ORIGINAL ARTICLE

Two new Andean species of Neocorynura (Hymenoptera: Halictidae:Augochlorini) with notes on their biology

VICTOR H. GONZALEZ1, ALLAN H. SMITH-PARDO1 & GIOVANNI BOGOTA2

1Division of Entomology, Natural History Museum, University of Kansas, Lawrence, KS, USA, and 2Facultad del Medio

Ambiente y Recursos Naturales, Universidad Francisco Jose de Caldas, Bogota, Colombia

(Received 27 May 2005; Accepted 17 January 2005)

AbstractWe describe and illustrate the male, female, and pupa of two new species of Neocorynura from the Colombian Andes, andprovide information on their nesting behavior, host plants and social organization. Nests of Neocorynura iguaquensis sp. nov.Smith-Pardo & Gonzalez and N. muiscae sp. nov. Smith-Pardo & Gonzalez were found in flat or vertical banks, andcontained one to three adult females. The percentage of multi-female nests varied considerably from 8.3% to 40% of thetotal of dissected nests at different times in 2000 and 2003. The ovarian development, wing and mandibular wear of femalesin the nests with two or more females indicated that they were the offspring that remained in the nest before dispersal or amother and a newly emerged daughter. Pollen taken from the nest cells and scopae of museum specimens showed that bothspecies rely heavily on pollen of asteraceans and may be active year-round. Details on the nest architecture and commentson the phylogenetic placement of both species are also provided. Direct observations on the multi-female nests are requiredto determine whether or not such nests contain temporary assemblages or truly semisocial colonies.

ResumenDescribimos e ilustramos el macho, hembra y pupa de dos especies nuevas de Neocorynura de los Andes colombianos, yproporcionamos informacion sobre su biologıa de nidificacion, plantas hospederas y organizacion social. Los nidos deNeocorynura iguaquensis sp. nov. Smith-Pardo & Gonzalez y N. muiscae sp. nov. Smith-Pardo & Gonzalez fueronencontrados en el suelo, en areas planas o barrancos verticales, y tenıan de una a tres hembras adultas. El porcentaje denidos con mas de una hembra vario considerablemente desde 8.3% al 40% del total de nidos disectados a diferentes mesesen el ano 2000 y 2003. El desarrollo ovarico, desgaste de las alas y mandıbulas de las hembras en los nidos con dos o mashembras indicaron que ellas fueron las crıas que permanecieron en el nido antes de dispersarse o una madre con la hija reciennacida. El polen tomado de las celdas de crıa y escopas de especimenes de museo mostro que ambas especies dependenprincipalmente de asteraceas y que podrıan estar activas todo el ano. Tambien oferecemos detalles de la arquitectura delnidos y comentarios sobre la posicion filogenetica de ambas especies. Observaciones directas de los nido con mas de unahembra son necesarias para determinar si tales nidos son agrupaciones temporales o verdaderas colonias semisociales.

Keywords: Neocorynura, nesting biology, Andes, Colombia

Introduction

Neocorynura Schrottky is one of the larger genera of

the primarily Neotropical tribe Augochlorini

(Hymenoptera: Halictidae), containing more than

60 named (Engel, 2000) and perhaps an equal

number of undescribed species (Smith-Pardo,

2005a). Neocorynura ranges from Argentina to

Mexico, but is perhaps most diverse in northern

South America, where several species reach high

altitudes in the Andes. In fact, Neocorynura and

Caenohalictus (Halictidae, Halictini) are likely the

most diverse groups among those bee taxa occurring

in the higher Andes (Gonzalez & Engel, 2004).

Most species of Neocorynura nest in the ground

but some species nest in rotten wood (Engel, 2000).

The purpose of this paper is to describe two new

ground-nesting species of Neocorynura from the

Colombian Andes, and provide information on their

nesting biology, host plants, and social organization.

Neocorynura iguaquensis sp. nov and N. muiscae sp.

Correspondence: V. H. Gonzalez, Division of Entomology, Snow Hall, 1460 Jayhawk Blvd., University of Kansas, Lawrence, KS 66045, USA. Fax: . Fax: +1

(785) 864-5260. Email: vhgonza@ku.edu

Studies on Neotropical Fauna and Environment, December 2006; 41(3): 197–208

ISSN 0165-0521 print/ISSN 1744-5140 online # 2006 Taylor & Francis

DOI: 10.1080/01650520600630782

nov. appeared as Neocorynura sp. 1 and Neocorynura

sp. 2, respectively, in a paper regarding the

kleptoparasitic behavior of the phorid fly Megaselia

on the brood cells of both species (Gonzalez et al.,

2002).

Material and methods

Taxonomic descriptions

Morphological observations, measurements, and

illustrations were made using an ocular micrometer

on an Olympus SZ60 microscope. The abbreviations

F, S, T, OD and PD are used for flagellomere,

metasomal sternum and tergum, and ocellar dia-

meter and puncture diameter, respectively.

Morphological terminology follows that of

Michener (2000) and Harris (1979) for surface

sculpturing. Description format generally follows

that used for other augochlorine bees (e.g. Engel,

2000; Engel & Smith-Pardo, 2004). Mean values are

given with standard errors.

Type specimens and immature stages of the bees

and associated organisms are deposited in the

following institutions: AMNH American Museum

of Natural History, New York (J. G. Rozen); IAVH

Instituto Alexander von Humboldt, Villa de Leyva,

Boyaca, Colombia (J. E. Castillo); ICN Museo de

Historia Natural, Instituto de Ciencias Naturales,

Universidad Nacional de Colombia, Bogota,

Colombia (G. Amat); LABUN Laboratorio de

Investigaciones en Abejas, Universidad Nacional de

Colombia, Santa Fe de Bogota (G. Nates-Parra),

MEFLG Museo de Entomologıa Francisco Luis

Gallego, Universidad Nacional de Colombia, Sede

Medellın, Colombia (G. Morales), SEMC

Entomology Division, Natural History Museum,

University of Kansas, Lawrence, KS, 66045-7523,

USA (Z. Falin).

Nest excavations

Nests of Neocorynura iguaquensis sp. nov. and N.

muiscae sp. nov. were collected in the middle and

end of the wet season, 22–26 June and 16–20 July

2000, and 22–24 August 2003, at an altitude of

2850 m in the Eastern Andes of Colombia, at the

Santuario de Fauna y Flora de Iguaque National

Park (Carrizal) in the Department of Boyaca (5.70uN, 73.46u W). The site is located in the transition

between cloud forest and Paramo; in this area, the

dry season begins in September and extends until

the end of March when rains begin. There are on

average 84 mm of rainfall during the dry season. The

rainy season continues through August, with an

average of 209 mm of rainfall during this season, the

annual average being 294 mm. The median annual

temperature is 12uC (IGAC, 1996).

