Nesting Biology of a New High Andean Bee, Anthophora walteri

Gonzalez (Hymenoptera: Apidae: Anthophorini)

Author(s): Victor H. Gonzalez and Francis Chavez
Source: Journal of the Kansas Entomological Society, 77(4):584-592. 2004.
Published By: Kansas Entomological Society
DOI: http://dx.doi.org/10.2317/E-8.1
URL: http://www.bioone.org/doi/full/10.2317/E-8.1

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 JOURNAL OF THE KANSAS ENTOMOLOGICAL SOCIETY
77(4), 2004, pp. 584–592

Nesting Biology of a New High Andean Bee, Anthophora walteri

Gonzalez (Hymenoptera: Apidae: Anthophorini)
VICTOR H. GONZALEZ1 AND FRANCIS CHAVEZ2

ABSTRACT: Males and females of Anthophora walteri Gonzalez, new species, from the Eastern
Andes of Colombia are described and illustrated. The pupa is described, and information on the nesting
behavior, natural enemies and host plants of this species are also given. Females of A. walteri can be
distinguished from other Neotropical Anthophora by the predominantly white pubescence on the head
and mesosoma, and orange setae on the metasoma which are denser and posteriorly-directed on apical
margins of terga, thus forming apical bands. Males can be recognized by the sparse pubescence on the
metasoma and the translucent apical margins of terga one to six. Bees nested in flat ground or small
banks in very hard soils of semiarid and highly disturbed areas above 2500 m; nest entrances lacked
turrets and cells were arranged in linear series as reported for other Mystacanthophora species. Pollen
taken from nest cells was almost exclusively (.95%) from a single species of Lamiaceae (Salvia
bogotensis). Adults of A. walteri fly from about October at least through July. Presumably immatures
spend most of the year as postdefecating larvae and the period of adult emergence corresponds with
blooming of S. bogotensis and the second peak of maximum rain in October.
KEY WORDS: Anthophora, taxonomy, Salvia, Andes, Colombia, nesting behavior

RESUMEN: Hembras y machos de Anthophora walteri Gonzalez, especie nueva, de la cordillera

Oriental de Colombia son descritos e ilustrados. La pupa también se describe y se presenta
información sobre la biologı́a de nidificación, enemigos naturales y plantas hospederas de esta especie.
Las hembras de A. walteri pueden distinguirse de otras Anthophora neotropicales por la pubescencia
predominantemente blanca en la cabeza y mesosoma, y las setas anaranjadas sobre el metasoma, las
cuales son densas y están dirigidas posteriormente sobre las márgenes apicales de los tergos y por lo
tanto formando bandas apicales. Los machos pueden reconocerse por la pubescencia esparcida del
metasoma y las márgenes apicales translucidas de los tergos uno a seis. Las abejas nidifican en
superficies planas o en pequeños barrancos de suelos muy duros, en áreas semiáridas y altamente
perturbadas por encima de los 2500 m; las entradas de los nidos carecen de torres y las celdas están
dispuestas en series lineares como ha sido registrado para otras especies del subgénero
Mystacanthophora. Las muestras de polen tomadas de las celdas contenı́an casi exclusivamente
(.95%) polen de una sola especie de Lamiaceae (Salvia bogotensis). Los adultos de A. walteri vuelan
desde alrededor de Octubre, por lo menos, hasta Julio. Probablemente los inmaduros pasan la mayor
parte del año como larvas posdefecantes y el periodo de la emergencia de los adultos corresponde con
la floración de S. bogotensis y el segundo pico de máxima lluvia en Octubre.
PALABRAS CLAVE: Anthophora, taxonomı́a, Salvia, Andes, Colombia, comportamiento de
nidificación

