeth, apical tooth acute, other
teeth as in figure 17; acetabular carina ending at base of
more internal tooth and with branches of carina extending
toward other tooth of pollex. Labrum wider than long,
yellow. Clypeus yellow, with black spots toward the upper
margin (some species with upper margin of clypeus black).
Clypeocular distance shorter than minimum width of Fl.
Lower paraocular area yellow. Compound eyes
converging dorsally (Fig. 2). Ocellocular space narrow,
about one-half of ocellar diameter (based upon median
ocellus). Upper margin of the head (frontal view) at level
of upper ocular tangent. Scutellum with two lobes rounded
and projected backwards. Hind leg with femur swollen
and tibia slightly swollen, tibia and basitarsus with
pubescence scopa-like (Fig. 19). Metasomal terga with
integument shiny but not polished, brown or dark brown;
pygidial tergum with well defined plate with posterior
margin bilobulate forming strong notch, the plate
trapezoidal with margin elevated and with little
pubescence.
Description. — Female: Maxillary palpus with three
free segments; first free segment longer than second. Galea
with basal inferior margin with a well-defined lobe.
Mandible with inner subapical area lacking tooth.
Acetabular carina reaching base of fourth tooth; trimma
with a tooth on two- thirds or three-fourths of distance from
inner tooth to mandibular base; rastellum with three teeth;
pollex with two teeth. Labrum wider than long, with little
pubescence. Clypeus projecting or little projecting with
rather flat discal area, discal surface with rounded margins,
with or without central depression; lower margin and
submargin of clypeus with weak sulcus, defined by hyaline
area. Clypeus with black macula on upper margin (Fig.
1). Fl at least slightly longer than scape. Alveolus-orbital
space as long as diameter of alveolus. Lateral ocelli in facial
view below upper tangent of eyes. Vertex in facial view
above upper tangent of eyes (Fig. 1). Mesoscutum with
pubescence plumose, abundant or dense, completely
covering integument. Scutellum with two tubercles (or
lobes) little or slightly elevated. Axillar surface elevated
above level of scutellum. Forewing papillae each ending
with a seta; marginal cell shorter than distance from its
apex to wing tip; wing membrane translucent, ochre or
dark ochre, without or with weak iridescence. Fore
basitarsLis with elaiospathe on anterior margin and long
hair forming comb on posterior margin. Mid tibia with
elaiospathe; spur flat and short, as long as Fl or combined
Scientific Papi;rs, Natural Hisiory Musuum, Thi; Univi-rshy oi Kansas
I igs. 1.) 15. Mesosomal tharactfrs of Ci'/ifnV species. 13. Scutelhini of female Ci'/ifiis (/\;>/;('m/s;ij)f)/i/mi;)t'.'i Smith. 14. IJiisitibial pl.Ue oi l.
(/I.) phimipes. 15. Basitibial plate oi C. {Scliistlwinisia) fltwiltihrif Mocsar\-.
length of Fl and F2. Inner hind tibial spur pectinate; inner
and outer spurs similar in length. Scopa on hind tibia with
plumose hairs emerging from strong, central rachis;
basitibial plate with secondary plate elevated, its margin
defined, without yellow markings; secondary plate with
apex rounded, inside margin of principal plate. Hind
basitarsus with simple, robust hairs, with bent apices, on
the distal internal area of posterior margin. Terga without
metallic iricHescence; T2 with weak punctation not strong,
weaklv shin\'; T2 and T3 with short but not dense
Nbw Subgenera of Bees
Figs. 16-23. Ccntris [Schi^^thcniiiia) flnvilnbris Mocsary, male. 16. Lateral view of galeal base, maxillary palpus, and apex of
stipes. 17. Mandible. 18. Pygidial plate. 19. Hind leg. 20. Genital capsule, dorsal aspect. 21. Genital capsule, ventral aspect. 22.
Metasomal sternum 7. 23. Metasomal sternum 8.
pubescence; T2-4 without apical fringes of hair; T5 with
fringe on margin of strong appressed hairs; pygidial plate
with margins converging (Fig. 18), apex truncate with
strong median notch; secondary pygidial plate rounded
(Fig. 18); S4-5 with plumose hairs only on distal margins,
not obscuring surface; S6 with apex rounded, short, or
ending in a tuft of hairs.