A total of 68 nests of Neocorynura iguaquensis sp.

nov. and 31 of N. muiscae sp. nov. were excavated

(Table I) during cold, cloudy weather when the bees

should have been in their nests. Cell clusters were

carefully transferred to the laboratory, cells were

opened with forceps and their contents examined

under a stereomicroscope. Adults and immatures

were fixed in Kahle’s solution (Triplehorn &

Johnson, 2005) for later dissection and examination,

and pollen was transferred to a plastic tray for later

identification. Cells contents and female’s ovarian

development were only examined in the samples

collected in 2000. The ovarian development was

examined in females collected from nests that had

more than one female and is provided according to

the types defined by Michener and Wille (1961).

Finally, the presence of hydrophobic glandular

secretions on the inner wall of turrets, tunnels, and

cell cavities of the nests was tested under the

stereomicroscope by putting a drop of water on

the surface: if glandular secretions were present,

the water would not be immediately absorbed by

the substrate.

Most nest features were measured in the field

using dial calipers, except the diameter of cell

entrances and thickness of cell closures, which were

taken with an ocular micrometer on an Olympus

SZ60 microscope. Nest descriptions follow

Sakagami and Michener (1962).

Palynological analysis

Pollen samples from brood cells of Neocorynura

iguaquensis sp. nov. nests 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.

Additionally, pollen from the bodies of four

Table I. Number of nests (percentage in parenthesis) of

Neocorynura iguaquensis sp. nov. and N. muiscae sp. nov. collected

in 2000 and 2003 at Iguaque National Park (Colombia) contain-

ing one, two and three adult females.

Number of nests (% in parentheses)

N. iguaquensis sp. nov. 1 R 2 RR 3 RR Total

June 2000 9 (75) 3 (25) 0 12

July 2000 11 (91.7) 0 1 (8.3) 12

August 2003 27 (61.3) 12 (27.3) 5 (11.4) 44

N. muiscae sp. nov.

June 2000 7 (77.8) 2 (22.2) 0 9

July 2000 11 (91.7) 1 (8.3) 0 12

August 2003 6 (60) 4 (40) 0 10

198 V. H. Gonzalez et al.

museum specimens (females) of N. muiscae sp. nov.

collected on 4 January 1992 from the same locality

was examined. Bees were placed in individual vials

with 95% ethanol, and then gently shaken for about

5 min to remove the pollen from their bodies. The

ethanol was discharged and the pollen treated as

explained above.

Pollen grains were identified and counted by

sampling all grains found in a row until 1000 grains

were reached. Pollen identification was done using

the following palynological atlases for the Paramo

vegetation of the Colombian Eastern Andes:

Velasquez and Rangel (1995), Bogota et al.

(1996), Velasquez (1999) and Bogota (2002).

Examination of pollen was at 40X and 100X

magnification with a Leitz DIALUX 22 EB micro-

scope. Pollen samples are deposited in the

Palynological collection of the Instituto de Ciencias

Naturales, Universidad Nacional de Colombia,

Bogota. Mean values are given with standard errors.

Results

Systematics

Neocorynura iguaquensis Smith-Pardo &

Gonzalez, new species (Figure 1)

Diagnosis

Males and females of N. iguaquensis sp. nov. can be

easily recognized by the distinctive color pattern,

with the head and mesosoma mainly black, con-

trasting with the legs, sterna and T1 and T2 largely

orange; remainder of terga mainly dark brown to

black. This species can be distinguished from other

species with similar body color such as N. papallac-

tensis Engel by the sulcate gena and postgena, and

the base of propodeum metallic green, basally

sulcate and distally imbricate. In N. papallactensis,

the gena is strongly imbricate to rugose, and the

basal area of the propodeum is microareolate to

rugose, mainly dull green with weak coppery

reflections.

Description

Female. Total body length, 9.7 mm (¡0.64, 8–11,

n517); forewing length 8.1 mm (¡0.3, 7.3–8.4,

n518). Head wider than long. Scape longer than

combined lengths of F1–F6; F10 longer than

preceding flagellomeres; F2 subequal in length to

F1. Clypeus with upper margin above lower tangent

of compound eyes, paraocular lobe obtuse (,110u).Mesoscutum slightly wider than long; metanotum

more than half length of mesoscutellum. Meso-

tibial spur serrate, more than half length of

mesobasitarsus; inner metatibial spur with five

teeth, the most distal almost completely fused with

spur apex (fifth); 1m-cu confluent with 1rs-m;

2m-cu basad 2rs-m by approximately five times

vein width; first submarginal cell as long as

combined lengths of second and third submarginal

cells; second submarginal cell with anterior margin

slightly longer than anterior margin of third marginal

cell; hamuli spaced 3-1-1-1-1-1. Basal area of

propodeum about as long as mesoscutellum.

Metasoma elongate, semi-petiolate.

Mandible with strong acetabular groove. Labrum

with weak sulcus and small, more or less narrow

central process (orbiculate elevation of medial

region). Clypeus, supraclypeal and lower paraocular

areas dull, minutely but strongly imbricate, punctu-

res sparser (2 PD) and larger on clypeus, denser on

lower paraocular areas; frons, remainder of face and

vertex strongly punctate; gena and postgena strongly

imbricate and striate. Scape imbricate with fine

punctures irregularly separated. Pronotum and

propleuron imbricate. Mesoscutum dull, densely

punctured (( PD); punctures sparser on slightly

shining medio-anterior area, integument between

punctures imbricate; mesoscutellum and metano-

tum dull, strongly punctate. Metanotum faintly

imbricate, punctures separated by 1–2 PD. Pleura

strongly imbricate, largely dull, punctures on pro-

pleuron coarser than on mesoscutum. Tegula faintly

imbricate. Basal two-thirds of propodeum striate,

apical one-third imbricate but shining.

Figure 1. Male terminalia of Neocorynura iguaquensis sp.

nov. A5sixth sternum, B5fused seventh and eighth sterna, C and

D5genital capsule, ventral (left half) and dorsal (right half), and

in lateral views, respectively.

Two new Andean Neocorynura 199

Head and mesosoma mainly black, except as

follows: apex of mandible and labrum light brown;

basal half of clypeus, supraclypeal and paraocular

areas, gena, pronotum, lateral and posterior margins

of mesoscutum, all margins of mesoscutellum,

metanotum, and base of propodeum weakly metallic

green, with coppery reflections; lateroposterior area

of propodeum mainly with coppery highlights. Legs

orange. T1 and T2 mainly orange, apical margins

dark brown; T3–T5 mainly dark brown to blackish;

T2–T5 with basal margins feebly metallic green or

coppery. Sterna mainly orange, distal sterna dark

brown. Wing veins and pterostigma dusky brown.