Introduction
Anthophora, a genus of solitary bees (Apidae: Anthophorini), contains more than 300
species and 14 subgenera worldwide. They are particularly abundant in xeric climates and
scarce in tropical forests, but are often found in tropical montane environments (Brooks,
1988). The great majority of the Anthophora species that occur in the neotropics belong to
the subgenus Mystacanthophora (Michener, 2000); unlike their temperate congeners little
is known about their biology.
1
Entomology Program, Department of Ecology and Evolutionary Biology, Snow Hall, 1460 Jayhawk
Boulevard, University of Kansas, Lawrence, Kansas 66045-7523, e-mail: vhgonza@ku.edu
2
Instituto de Ciencias Naturales, Universidad Nacional de Colombia, Santa Fé de Bogotá, Colombia, A.A.
7495, e-mail: francesca@ciencias.unal.edu.co
Accepted 6 November 2003; revised 21 April 2004
Ó 2004 Kansas Entomological Society
VOLUME 77, ISSUE 4 585

In this paper we provide information on the nesting biology, natural enemies and host
plants of a new Mystacanthophora species that occurs at high altitudes in the Andes. The
novelty of this species was first recognized by Dr. Robert Brooks who urged one of us
(VG) to describe and illustrate it.

Materials and Methods

TAXONOMIC DESCRIPTION: The morphological descriptions and illustrations were made
using an Olympus SZ microscope. The description format and morphological terminology
follow Brooks (1988) and terminology for surface sculpturing follows Harris (1979). Setal
length is given relative to the diameter of median ocellus. The abbreviations F, S, OD, and
T are used for flagellomere, metasomal sternum, ocellar diameter, and metasomal tergum,
respectively. Type specimens, immature stages and nest structures are deposited in the
following institutions:
IAVH Instituto Alexander von Humboldt, Villa de Leyva, Boyacá, Colombia (J. E.
Castillo).
SEMC Snow Entomological Collection, Entomology Division, Natural History
Museum, University of Kansas, Lawrence, Kansas, 66045-7523, USA (Z. Falin).
LABUN Laboratorio de Investigaciones en Abejas, Universidad Nacional de
Colombia, A.A. 144909, Santa Fé de Bogotá, Colombia (Guiomar Nates-
Parra).
NMNH National Museum of Natural History, Smithsonian Institution, Washington
D.C., USA.
UNCB Museo de Historia Natural, Instituto de Ciencias Naturales, Universidad
Nacional de Colombia, A.A. 7495, Santa Fé de Bogotá, Colombia (E. Flórez).

STUDY SITE AND NEST EXCAVATIONS: Observations were made in the area of Mondoñedo,
between Mosquera and Bojacá (Departamento of Cundinamarca), Colombia, at 2500–
2700 m. This is a semiarid area that is highly disturbed by cattle ranching, gravel
extraction and waste dumping. The soil is very compacted, shallow, and strongly eroded,
with a pH of 6.0. The median temperature is approximately 138C (IGAC, 1996). The
vegetation of this area is mainly characterized by sparse, low gramineas and shrubs usually
less than 1 meter tall. Plants such as Senecio spp. (Asteraceae), Solanum spp. (Solanaceae),
Opuntia schumannii Weber ex Berger, O. ficus-indica (Linnaeus) (Cactaceae), and Salvia
bogotensis Benth (Lamiaceae) are predominant in the area. The rainy season is bimodal
with a maximum in April–June and another in September–October, never reaching beyond
100 mm of precipitation monthly (IGAC, 1996).
Nest excavations were made on 16 December 1999; 13 August, 10 September, 11, 12, 13
October, 25 November, and 26 December 2000. Nests were excavated using a geological
pick and knife. Adults and immatures were fixed in Kahle’s solution and pollen from cells
was transferred to plastic trays for identification. Mean values are given with standard errors.
Most of the paratypes were from other localities but from environments and within the
season indicated above.
PALYNOLOGICAL ANALYSIS: Pollen samples from cells were treated and preserved
following Erdtman’s (1986) acetolysis method and then mounted on glass slides. In order
to estimate the relative percentage of pollen of each plant species, we randomly selected
one slide among several slides that had been prepared for each cell. Then, pollen grains
were identified and counted by sampling all grains found in a row until 500 grains were
586 JOURNAL OF THE KANSAS ENTOMOLOGICAL SOCIETY