Male: Similar to female. Labrum broader than long.
Malar space short, as long as clypeocular space. Scape
shorter than Fl; Fl longer than combined length of F2 and
F3. Compound eyes converging dorsally (Fig. 14). Vertex
at level of upper ocular tangent (Fig. 2). Hind femur
swollen; hind basitarsus with posterior margin lacking
carina and denticulate projection. Pygidial plate with apex
8
3 2044 072 228 760
Scientific Papers. Natural History Museum, The University oi- Kansas
bilobulate. S7 slender with elongate medial projection with
strong notch at apex (Fig. 22); S8 with medial projection of
distal margin elongate, widened in mid distal part (Fig.
23); genitalia as in figures 20 and 21.
Comments. — The name of this subgenus was
suggested by Pe. J. S. Moure more than 40 years ago
when he independently recognized this subgenus as
new but never published on it. I found it on labels of
specimens in CUIC and SEMC, and publish it here with
his permission, given in 1998. See notes under
Aphemisia.
The species of this subgenus were pre\'iously included
in Mcliicentris (= Melanocentris sensu Snelling, 1984). This
is the sister subgenus of Aphemisia. The subgenus
Schi:>thi'nii^ia can be recognized by the shape of the
mandibles, scutellum, hind tibia, and pygidium. The male
can be separated from other Centris by the shape of the
clypeus and scutellum.
Distribution. — The only known specimens of this
subgenus were collected in Brazil (Amazonas, Guapore)
and Bolivia (La Paz, Beni).
Included species. — Centris (S.) flnvilabris Mocsary, 1899
and C. (S.) fusciventris Mocsarv, 1899.
LITERATURE CITED
Ayala, R. 1998. Sistematica supraespecifica de las abejas de la tribu
Centridini (Hvmenoptera: Apoidea). Doctoral Dissertation in
Sciences (Zoology), Universidad Nacional Autonoma de Mexico,
iv+280 pp.
Michencr, C. D. 1944. Comparative external morphology, phylogeny,
and a classification of the bees. Bulletin of the American Museum
of Natural Histor\' 82:151-326.
Michener, C. D. 1951. Subgeneric groups of Hcinitna (Hymenoptera,
Apidae). Journal of the Kansas Entomological Society 24:1-11.
Michener, C. D. 1954. Bees of Panama. Bulletin of the American Museum
of Natural History 104:1-176.
Michener, C. D. 2000.' The Bees of the Worhl. Baltimore, Maryland: The
Johns Hopkins University Press, xiv+|ll+913 pp.
Michener, C. D., and R. VV. Brooks. 1984. Comparative study of the glossae
of bees. Contributions of the American Entomological Institute 22:1-
73.
Moure, J. S. 1945. Notes sobre os Epicharitina (Hvmenoptera, Apoidea).
Revista de Entomologia [Rio de Janeiro] 16:293-314.
Neff, J. L., and B. B. Simpson. 1981. Oil-collecting structures in the
Anthophoridae (Hymenoptera): Morphology, function, and use in
systematics. Journal of the Kansas Entomological Society 54:95-123.
Snelling, R. R. 1966. The taxonomy and nomenclature of some North
American bees of the genus Centris with descriptions of new species
(Hymenoptera: Anthophoridae). Contributions to Science, Los
Angeles County Museum of Natural History 112:1-33.
Snelling, R. R. 1974. Notes on the distribution and taxonomy of some
North American Coifr/'s (Hymenoptera: Anthophoridae).
Contributions to Science, Los Angeles Countv Museum of Natural
History 259:1^1.
Snelling, R. R. 1984. Studies on the taxonomv and distribution of
."Xmerican eentridine bees (Hymenoptera: Anthophoridae).
Contributions to Science, Los .Angeles Countv Museum of Natural
History 347:1-69.
Stephen, W. P., G. E. Bohart, and P. R Torchio. 1969. The Biohgi/ and
External Morpholcgi/ of Bees. With a S\/nop'sis of the Genera of
Nortliwesteni America. Corvallis, Oregon: Agricultural Experimental
Station, Oregon State University [2J+140 pp.
Winston, M. L. 1979. The proboscis of the long-tongued bees: A
comparative study. University of Kansas Science Bulletin 51:631-
667.