Face with erect and mostly simple setae, longer

and denser on paraocular areas. Area along inner

margin of compound eye, above antennal sockets

and vertex with sparse brown setae. Gena densely

covered with highly branched pale yellow setae;

longer, thinner and sparser on postgena.

Mesoscutum, mesoscutellum and metanotum den-

sely covered with erect, short brown setae, more

densely covering the metanotum. Pronotum, prono-

tal lobe, subalar area, pre-episternum, mesoscutum

and mesoscutellum with long, erect and simple or

minutely branched black or brown setae, longest and

sparser on mesoscutellum; lateral and dorsal ridges

of pronotum, surrounding areas of pronotal lobe and

subalar area densely covered with whitish tomen-

tum. Ventral area of mesepisternum, lateral poster-

ior surfaces of propodeum, and sides of metanotum

with dense, long, branched orange setae, shorter on

the posterior surface of propodeum, projecting from

an appressed and dense whitish pubescence.

Pubescence of legs mainly orange. T1 with dense,

long (1–1.5 OD) branched yellowish setae on basal

half, shorter on sides, distal half almost bare. T2–T6

with two types of setae: dense, appressed, short

blackish or dark brown setae, longer on T5–T6; long

pale yellowish setae, denser and longer on sides of

terga and mostly absent dorsally, such setae darker,

longer and denser on terminal terga. Basal zones of

T2–5 with yellowish tomentum forming distinct

basal bands. Sterna with long orange scopal setae.

Male. As described for the female, except for usual

sexual characters, and as follows: Body length

13 mm (¡0.60, 12.5–14.0, n54); forewing length

10 mm (¡0.45, 9.1–11.3, n54). Head about as

wide as long. F1 shorter than F2. Paraocular lobe

more obtuse. Mesoscutum slightly wider than long;

inner metatibial spur serrate, one third length of

metabasitarsus, rather slender and slightly curved.

Hamuli with a spacing pattern 4-1-4. Metasoma

slightly more elongate than in female; S6, S7+S8,

and genital capsule as in Figure 1.

Clypeus metallic green, with copper-red high-

lights; upper frons and vertex black with blue-green

highlights. Scape dark brown with strong green

reflections on outer surface. Metepisternum black,

with weak blue-green reflections. Legs black (green-

ish reflections on front and hind coxae) with the

following parts orange: tarsi, apices of tibiae, base of

hind tibiae, apex of hind femur. T1 with dark brown

spots on base and apical margin, T2 with apical third

dark brown. T4–T6 with green reflections. S1–S3

orange, remainder of sterna dark brown.

Pubescence yellowish, longer and denser than

female. Scape densely covered with branched setae

(about twice as long as scape diameter). Gena and

postgena densely covered with highly branched

orange setae; longest on postgena. Dorsal and lateral

ridge of pronotum, pronotal lobe, mesoscutum,

mesoscutellum, and metanotum densely covered

with long, erect, branched brown setae; longest on

mesoscutellum. Mesepisternum, and lateral and

posterior surfaces of propodeum with long,

branched, erect orange setae. T1 and basal half of

T2 densely covered with long, branched, semi-erect

orange setae; apical half of T2 and remaining terga

with shorter, sparser, semierect setae. Basal bands of

pale tomentum weak on T1 and T3, otherwise

absent. Sterna densely covered with small,

appressed, orange setae.

Type material

Holotype. R, Colombia. Boyaca, Santuario de Flora

y Fauna de Iguaque, 5.70u N, 73.46u W, 2850 m, 19

July 2000. V. H. Gonzalez [SEMC]. Paratypes. 12Rwith same data as the holotype but collected on 18

July, 23–25 June 2000 [AMNH, IAVH, ICN,

LABUN, MEFLG, SEMC,]; 1R, idem, 1

November 1987, C. Sarmiento, M. Guevara

[LABUN]; 1R, idem, La Laguna, 9 January 1989

[LABUN]; 1R, idem, Morronegro, 15 January 1991.

H. Alonso [SEMC]; 1R, 2„, idem, Francos, 3000 m,

5 January 1992. A. Amarillo [AMNH, LABUN];

1„, idem, C. Sarmiento [ICN]. Cundinamarca, 1R,

La Calera, Vereda Buenos Aires, 5 May 1991,

2880 m. C. Sarmiento [MEFLG]; 1„, San Antonio

de Tena, PNN. Chicaque, 9 June 1993, 2150 m,

C. Sarmiento [LABUN].

Etymology

The specific epithet refers to the type locality, the

National Park Santuario de Fauna y Flora de

Iguaque, Colombia.

Variation

Variation noted in the following characters: vein 1m-

cu basad or confluent with 1rs-m; length of anterior

margin of third submarginal cell as long as or longer

200 V. H. Gonzalez et al.

than anterior margin of second submarginal cell;

coloration of trochanter, femur and basitarsus of

forelegs, and T1 and T2 ranging from orange

to largely dark brown. One female paratype

from Iguaque has both forewings with only two

submarginal cells.

Pupa. Pupae of both sexes (n57 of each sex) agree

closely with pupal description of Augochlora

nominata Michener (Eickwort & Eickwort, 1972),

except for presence of small rounded tubercles on

base and apex of outer surface of scape, all coxae

with long rounded spines, fore and mid-femora

with rounded protuberances, S6 of female with

rounded apical tubercle. Number and arrangement

of tubercles on metasomal terga as follows: no

tubercles on T1; T2–T5 with pointed tubercles

progressively increasing in size toward the end of

metasoma, except T5 in female and T7 in male,

with shortest tubercles; 6–9 tubercles per side on

T2; 6–10 on T3; 7–9 on T4; the most lateral

tubercle on each side of T3 and T4 longest; 5–7 on

T5; 2–4 on T6 and 2–3 on T7 in male. On all terga,

tubercle series broken by median gap. Hind tibia of

female with apical tubercle virtually absent.

Metasomal S4 of male without long spines. Caudal

extremity of both sexes without doorknob-shaped

protuberance.