reached. Pollen was identified by comparison with a palynological reference collection

made from plants collected at the study area. Examination of pollen was done under the
microscope (Leitz DIALUX 22 EB) at 403 and 1003. Pollen samples are deposited in the
Palynological collection of the Instituto de Ciencias Naturales, Universidad Nacional de
Colombia, Santa Fé de Bogotá.

Results
Anthophora (Mystacanthophora) walteri Gonzalez, new species
(Fig. 1)
DIAGNOSIS: This species belongs to the A. pilosifrons species-group, whose members
have an almost flat clypeus bearing stout hooked setae and a transverse row of blunt,
spine-like projections (pecten) across the base. Females of A. walteri can be easily
distinguished from any other Neotropical Mystacanthophora by the predominantly white
pubescence on the head and mesosoma, and the long, minutely branched, orange setae
curving anteriorly on the metasoma, these orange setae becoming denser on the apical
margins of T1–T4 and posteriorly-directed, forming apical bands. Males can be recognized
by the sparse pubescence on the metasoma, the translucent apical margins of T1–T6, and
the shapes of the genital capsule and associated sterna.
DESCRIPTION: Female. Body length 12.3–15.0 mm (x ¼ 13.5 6 0.4, n ¼ 5); forewing
length 9.5–10.4 mm (x ¼ 9.8 6 0.2, n ¼ 5). Inner orbits diverging above; shortest distance
between compound eyes about as long as eye; malar space short, about half as long as
maximum diameter of F3; head wider than long; clypeus flattened in profile, projecting
about 0.6 compound eye width in lateral view; F1 equal in length to next 3.2 flagellomeres
taken together, 0.8 as long as scape and 1.6 times as long as last flagellomere, F2 0.3
length of F1, F3 slightly longer than F2 and shorter than F4–F9 individually; ocellocular
distance as long as interocellar distance; distance from median ocellus to lateral ocellus 0.4
times the ocelloocular distance. Marginal cell length 0.8 times the distance from its apex to
wing tip; hind wing with cu-v 0.7 times length of second abscissa of MþCu; vein M about
2.1 times as long as second abscissa of MþCu. Coloration. Mainly black, except apex of
mandible, legs and ventral surface of metasoma dark reddish-brown; tegula brown; wings
slightly dusky and veins dark brown to black. Pubescence. Head mainly white, with black
setae intermixed on vertex, ocellar area and superior paraocular area; clypeal pecten
consisting of a row of 15–17 stout, simple, apically hooked, pale ferruginous setae; rest of
clypeus, supraclypeal area and just below antennal sockets covered with less stout, apically
hooked, simple, pale, ferruginous setae, stouter than plumose setae on the rest of head;
labrum covered with pale, hooked, simple setae, less stout and shorter than those on
clypeus; paraocular area and lateroapical area of clypeus covered with long (.3 OD),
white, minutely branched setae, longest on inferior gena. Scutum, scutellum and
metanotum and margins of tegula densely covered (integument still visible) with long
(about 3 OD), branched, pale gray setae, mixed with long black setae; remaining areas of
thorax with white setae as on gena, mesepisternum mixed with black setae. Pubescence on
legs white except as follows: inner surface of front tibia and tarsus with ferruginous setae;
middle and hind tibiae and tarsi with dark brown to black setae; dark brown setae
bordering basitibial plate and along posterior margin of hind tibia; penicillus dark brown.
Apical margin of T1 and all T2–T4 covered with minutely branched, long (about OD),
semierect, orange setae curving anteriorly; apical margins with denser, posteriorly directed
setae, forming apical bands. Lateroventral parts of all terga with tufts of long (.3 OD),
VOLUME 77, ISSUE 4 587