Neocorynura muiscae Smith-Pardo & Gonzalez,

new species (Figure 2)

Diagnosis

Males and females of N. muiscae sp. nov. can be

recognized by the combination of the following

characters: head and mesosoma metallic green,

metasoma largely dull green, with weak orange to

coppery highlights (looks darker with little or no

green reflections at lower magnifications); base of

propodeum sulcate, integument otherwise imbricate,

and T1 imbricate, with sparser punctures on basal

half than distal half. Neocorynura muiscae sp. nov.

belongs to the same group as N. papallactensis

(Ecuador) and N. iguaquensis sp. nov., differing from

these species, in addition to the characters mentioned

above, by the strongly sulcate gena, and the presence

of basal bands of appressed setae on T2–T5.

Description

Female. In general as described for N. iguaquensis sp.

nov. except: total body length, 9.1 mm (¡0.45, 8.9–

10.2, n 510); forewing length 7.5 mm (¡0.2, 7.2–

7.7, n 510). Head slightly wider than long. Scape

longer than combined lengths of F1–F5; F2 slightly

longer than F1. Clypeus wider than long, ocular angle

(lateral angle formed by epistomal sulcus) obtuse.

Inner metatibial spur pectinate, with four teeth (not

including apex), the most distal separated from spur

apex; second submarginal cell rectangular; anterior

margin of second submarginal cell as long as anterior

margin of third marginal cell; hamuli spaced 4:1:3.

Head and mesosoma metallic green, except for

mandible (apex reddish), labrum, apical third of

clypeus, antenna, upper paraocular area, along inner

margin of compound eye, tegula, legs and sterna

dark brown to black; face, pronotum and midanter-

ior area of mesoscutum with weak coppery reflec-

tions, mesopleura with blue reflections. Metasoma

largely dull green, with weak orange to coppery

highlights, especially on basal half of T1 and T5.

Wings slightly dark, darker along costal margin;

veins brown, pterostigma honey colored.

Mandible with few simple, long setae (0.5–2.5

OD). Labrum mostly bare with stiff, brown, simple

setae along lower margin. Clypeus with brown,

simple setae (1 OD) along apical margin, sides with

branched, denser and shorter setae (0.5–1 OD).

Supraclypeal and subantennal areas with very sparse

and short (%OD) branched setae; sparser on latter.

Paraocular area and frons with longer and denser

setae. Scape with stiff, dark brown and poorly

branched setae (( 1.5 OD), longest on basal third.

Vertex with stiff, minutely branched, dark brown

setae (0.5–1 OD) in length; postgena with sparse,

Figure 2. Male terminalia of Neocorynura muiscae sp. nov.

A5sixth sternum, B5fused seventh and eighth sterna, C and

D5genital capsule, ventral (left half) and dorsal (right half), and

in lateral views, respectively.

Two new Andean Neocorynura 201

branched, light brown setae (1–2.5 OD). Pronotum

and propleuron with minute, light brown setae,

stiffer and lighter in color on posterior margin of

pronotal lobe. Mesoscutum covered with stiff,

minutely branched light brown setae (0.5–2 OD).

Mesoscutellum mostly covered with sparse, minute

(1–2), dark brown setae. Metanotum as mesoscu-

tellum but with shorter (0.5–1.5 OD) and lighter

setae. Base of propodeum nearly bare, sides with

branched, minute to 1 OD, light brown setae.

Mesepisternum with mostly minute and scattered

light brown setae, denser towards venter, remainder

areas covered with minute, light brown setae mixed

with some scattered dark setae; metepisternum with

light brown setae, minutely branched and uniformly

abundant. Procoxa with branched light brown setae

(1.5–2.0 OD); protrochanter with scattered, poorly

branched setae brown; profemur with inner side

mostly bare, outer side covered with stiff, branched,

brown setae, posterior margin with longer and

darker setae (1–1.5 OD); protibia covered with

dense, minute, light brown setae, inner surface more

densely cover with longer setae (%1.5 OD); tarsi as

protibia, but with longer setae on outer margins;

mesocoxa mostly bare, a few thick, simple, light

brown setae (2–2.5 OD) near articulation with

mesotrochanter; mesotrochanter with stiff, branched

yellow setae (1–1.5 OD), sparser on inner margin;

mesofemur with most of outer side covered with

minute, light brown setae, inner side mostly bare, a

dense patch of longer setae on posterior margin;

mesotibia covered with minute, simple setae (%OD),

uniformly distributed, inner side with row of longer

and simple setae; metatrochanter and metacoxae

covered with light brown and branched setae, 1–1.5

OD in length; metafemur with light brown setae

(2–4 OD), each seta with 2–4 branches on apex,

shorter on posterior margin; metatibia covered with

stiff, mostly simple, light brown setae (%OD),

margins with longer setae (1–1.5 OD), 5–6 branches

on apex on each setae. T1 with basal half with sparse,

light brown branched setae (1–1.5 OD); T2–T4 with

basal band of minute white, branched setae, minute

brown setae on remainder of tergum; setae longer

(1–1.5 OD) on T3 and T4; T5 with basal band of

white minute setae, remainder of T5 densely covered

with stiff, branched brown setae (,2.5 OD) in length,

the longer on posterior margins.

Male. As described for female except as follows

(n54): total body length 9 mm (¡0.44, 8.2–9.1);

forewing length 7.4 mm (¡0.3, 7.0–7.8). Mandible

slender and pointed, without preapical teeth, outer

surface without grooves, lower margin slightly

carinate, upper margin continuous. Labrum

smooth and shiny, without tubercle. Paraocular

lobe more angular than female. Inner metatibial

spur slender and weakly curved, one fourth of

metabasitarsus length. Hamuli spaced 3-1-3.

Metasoma slightly more elongate in appearance;

S6, S7+S8, and genital capsule as in Figure 2.

Subantennal and lower paraocular areas, and

metanotum strongly punctured. Clypeus metallic

green. Legs dark brown with green reflections. Terga

with bases metallic green, especially on T1 and T5,

remainder brown, with yellowish reflections.

Mandible with sparser, simple yellowish setae

(1.5–2.5 OD). Postgena with sparse, branched setae

(2–3 OD). Mesoscutum and mesoscutellum covered

with light brown setae, darker, sparser and longer

on latter. Legs in general less pubescent than in

female; profemur and protibia with some dark

brown (0.5–1 OD) setae along posterior margins,

much sparser on latter; metafemur covered with

shorter setae (minute-1 OD), especially on outer

side. Metasomal pubescence as in the female, except

setae lighter in color, denser on T1; sterna with

sparser pubescence than in female.

Type material

Holotype. R, Colombia: Boyaca, Santuario de Flora

y Fauna de Iguaque, 5.70u N, 73.46u W, 2850 m, 20

July 2000. V. H. Gonzalez [SEMC]. Paratypes. 7Rand 3 „ with the same data of the holotype but

collected on 19 and 25 June 2000 [AMNH, ICN,

MEFLG, SEMC]; 1R, idem, 4 January 1992, A.

Tamayo; 1R, 2„, idem, C. Sarmiento [LABUN,

SEMC].

Etymology

The specific epithet honors the Muiscas, an indi-

genous Colombian nation that formerly inhabited

the high plains in the Central Andes, between the

departments of Boyaca and Cundinamarca.

Variation

The coloration of clypeus, mesoscutellum, and T2–

T4 ranges from dull to metallic green with or

without orange highlights in the female.

Pupa (n5three males and four females). As

described for N. iguaquensis sp. nov. except for the

presence of larger tubercles, particularly those on the

head and mesoscutellum; number of tubercles on

metasomal terga as follows: T2 with 5–7 tubercles

per side; 7–9 on T3; 6–8 on T4; 3–5 on T5; 2–4 on

T6; and 1–2 on T7 of male.

Comments

Pupae of N. iguaquensis sp. nov. and N. muiscae sp.

nov. differ from that of N. colombiana (Eickwort,

202 V. H. Gonzalez et al.

1979) by the absence of long spines on S4 of male

and the presence of tubercles on eye summit, base of

outer surface of scape, inner surfaces of scape and

pedicel in both sexes, and rounded apical tubercle

on S6 of female. However, the strongest similarity to

N. colombiana is the caudal extremities in both sexes,

which lack a doorknob-shaped protuberance, sup-

porting its absence as characteristic of the genus

Neocorynura.

Nest architecture

The nests of N. iguaquensis sp. nov. were usually

found scattered in nearly vertical banks (20–40 cm

high) along ditches and trails, and in grazed pastures

around the visitor’s center at Iguaque National Park.

The banks with nests face east and were sparsely

covered with mosses. The soil in flat ground and

banks was loose, black, wet to very wet clay.

Nests of N. iguaquensis sp. nov. were shallow

(Table II), and consisted of a short main burrow

ending in a single chamber containing only one

earthen cell cluster supported by pillars and rootlets.

The nest entrance was round and constricted, and

lacked an earthen turret; a nearly circular tumulus

was usually deposited outside the entrance. The

main burrow widened gradually in diameter toward

the bottom of the nest and entered the cell cavity

commonly above the cell cluster. The general

direction of the main burrow was variable, being

horizontally inclined upward or downward in those

nests located in the banks, to almost vertical for

those nests in flat ground. Two of the nests

excavated in 2000 had a single blind burrow

extending beyond the nearly spheroid cell cavity.

The inner wall of the main burrow and cell cavity

was smooth, presumably not lined with hydrophobic

secreted material. The cell cluster was separated

from the wall of the cell cavity and was supported by

1–5 earthen pillars; cell clusters in the pasture were

frequently tightly supported by rootlets in addition

to pillars. Cell clusters were rough on the outside

and cells were generally not externally discernible.

As in most halictine bees, cells were slightly flattened

on the bottom surface with a rounded oval upper

surface; their inner walls were smooth and lined with

a yellowish waterproof substance. Most cells were

vertical with their bases slightly diverging within the

cluster, although some were nearly horizontal; cell

closures were loose and concave on the inner face.

Complete pollen masses were nearly spherical but

were flattened on the bottom (,5 mm in diameter

and 3 mm high), and were placed on the flatter side

of the cell at the end opposite the entrance.

Nesting structures of Neocorynura muiscae sp. nov.

were as described for N. iguaquensis sp. nov. except

that the entrance usually had an earthen turret

projecting above the ground surface and was

surrounded by an elliptical tumulus. The surfaces

of the turrets were rough externally and smooth

internally but not polished. As in N. iguaquensis sp.

nov., there was no evidence of hydrophobic gland-

ular secretions lining the inner walls of turret, tunnel

and cell cavity. The general direction of the main

burrow was usually vertical in flat ground and it

entered the cell cavity at the top; however, the main

burrow sloped horizontally downward or upward in

those nests found in vertical banks. A single blind

burrow per cell cavity was present in 21% of the

nests excavated in 2000. The cell cavity was rather

ovoid in longitudinal section, and because the

majority of the nests were found in the pasture, cells

clusters were usually supported by rootlets, but also

had up to six pillars.

Contents of the nests

Based on our limited collection, the percentage of

nests of N. iguaquensis sp. nov. and N. muiscae sp.

nov. containing more than one adult female varied

considerably from 8.3% to 40% of the total of

dissected nests at different months (Table I).

Females in those multiple female nests had about

the same head width and differed in the degree of

wing and mandibular wear and ovarian development

as indicated in Table III.

Pooled across all nests of N. iguaquensis sp. nov.

collected in 2000, 10% of the cells contained eggs,

50% had larvae, 30% pupae, and 7.% had une-

merged adults (n540 sealed cells). A single adult

male was found in three of the nests excavated in

2000 and in one of the nests excavated in 2003.

Similarly, pooled across all nests of N. muiscae sp.

nov. excavated in 2000, 78% of cells had larvae,

19% had pupae, and 3% had unemerged adults

(n532 sealed cells). A single adult male was found in

three of the 31 nests excavated in both years.

Adult females were frequently seen flying over the

grazing area and some of them were excavating the

soil, presumably starting new nests. Before excava-

tions no females were seen either guarding nest

entrances or exhibiting any kind of aggressive

behavior during the excavations; instead they ran

away and hid at the bottom of the cell cavity.

Host plants

Pollen grains from 32 plant species belonging to 18

families were found within Neocorynura iguaquensis

sp. nov. cells (Table IV). The family Asteraceae

was represented in the pollen samples with the

highest number of species. The spectrum of pollen

types varied among cells and each cell had on

average identifiable pollen from 9.2 plant species

Two new Andean Neocorynura 203

Table II. Measurements (mm) of some structures of nests of Neocorynura iguaquensis sp. nov. and N. muiscae sp. nov. collected in 2000 at

Iguaque National Park (Colombia). Means are given with standard error followed by the range and sample size in parenthesis. N/A5no

applicable; *5tumulus of N. muiscae sp. nov. was elliptical, thus we provide a maximum and minimum diameter.

Nest structure N. iguaquensis sp. nov. N. muiscae sp. nov.