Fig. 1. Male of Anthophora (Mystacanthophora) walteri n. sp. A ¼ S5, ventral view; B ¼ S6, ventral view; C ¼
S7, ventral view; D ¼ S8, ventral view; E, F ¼ Genital capsule, dorsal (left), ventral (right) and side views; G ¼
inner surface of hind femur and trochanter; H ¼ face showing integumental color. Stippling indicates black,
remainder whitish cream; pubescence omitted (Scale: A, B, E–H ¼ 1 mm; C, D ¼ 0.5 mm).
588 JOURNAL OF THE KANSAS ENTOMOLOGICAL SOCIETY

branched, white setae, progressively increasing in length toward apex of metasoma. T1–T5
with sparse, long (2 OD), erect, simple, dark brown to black setae arising among orange
setae, such dark setae longest and sparser on T1, denser on sides of T3–T4; sides of T5
with white branched setae, medioapically with long (,2 OD), stouter and mostly branched
black setae like those bordering pygidial plate of T6. Metasomal sterna with long (.2
OD), branched, white setae on apical margins, except S5 and S6 where setae longer, stiffer
and dark brown. Punctation. Integument microareolate between punctures. Labrum
strongly punctate, almost areolate-rugose; clypeus and supraclypeal area with strong
punctures, separated by 1–2 puncture widths; above antennal sockets to ocelli as on
paraocular area with punctures almost contiguous, finer on latter. Mesosoma with fine
punctures, almost contiguous; metasoma with faint punctures, separated by 2–3 puncture
widths, except denser and stronger on T5.
Male. As described for female except: body length 11.5 mm; forewing length 8.2 mm.
Malar space shorter than in female, length about one third maximum diameter of F3;
clypeus more protuberant in profile, about 0.8 times width of compound eye in lateral view;
F1 length equal to next 1.5 flagellomeres taken together, 0.6 as long as scape and about as
long as last flagellomere, F2 0.6 length of F1, F3 length about 1.5 times greater than F2 and
about as long as F4 and F5 individually, F6–F10 each slightly shorter than F3; distance
from median ocellus to posterior ocellus about half of ocellocular distance. Marginal cell
length 0.8 times the distance from its apex to wing tip; hind wing with cu-v half of length of
second abscissa of MþCu. Middistarsal and midtarsal brush absent; circular tuft of long
black setae on basal inner surface of hind femur as in Fig. 1G. S5–S8 and genital capsule as
in Fig. 1A–F. Coloration. Whitish-cream maculations on face as shown in Fig. 1H.
Mediotarsi and distitarsi of all legs dusky ferruginous, contrasting with reddish-brown rest
of legs; wings hyaline; apical half of T1–T6 reddish-brown, vanishing toward margin;
apical margin translucent and slightly yellow, especially on T1. Pubescence. Mainly white;
paraocular area, apicolateral area of clypeus and labrum with long (3 OD), minutely
branched setae; rest of clypeus and labrum with short (,OD), sparse, semidecumbent setae,
longer on labrum; vertex, ocellar area and paraocular area intermixed with black setae,
sparser than in female. Scutum, pronotal lobe, tegular area, scutellum and metanotum
mixed with long (.3 OD) black setae. Pubescence on legs white, except as follows: inner
surface of front tibia and tarsus, middle tarsus and hind distitarsus with ferruginous setae;
inner surface of hind tibia and basitarsus with dark brown to black setae. Metasomal
pubescence longer and sparser than in female. Apical margins of T1–T4 with denser,
posteriorly directed yellowish setae, forming apical bands; apical margins of T5–T6 nearly
bare; T1–T6 sparsely covered with long (2 OD), branched, yellowish setae, denser and
longer on T1; T2–T5 with long (2 OD), simple, erect dark brown setae sparsely mixed with
the yellowish to white setae, white setae denser on T6 and T7, especially laterally. Apical
margin of S1 and apicolateral areas of S3–S5 with long (2 OD), branched, white setae; disc
of S3 densely covered with short, decumbent white branched setae. Medioapical margins of
S4–S5 with dense ferruginous setae. Punctation. Labrum and clypeus with faint punctures,
separated by about a puncture width on latter. Metasoma with fine punctures separated by
1–5 puncture widths, denser and stronger on T7.
Pupa. As described for A. linsleyi Timberlake (Michener, 1954), except for the presence
of a small tubercle on disc of tegula and basal outer surface of hind tibia, large tubercle
weakly divided medially on middle line of metanotum (lower in females), and very small
tubercle on each axilla (more evident in males). T2–T4 of both sexes with medially
interrupted subapical row of spicules, 11–16 spicules on each side; T5 of female with 8–11
VOLUME 77, ISSUE 4 589

small spicules, almost forming two subapical rows on each side of tergum, smaller than on
previous terga; smallest spicules (6–8) on each side of T6. T6 of male with 10–12 spicules,
smallest on T7 (6–8).
HOLOTYPE: COLOMBIA: CUNDINAMARCA: Mosquera, Vereda Balsillas, Hacienda
Mondoñedo, X-11-1999, 2700 m, M. Ospina [LABUN].
PARATYPES: COLOMBIA: CUNDINAMARCA: 2 $$: Bogotá, IX-29-1974, 2600 m
[label erroneously says 20 m] G.E. Bohart; 1 #, 4 $$, Bogotá, VI-21-1936 [two $$ with
no date], 2600 m, J. Bequaert; 2 $$, 2 ##: Subachoque, VII-26-1936, 2836 m, J.
Bequaert; 1 #, same but no date, 1936; 1 $, 1 #, 4 km from Mosquera, XII-2-1988, 2000
m, R. Pardo and G. Castaño; 1 $, 1 #, Alto de Mondoñedo, XI-26-2000, 2710 m, M.
Ospina; 1 $, X-8-2000, 2400 m, V. Gonzalez; 2 ##, Mosquera, Hacienda Pueblito
Español, XII-28-1981, R. Ospina; 1 $, Tocaima, about 25 km NE from Girardot, II-17-
1948, 600 m, L. Richter. BOYACÁ: 1 $, Tunja, Santuario de Fauna y Flora de Iguaque,
Chaina, I-11-1992, 2580 m, A. Amarillo [IAVH, LABUN, NMNH, SEMC and UNCB].
ETYMOLOGY: The specific name is a patronymic honoring Walter Van Sickle, a biologist
who understands the importance of nature, in gratitude for his inspirational efforts towards
the conservation of the biodiversity on our planet.
COMMENTS: Anthophora paranensis Holmberg is probably the closest relative of A.
walteri as is suggested by the similarity of the male genitalia and associated sterna. Males
of only these two species share a well-defined nose-like medioapical tubercle on the S6
which is more pronounced in A. walteri (Fig. 1B). However, the apical margin of S6 is
straight in A. walteri as opposed to a shallow, broad emargination found in males of A.
paranensis and in other North American species of Mystacanthophora. Besides the
characters given above and in the diagnosis, males of these species differ by the
pubescence on S5–S6, pubescence on lateral lobes and apical margin of S7, which is
longer and denser in A. paranensis. In A. walteri, the lateral carinal process of the
gonocoxite (terminology of Brooks, 1988) is short relative to its maximum width and
gonostylus length (about 1.4 times as long as wide and 0.7 as long as gonostylus). In A.
paranensis the lateral carinal process is longer, about twice as long as wide and 0.8 as long
as the gonostylus. Also, in the male the basal dense patch of black setae on the inner
surface of the hind femur is large and rounded in A. walteri; it is smaller and forming
a band next to the trochanter-femur articulation in A. paranensis.
Two females (not designated as paratypes) from Mérida, Venezuela with the following
labels: ‘‘Viena, 1883’’ and ‘‘Timotes, 1 June 1976, A. S. Menke and D. Vicent coll’’,
deposited at SEMC, closely resemble females of A. walteri. There are, however, major
differences from A. walteri such as the darker pubescence on mesosoma and legs, the
hairier and more fulvous metasomal terga, and more robust female gonostyli. They could
be the same species but examination of the male is required to confirm their identity.
NEST ARCHITECTURE: Nests were found forming aggregations mostly in flat ground or
gently sloping surfaces. The nesting area was usually barren and well exposed to the sun.
Numerous hardened cells and remains of cells from older generations were commonly
exposed in the study area as a result of erosion. The density of nests in aggregations ranged
from 4.6 to 65 nests per square meter (x̄ ¼ 19.6 6 6.9, n ¼ 8 aggregations). The inter-nest
distance in those aggregations ranged from 0.9 to 109 cm (x̄ ¼ 20.4 6 19.2, n ¼ 88). Nest
entrances were almost circular, ranged from 6 to 12 mm in diameter (x̄ ¼ 7.9 6 0.2, n ¼
53), and lacked turrets (Fig. 2). When nests were active, irregular tumuli approximately 2.4
cm in length (60.4, n ¼ 20) and 1.8 cm in width (60.3, n ¼ 20) extended downward from
the nest entrances. Burrows were rough, not lined, and on average 7.8 mm in diameter (6
590 JOURNAL OF THE KANSAS ENTOMOLOGICAL SOCIETY