Maximum nest depth 80.6¡0.6 (64.6–110.9, n515) 87.3¡0.5 (75–113.0, n513)

Turret Absent Present

Height N/A 17.7¡1.0 (8.1–38, n56)

Inner diameter N/A 4.1¡0.03 (3.3–5.0, n510)

Outer diameter N/A 9.3¡0.08 (8.0–10.6, n57)

Tumulus diameter * 44.5¡0.1 (41.7–50.5, n58) max. 34.2¡0.9; min 29.3¡1.0 (n53)

Nest entrance (diameter) 3.7¡0.4 (3.3–4.8, n511) N/A

Main burrow (diameter)

Below entrance 4.3¡0.4 (n54) 4.0¡0.04 (n56)

Midpoint 4.2¡0.4 (n510) 4.4¡0.06 (n57)

At entrance of cell cavity 7.3¡1.1 (n514) 6.2¡0.06 (n510)

Main burrow length 46.8¡8.8 (36.0–69.6, n513) 50.6¡0.8 (22.0–68.0, n510)

Blind burrows

Length 16.3–20 (n52) 14.8¡0.1 (6.8–26.1, n55)

Diameter 4.6–7.8 (n52) 6.3¡0.07 (4.7–7.0, n58)

Cell cavity dimensions

Width 27.8¡6.1 (18.1–35.0, n512) 23.4¡0.2 (18.0–26.4, n58)

Length 26.4¡5.5 (22.0–36.5, n515) 24.7¡0.2 (16.0–32.7, n513)

Height 28.5¡4.5 (22.6–36.5, n516) 28.1¡0.2 (22.5–37.5, n513)

Space between cavity & cell cluster 5.3¡0.6 (4.7–6.2, n58) 4.7¡0.1 (n54)

Pillars

Diameter 3.1¡0.9 (2.3–4.0, n53) 5.8–7.1 (n52)

Length 4.3¡1.0 (3.0–5.4, n56) 3.8¡1.2 (2.8–5.5, n54)

Number of cells per cluster 4.1¡1.9 (2–9, n517) 2,9¡0.4 (1–6, n517)

Cell cluster dimensions

Width 14.3¡0.4 (14.0–24.3, n57) 17.7¡0.6 (14.0–24.0, n54)

Length 20.9¡3.4 (13.0–23.8, n57) 20.8¡0.5 (17.0–26.4, n54)

Height 16.0¡1.8 (14.2–18.2, n57) 16.4¡0.4 (11.8–18.7, n54)

Thickness of outer walls 1.3¡0.5 (0.8–2.2, n510) 1.3¡0.3 (0.9–2.1, n58)

Thickness of wall between cells 1.9¡0.8 (1.0–3.2, n57) 1.8¡0.5 (1.0–3.5, n510)

Cell dimensions

Length 12.7¡0.5 (12.1–13.8, n510) 12.3¡0.2 (11.7–13.3, n513)

Max. diameter 6.0¡0.4 (5.4–6.4, n512) 5.6¡0.3 (4.9–6.4, n512)

Diameter at neck 3.7¡0.2 (3.4–3.9, n55) 3.4¡0.2 (3.3–3.5, n53)

Cell entrance (diameter) 3.0¡0.1 (2.8–3.2, n522) 3.0¡0.1 (2.7–3.2, n511)

Cell closure thickness 1.7¡0.2 (1.6–2.1, n55) 1.6¡0.2 (1.4–2.0, n55)

Table III. Cell contents, mandibular and wing wear, and ovarian condition of females in multiple female nests of Neocorynura iguaquensis

sp. nov. and N. muiscae sp. nov. collected in 2000 at Iguaque National Park (Colombia). Developed and undeveloped ovaries were similar

to those ovary types A or B, and type E, of Michener and Wille (1961), respectively.

No.

females

Open

cells

Closed

cells Eggs Larvae Pupae

Callow

adults Wings Mandibles Ovaries

N. iguaquensis sp. nov.

A 2 0 3 0 2 0 1 Unworn,

Worn

Unworn,

Worn

Developed

Developed

B 2 2 7 2 3 2 0 Unworn,

Worn

Unworn,

Worn

Developed

Developed

C 2 2 4 0 4 0 0 Unworn,

Unworn

Unworn,

Unworn

Undeveloped

Developed

D 3 3 2 1 1 0 0 Unworn,

Unworn,

Unworn

Unworn,

Worn,

Unworn

Undeveloped,

Developed,

Developed

N. muiscae sp. nov.

A 2 0 3 0 3 0 0 Unworn,

Unworn

Unworn,

Unworn

Developed,

Developed

B 2 0 3 0 0 3 0 Unworn,

Unworn

Unworn,

Unworn

Developed,

Undeveloped

C 2 0 3 0 0 3 0 Unworn,

Unworn

Unworn,

Unworn

Developed,

Undeveloped

204 V. H. Gonzalez et al.

(¡0.5, 5–13). Although Viburnum sp.

(Caprifoliaceae) was present in all samples and

accounted for 42.5% of the total sampled pollen, it

was not always the dominant plant species in all

cells. Ageratina type, Gnaphalium-Lourteigia type,

Hypochaeris radicata, Loricaria complanata, Smilax

sp. and Espeletia type were present in percentages

above 1% and all together (including Viburnum sp.)

constituted more than 50% of the pollen found in

the sampled cells.

Similarly, pollen from 23 plant species of 14

families was identified from the museum specimens

of Neocorynura muiscae sp. nov. Almost all of the

species were also found in N. iguaquensis sp. nov.

Table IV. Spectrum of plant species and pollen types recorded in pollen samples (n515 brood cells) from nine nests of Neocorynura

iguaquensis sp. nov. collected in the middle of the wet season (June and July 2000) in Iguaque National Park (Colombia). Proportion (%)

per cell is given with standard error and range in parentheses.

Plant species Overall proportion (%) Proportion per cell (%) Number of cells

Apiaceae

Hydrocotyle sp. 0.02 0.3 1

Araliaceae

Oreopanax mutisianum (H.B.K.) Decne. & Planch. 0.04 0.3 1

Asteraceae

Asteraceae type 0.01 0.2 1

Ageratina type 16.1 20.1¡4.6 (0.06–45.3) 14

Espeletia type 1.3 1.4¡3.3 (0.2–9.7) 7

Bidens type 0.9 1.2¡3.9 (1.4–7) 4

Gnaphalium-Lourteigia type 13.8 14.3¡3.5 (0.5–39) 14

Hypochaeris radicata L. 9.2 7¡2.5 (0.1–19.4) 8

Loricaria complanata Wedd. 8.0 9.6¡3.4 (1.1–42.5) 12

Munnozia sp. 0.1 2.5 1

Betulaceae

Alnus sp. 0.4 2.6¡1.7 2

Caprifoliaceae

Viburnum sp. 42.5 46.5¡5 (11.9–78) 15

Chloranthaceae

Hedyosmun sp. 0.06 0.6¡0.1 2

Cunoniaceae

Weinmannia sp. 0.4 1.9¡1.3 (0.2–5.9) 4

Loranthaceae

Gaidendron sp. 0.02 0.1¡0.01 2

Melastomataceae

Bucquetia glutinosa DC. 0.3 0.8¡0.2 (0.2–1.3) 5

Miconia type 0.8 0.8¡0.1 (0.1–1.9) 14

Myrsinaceae

Myrsine coriacea (Sw.) R. Br. 0.01 0.1 1

Myrtaceae

Myrtaceae type 0.01 0.1 1

Poaceae

Poaceae type 0.2 0.9¡0.7 (0.1–2.3) 3

Polygonaceae

Rumex acetosella L. 0.01 0.1 1

Rosaceae

Rubus bogotensis Kunth 0.09 0.9¡0.3 2

Rubus robustus C. Presl. 0.1 1.1¡1.1 2

Rubiaceae

Arcytophyllum sp. 0.03 0.3 1

Borreria sp. 0.02 0.2¡0.1 2

Cinchona type 0.08 0.6¡0.2 (0.3–1) 3

Smilacaceae

Smilax sp. 4.9 6.5¡3.4 (0.06–29.2) 8

Solanaceae

Cestrum sp. 0.02 0.4 1

Solanum sp. 0.01 0.2 1

Valerianaceae

Valeriana pilosa Ruız & Pavon 0.02 0.3 1

Unknown

type 1 0.6 1.9¡1 2

type 2 0.06 0.8 1

Two new Andean Neocorynura 205

cells; however, about 80% of the pollen types is

present in trace amounts less than 1% and may be

contaminants or were gathered incidentally during

nectar visits (Table V). Gnaphalium-Lourteigia type

accounted for about 70% of the total pollen, and

ranged from 48% to 96% among bee specimens.

Discussion

Systematics

Neocorynura papallactensis, N. iguaquensis sp. nov.,

N. muiscae sp. nov. and at least one undescribed

species from lower elevations in Panama form part of

a well-supported clade of primarily high Andean

bees that occurs from Venezuela to Ecuador, and

which is closely related to a clade with species

ranging from central South America to central

Mexico (Smith-Pardo, 2005a). Most of the species

of the iguaquensis group have an unusual semi-obtuse

epistomal suture, very weak occipital carina, and

exhibit a distinctive color pattern that resembles the

polistine wasps Agelaia multipicta (Haliday)

(Hymenopera, Vespidae) (Smith-Pardo, 2005b), a

sympatric and very common social wasp in Iguaque

suggesting a possible mimicry complex around this

wasp species (Sarmiento, 1993).

Biology

The nesting biology of Neocorynura iguaquensis sp.

nov. and N. muiscae sp. nov. is similar to that

described for other ground-nesting species of the

genus (e.g., Michener et al., 1966; Michener, 1977),

consisting of shallow nests with a short main burrow

ending in a single chamber containing only one

earthen cell cluster, and usually occupied by a single

adult female.

Except for the nests A and B of N. iguaquensis sp.

nov., the unworn wings and mandibles of the

females in most of the multi-female nests

(Table III) suggest that individuals in most of these

nests were relatively young females of about the

same age; in addition, the ovarian development

shows that one and sometimes all of the females in a

nest were reproductively active. Assuming that the

females in those multi-female nests were related, our

observations suggest that they were likely sisters or as

in the case of the nests A and B of N. iguaquensis sp.

nov., a mother and a recently emerged daughter.

Therefore, the founder female in those nests

occupied by sisters may have died or left the nest.

It is likely that these multi-female nests are

temporary assemblages of offspring before one or

both of them disperses. Nonetheless, cell contents

indicate that at least one female was actively

provisioning. We have no data on the period that

newly emerged adults remain in their natal nests or

on whether or not females in such multiple female

nests were actively provisioning, mated (we did not

examine the spermatheca) and laying eggs.

However, anecdotal observations of a two-female

nest of N. iguaquensis sp. nov. suggest that both

individuals foraged for pollen (P. Montoya, pers.

comm.).

Neocorynura erinnys Schrottky, a lowland species,

has the highest reported number of adult females

(five) per nest recorded in the genus (Luderwaldt,

1911), and therefore it is usually referred to as a

semisocial species (Danforth & Eickwort, 1997;

Engel, 2000). However, the provisioning phase of

these females was never examined and nothing can

be said about the social behavior of this species.

Direct and detailed observations on these multi-

female nests in Neocorynura are thus required to

determine whether or not such nests contain

Table V. Spectrum of plant species and pollen types recorded in

pollen samples removed from the bodies of museum specimens of

Neocorynura muiscae (n54 females) collected in Iguaque on 4

January 1992.

Plant species Overall proportion (%)

Asteraceae

Ageratina type 21.3

Ambrosia type 0.5

Bidens type 0.2

Espeletia sp. 0.2

Jungia sp. 0.04

Loricaria complanata Wedd. 0.02

Gnaphalium-Lourteigia Type 72.4

Hypochaeris radicata L. 3

type 1 0.1

Caprifoliaceae

Viburnum sp. 0.3

Clusiaceae

Hypericum sp 1.3

Cunoniaceae

Weinmannia sp. 0.02

Cyperaceae

type 1 0.06

Fabaceae

Salvia sp. 0.02

Fagaceae

Quercus sp. 0.02

Melastomataceae

Bucquetia glutinosa DC. 0.4

Miconia type 0.02

Myrtaceae

Myrcianthes sp. 0.02

Poaceae

Poaceae type 0.02

Polygonaceae

Rumex acetosella L. 0.02

Rubiaceae

Arcytophyllum sp. 0.02

Scrophulariaceae

Castilleja sp. 0.02

Smilacaceae

Smilax sp. 0.02

206 V. H. Gonzalez et al.

temporary assemblages or are truly semisocial

colonies.

The palynological analysis of the brood provisions

of Neocorynura iguaquensis sp. nov. (Table IV)

indicated that this species collected almost exclu-

sively pollen from asteraceans and Viburnum sp.

(Caprifoliaceae). These results contrast with the

common view among melittologists that species of

Neocorynura may be polylectic as in other members

of the tribe Augochlorini, an idea that is also

supported by the generalized structure of the

mouthparts in Neocorynura in relation to other

augochlorines, and the scarce floral records

(Michener, 2000; Eickwort, 1969; Moure & Hurd,

1987; Smith-Pardo, 2005a). However, our sample

size was small and restricted to a single season and

locality, thus limiting conclusive interpretations on

the foraging behavior of the studied species.