Fig. 2. Schematic illustration of a nest of Anthophora (Mystacanthophora) walteri n. sp., a ¼ tumulus

(Scale ¼ 1 cm).

0.1, n ¼ 42) and 107.7 mm in length (67.3, n ¼ 20), meandering downward from the
surface, usually with the first centimeters from the entrance running parallel to the surface.
Nests were shallow with the maximum depth (measured from the nest entrance to the
bottom of the innermost cell) ranging from 52 to 111 mm (x̄ ¼ 71, 63.6, n ¼ 16). Cells
were constructed in linear series and separated from each other by plugs of soil. Cells were
excavated in the soil and were not separable from the substrate. Cells varied from almost
horizontal to nearly vertical and they were constructed at different depths ranging from
15 to 92 mm (x̄ ¼ 65.5, 62.9, n ¼ 33). The number of cells per nest ranged from 1 to 13
(x̄ ¼ 3.4, 60.6, n ¼ 99).
Cells resembled those of other Anthophora bees, being ovoid, wider posteriorly and
narrowing toward the anterior end; on average they were 7.6 mm in diameter (60.2, n ¼
51), 13.4 mm in length (60.3, n ¼ 25) and 6 mm in cell entrance diameter (60.3, n ¼ 13).
The inner wall was smooth and shiny, presumably lined with Dufour’s glands secretions.
The cell cap was on average 6.6 mm in diameter (60.1, n ¼ 12) and 1.8 mm thick (60.07,
n ¼ 13), concave and convex on its outer and inner surfaces, respectively; the inner surface
had a small pore. Provisions were liquid and had a strong cheesy odor. Eggs were floating
on the surface. Defecated larval feces were usually smeared as a continuous whitish layer
occupying about three quarters of the cell length, but were sometimes weakly
distinguished as individual flattened bands about 0.06 mm wide (60.01, n ¼ 10). Some
females used old nests, as evidenced by the facts that the first centimeters of the tunnel in
some of them were clearly old, followed by newly excavated tunnels, and such nests had
a mixture of new and old cells.
HOST PLANTS: Palynological analysis of the brood provisions of 58 cells from 15 different
nests showed that 90–99% (x̄ ¼ 95.4 6 0.3) of the pollen within each cell was from
VOLUME 77, ISSUE 4 591