Finally, the collections of individuals of both sexes

and of both species in Iguaque in every month of the

year suggest that the species are multivoltine.

Furthermore, pollen gathering activity by N. muiscae

sp. nov. females during the dry season (Table V)

indicates that they are nesting year-round.

Acknowledgments

This work is dedicated to the memory of Juan Pablo

Cardenas, beloved father of AHS-P. We thank

Daniel Bennett, William Wcislo, M. S. Engel and

C. D. Michener, Anne Zillikens and two anonymous

reviewers for their comments and suggestions on the

manuscript; Paula Montoya and Monica Ospina for

their help in the field and their comments on the

biology of the species; Carlos Sarmiento and

Guiomar Nates-Parra for lending us some of the

specimens for study, and finally The Instituto de

Ciencias Naturales, Universidad Nacional de

Colombia, for allowing GB to use the Palynological

laboratory. We are indebted to Fernando Fernandez,

Diana C. Arias and Jose E. Castillo from the Seccion

de Entomologıa of the Instituto de Investigacion

en Recursos Biologicos Alexander von Humboldt,

and El Sistema de Parques Nacionales Naturales

de Colombia for their outstanding logistical support

and for generously providing both permission to

work in the park and lodging to V.G. during

this study. Financial support for V.G. and AHS-P

was provided by Idea Wild. This is a contribution

of the Division of Entomology, University of

Kansas.

References

Bogota G. 2002. El polen de la subclase Asteridae en el paramo de

Monserrate. Bogota, Colombia: Universidad Distrital

Francisco Jose de Caldas, Centro de Investigaciones y

Desarrollo Cientıfico. 128 p.

Bogota G, Lamprea SJ, Rangel JO. 1996. Atlas palinologico de la

Clase Magnoliopsida (excepto Asteridae) en el Paramo de

Monserrate. In: Ecoandina, editor. El Paramo ecosistema a

proteger. Bogota, Colombia: CODICE. 131–191.

Danforth BN, Eickwort GC. 1997. The evolution of social

behavior in the augochlorine sweat bees (Hymenoptera:

Halictidae) based on a phylogenetic analysis of the genera.

In: Choe JC, Crespi BJ, editors. The evolution of social

behavior in insects and arachnids. Cambridge: Cambridge

University Press. 270–297.

Eickwort GC. 1969. A comparative morphological study and

generic revision of the augochlorine bees (Hymenoptera:

Halictidae). Univ Kans Science Bull 48:325–524.

Eickwort GC. 1979. A new species of wood-dwelling sweat

bee in the genus Neocorynura, with description of its larva

and pupa (Hymenoptera: Halictidae). Entomol Gen

5:143–148.

Eickwort GC, Eickwort KR. 1972. Aspects on the biology of

Costa Rican halictine bees, IV. Augochlora (Oxystoglossella)

(Hymenoptera: Halictidae). J Kans Entomol Soc 45:18–45.

Engel MS. 2000. Classification of the bee tribe Augochlorini

(Hymenoptera: Halictidae). B Am Mus Nat Hist 250:1–90.

Engel MS, Smith-Pardo AH. 2004. The bee genus

Andinaugochlora in Central America (Hymenoptera:

Halictidae). J Kans Entomol Soc 77:116–120.

Erdtman G. 1986. Pollen morphology and plant taxonomy:

Angiosperms. New York: Hafner Publisher Company. 553 p.

Gonzalez VH, Brown B, Ospina M. 2002. A new species of

Megaselia (Diptera: Phoridae) associated with brood provi-

sions of nests of Neocorynura (Hymenoptera: Halictidae).

J Kansas Entomol Soc 75:73–79.

Gonzalez VH, Engel MS. 2004. The tropical Andean bee fauna

(Insecta: Hymenoptera: Apoidea), with examples from

Colombia. Entomologische Abhandlungen 62(1):65–75.

Harris RA. 1979. A glossary of surface sculpturing. Occas Pap

Entomol California Department of Food and Agriculture

28:1–31.

IGAC 1996. Diccionario Geografico de Colombia. Bogota,

Colombia: Instituto Geografico Agustın Codazzi.

Luderwaldt H. 1911. Nestbau von Neocorynura erinnys Schrottky.

Z Wiss Insektenbiol 7:94–96.

Michener CD. 1977. Nests and seasonal cycle of Neocorynura

pubescens in Colombia. Rev Biol Trop 25:39–41.

Michener CD. 2000. The bees of the world. Baltimore, MD: The

Johns Hopkins University Press. 913 p.

Michener CD, Wille A. 1961. The bionomics of a primitively

social bee, Lasioglossum inconspicuum. Univ Kans Science Bull

42:1123–1202.

Michener CD, Kerfoot WB, Ramırez WB. 1966. Nests of

Neocorynura in Costa Rica (Hymenoptera: Halictidae).

J Kans Entomol Soc 39:245–258.

Moure JS, Hurd PD. 1987. An annotated catalog of the

halictid bees of the Western Hemisphere (Hymenoptera:

Halictidae). Washington, DC: Smithsonian Institution Press.

405 p.

Sakagami SF, Michener CD. 1962. The nest architecture of the

sweat bees (Halictinae): A comparative study of behavior.

Lawrence, KS: University of Kansas Press.

Sarmiento CE. 1993. Abejas y Avispas (Hymenoptera: Apidae,

Vespidae, Pompilidae y Sphecidae) del Santuario Nacional de

Flora y Fauna de Iguaque, Boyaca, Colombia. Bol Mus

Entomol Univ Valle 1:1–11.

Smith-Pardo A. 2005a. Phylogeny and classification of the

bee genus Neocorynura (Hymenoptera: Halictidae,

Augochlorini). PhD dissertation, Lawrence, KS: University

of Kansas. 268 p.

Two new Andean Neocorynura 207

Smith-Pardo A. 2005b. Systematics and mimicry of the

genus Neocorynura: an example of two species from Central

America (Hymenoptera: Halictidae). Acta Zool Cracov

48B(1–2):11–21.

Triplehorn CA, Johnson NF. 2005. Borror and DeLong’s

introduction to the study of insects. 7th ed. Belmont, CA:

Thomson Brooks/Cole. 864 p.

Velasquez RCA. 1999. Atlas palinologico de la flora vascular

paramuna de Colombia: Angiospermae. Colombia:

COLCIENCIAS, Universidad Nacional de Colombia,

Medellın.

Velasquez RCA, Rangel JO. 1995. Atlas palinologico de la flora

vascular del paramo I. Las familias mas ricas en especies.

Caldasia 17:509–568.

208 V. H. Gonzalez et al.