a single plant species, Salvia bogotensis (Lamiaceae); an unidentified Asteraceae species

accounted for 0.2–4.4% (x ¼ 1.35 6 0.1) of the pollen in all samples. Other 31
unidentified types of pollen grains and spores were present in trace amounts (
1% of the
total per cell) and may have been contaminants.
Observations of pollen collecting behavior were made on two occasions. Pollen from
flowers of S. bogotensis was initially collected on the stout hooked setae of the clypeus and
supraclypeal area when the females visited a flower. After visiting several flowers, the
females flew to a tiny branch of S. bogotensis and gripped it firmly with their mandibles.
Then, they raised their body and used the forelegs to remove pollen from the face while
actively moving the body laterally and passing the pollen to the middle legs and finally, to
the hind tibial scopae. Both females and males of A. walteri also visited S. bogotensis
flowers for nectar.
NATURAL ENEMIES: Four fly larvae of the family Tachinidae (Diptera) were found feeding
on the carcass of an unemerged female on 26, December 2000. This nest also had two
more cells, each one containing a predefecating bee larva.
SEASONAL CYCLE: On 13 August, nest entrances were closed. On this day, about 25 nests
were excavated and 56 postdefecating larvae and 14 pupae ranging from brown and black
compound eyes, and four totally colored pupae were found. Four of these pupae (;22%)
with brown eyes were females. In the middle October most of the nests remained inactive
but males were first seen on S. bogotensis. Both female (n ¼ 4) and male colored pupae
(n ¼ 5) were present in three inactive nests excavated at this time. By November, both
adult males and females were flying in Mondoñedo. Although no adult bees from
Mondoñedo have yet been observed or collected between March and October, females and
males were collected on September 28, June 21, and July 26 from other localities (see data
for paratypes).

Discussion
The biology of Neotropical Anthophora is poorly known. Only brief accounts on the
biology for four of the 12 known species that occur in the neotropics are available (Brooks,
1988). The adults of Neotropical Mystacanthophora fly from around October through July;
they nest in very hard soils, either in flat ground or banks; their nest entrances lack turrets
and cells are arranged in linear series (Janvier, 1955; Michener and Lange, 1958; Alayo,
1982; Brooks, 1988, 1999). However, a trifid arrangement of cells, not in linear series, has
been described for A. incerta (Spinola) by Claude-Joseph (1926). Misinterpretations of the
nest architecture in aggregations of nests can sometimes occur because of the proximity of
nests and may be the case in Claude-Joseph’s (1926) account. Nevertheless, variations in
the nest architecture among populations are also likely to occur in Anthophora, as reported
for A. urbana Cresson in North America (Torchio and Trostle, 1986).
From the surveyed nests, apparently A. walteri is a protandrous species and spends most
of the year as postdefecating larvae. The emergence period corresponds with the blooming
of S. bogotensis and the second peak of maximum rain in October. Although a systematic
sampling throughout the whole year is necessary, the palynological analysis showed that
the study population of A. walteri specializes on pollen of S. bogotensis. Thus, it is likely
that the A. walteri life cycle is somewhat synchronized with the maximum blooming
period of S. bogotensis.
The presence of apically hooked setae on the clypeus and supraclypeal area of the female is
apparently related to pollen collection from Salvia. Other South American Mystacantho-
phora species (e.g., A. paranensis Holmberg) with hooked setae on the face have also been
592 JOURNAL OF THE KANSAS ENTOMOLOGICAL SOCIETY

collected on Salvia as indicated by the label data of several specimens at SEMC, while other
species that lack them such as A. urbana are, in contrast, highly polylectic (Mayer and
Johansen, 1976). Unrelated bee species even from different families [e.g., Andrena
(Andrenidae), Caupolicana (Zikanapis) (Colletidae), Trachusa (Ulanthidium), Megachile
(Megachilidae), one species of Amegilla, two species of Habropoda, many species of
Deltoptila, Tetraloniella (Pectinapis), and Tetraloniella s. str. (Apidae)] that also exhibit
such hooked setae are specialized pollen collectors from Salvia (Brooks, 1988; Michener,
2000; Michener et al., 2003). This indicates independent origins for such bee-flower
associations. Besides the observations reported here, nothing is known about the foraging
behavior of these bees on Salvia and a comparative study across taxa would be interesting to
learn if all of them converged to the same repertoire of behaviors.

Acknowledgments
This paper is dedicated to the memory of Dr. Howard E. Evans. We gratefully
acknowledge the financial support of Ideawild and the William J. Bell Fund of the
Program in Entomology of the University of Kansas (to VG). We thank Fagua Alvarez,
Jaime Pinzón and Alex Sabogal for their help in the field; Dr. F. Koch from Zoologisches
Museum an der Humboldt-Universität zu Berlin for lending us the holotypes of A.
versicolor and A. albiceps for study; Chinguen Wang, Molly Rightmyer and Charles
Michener for reading and offering valuable comments on the manuscript; George Byers
for the fly identification and Robert Brooks who encouraged us to study this bee. This is
contribution Nr. 3348 of the Division of Entomology, Natural History Museum and
Biodiversity Research Center, University of Kansas.
Literature Cited
Alayo, S. R. 1982. Observaciones en himenópteros cubanos. 1. Apoidea y Sphecoidea. Poeyana 241:1–7.
Brooks, R. W. 1988. Systematics and phylogeny of the anthophorine bees (Hymenoptera: Anthophoridae;
Anthophorini). University of Kansas Science Bulletin 53(9):436–575.
Brooks, R. W. 1999. Bees of the genus Anthophora Latreille 1803 (Hymenoptera, Apidae, Anthophorini) of the
West Indies. Tropical Zoology 12:105–124.
Claude-Joseph, F. 1926. Recherches biologiques sur les hyménoptères du Chili (Mellifères). Annales des
Sciences Naturelles, Zoologie 9(10):113–268.
Erdtman, G. 1986. Pollen Morphology and Plant taxonomy (An introduction to Palynology). E. J. Brill, Hafner
Publishing Company, Amsterdam. 553 pp.
Harris, R. A. 1979. A glossary of surface sculpturing. Occasional Papers in Entomology, California Department
of Food and Agriculture 28:1–31.
IGAC, 1996. Diccionario Geográfico de Colombia. Instituto Geográfico Agustı́n Codazzi, Bogotá, Colombia.
Janvier, H. 1955. Le nid et la nidification chez quelques abeilles des Andes tropicales. Annales des Sciences
Naturelles, Zoologie 17(11):311–349.
Mayer, D. F., and C. A. Johansen. 1976. Biological observations on Anthophora urbana urbana Cresson
(Hymenoptera: Anthophoridae). Pan-Pacific Entomologist 52:120–125.
Michener, C. D. 1954. Observations on the pupae of bees. Pan-Pacific Entomologist 30:63–70.
Michener, C. D. 2000. The Bees of the World. Johns Hopkins University Press, Baltimore, MD. xivþ[1]þ913 pp.
Michener, C. D., and R. B. Lange. 1958. Observations on the ethology of neotropical anthophorine bees
(Hymenoptera: Apoidea). University of Kansas Science Bulletin 39(3):69–96.
Michener, C. D., M. S. Engel, and R. Ayala. 2003. The bee genus Caupolicana in Central America
(Hymenoptera: Colletidae). Journal of the Kansas Entomological Society 76(2):160–171.
Torchio, P. F., and G. E. Trostle. 1986. Biological notes on Anthophora urbana urbana and its parasite,
Xeromelecta californica (Hymenoptera: Anthophoridae), including descriptions of late embryogenesis and
hatching. Annals of the Entomological Society of America 79:434–447.