The Bee Family Oxaeidae

with a Revision of the

North American Species

(Hymenoptera: Apoidea)

PAUL D. HURD, JR.

and

E. GORTON LINSLEY

m

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY • NUMBER 220
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SMITHSONIA N CONTRIBUTION S T O ZOOLOG Y • NUMBE R 22 0

The Bee Family Oxaeidae

with a Revision of the

North American Species

(Hymenoptera: Apoidea)

Paul D. Hurd, Jr.

and E. Gorton Linsley

SMITHSONIAN INSTITUTION PRESS

City of Washington

1976
ABSTRAC T

Hurd, Paul D., Jr., and E. Gorton Linsley. The Bee Family Oxaeidae with a

Revision of the North American Species (Hymenoptera: Apoidea). Smithsonian

Contributions to Zoology, number 220, 75 pages, 68 figures, 3 plates, 3 maps, 2

tables, 1976.—This study is a comparative treatment of the natural history of

the family Oxaeidae. Information is presented on the biology, behavior, intrafloral ecology, and systematics of these bees together with a historical review of

their classification and phylogeny. Included are diagnoses of the family and

genera with a revision of the North American species. Characters useful for

identification of the species are illustrated and complete distributional, biological, and synonymical data are provided and discussed. New taxa are Mesoxaea

(type-species: Oxaea nigerrima Friese), Notoxaea (type-species: Oxaea ferruginea

Friese) and Protoxaea australis, P. micheneri, Mesoxaea clypeata, and M.

rufescens.

OFFICIAL PUBLICATION DATE is handstamped in a limited number of initial copies and is recorded

in the Institution's annual report, Smithsonian Year. SI PRESS NUMBER 6129. SERIES COVER DESIGN:

The coral Montastrea cavernosa (Linnaeus).

Library of Congress Cataloging in Publication Data

Hurd, Paul David, 1921-

The bee family Oxaeidae with a revision of the North American species (Hymenoptera, Apoidea)

(Smithsonian contributions to zoology ; no. 220)

Bibliography: p.

Supt. of Docs, no.: SI 1.27:220.

1. Oxaeidae. 2. Insects—Classification. 3. Insects—North America. I. Linsley, Earle Gorton,

1910- joint author. II. Title. III. Series: Smithsonian Institution. Smithsonian contributions

to zoology ; no. 220.

QL1.S54 no. 220 [QL568.08] 591'.08s [595.7'99] 75-619293
Contents

Page

Introduction 1

Acknowledgments 1

Historical Review 3

Comparative Biology of the Family Oxaeidae 5

Flower Preferences 5

Territoriality of Males 11

Male "Sleeping" Aggregations 11

Nest Sites 11

Nest Architecture 12

Female Aggressiveness at the Nest Site 12

Nest Parasites 13

Immature Stages 13

Some Behavioral Characteristics of Adult Protoxaea gloriosa (Fox) 14

Emergence of Males, Establishment of Territories, and Other Premating

Behavior 15

Male "Sleeping" Aggregations 15

Emergence of Females, Feeding and Mating 16

Systematics 16

Family OXAEIDAE 16

Key to the Genera of the Family Oxaeidae 19

Genus Oxaea Klug 19

Notoxaea, new genus 21

Genus Protoxaea Cockerell and Porter 23

Key to the Species of the Genus Protoxaea 24

1. Protoxaea australis, new species 24

2. Protoxaea gloriosa (Fox) 29

3. Protoxaea micheneri, new species 38

Mesoxaea, new genus 41

Key to the Species of the Genus Mesoxaea 43

1. Mesoxaea tachytiformis (Cameron), new combination 44

2. Mesoxaea rufescens, new species 47

3. Mesoxaea vagans (Fox), new combination 51

4. Mesoxaea texana (Friese), new combination 54

5. Mesoxaea nigerrima (Friese), new combination 58

6. Mesoxaea arizonica (Cockerell), new combination and new status . 61

7. Mesoxaea clypeata, new species 64

Literature Cited 67

in

The Bee Family Oxaeidae

with a Revision of the

North American Species

(Hymenoptera: Apoidea)

Paul D. Hurd, Jr.

and E. Gorton Linsley

Introduction

This study is intended as a contribution to our

knowledge of the family Oxaeidae, a group of moderately large, rather robust, fast-flying bees found

only in the Western Hemisphere. Although as a

group the Oxaeidae are primarily centered in the

tropics, a few species occur or are centered in the

warm temperate areas of both continents. None of

the species is known to inhabit the West Indies or

for that matter any other islands.

The group is one for which the taxonomic status

and phylogenetic placement has long been in dispute. Twenty species are presently known, including those described below, and these are equally

divided between North and South America, only

one of which is shared between the continents.

Their known intrafloral relationships suggest a

rather narrow dependence upon relatively few

sources of pollen. These include only certain genera within the plant families Fabaceae, Solanaceae, and Zygophyllaceae. However, the males and

females seek nectar from a comparatively wide variety of plants since the flowers of some of the preferred pollen sources produce little or no nectar

Paul D. Hurd, Jr., Department of Entomology, National

Museum of Natural History, Smithsonian Institution, Washington, D.C. 20560. E. Gorton Linsley, Department of Entomological Sciences, University of California, Berkeley 94720.

and large quantities of nectar are required to meet

the bioenergetic requirements of these fast-flying

bees.

ACKNOWLEDGMENTS.—This investigation has been

made possible and was carried out under the joint

support of our respective institutions, the Smithsonian Institution, Washington, D.C. and the University of California, Berkeley. This arrangement

accorded us not only the opportunity to pursue our

program of research by working independently in

Berkeley and Washington, but also allowed us to

work jointly in Berkeley during the early part of

1975. We therefore express our appreciation to the

authorities of our institutions for their support of

the project and for the use of the requisite facilities

and supporting staffs. For the sponsorship and

financial support of our field and laboratory studies

we acknowledge with gratitude the Leopold Edward

Wrasse and Elvenia J. Slosson Endowment Funds

of the Division of Agricultural Sciences, University

of California, and grants from the Smithsonian Research Foundation (SRF-430039) and the Fluid

Research Fund of the Smithsonian Institution.

Throughout the course of this investigation we

have been ably assisted by Kathleen Sorensen in the

transcription and recording of data, curation of

specimens, preparation of maps, and assembly of

bibliographic materials.

The illustrations which accompany this study

1
SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

were kindly prepared by Celeste Green, Senior

Scientific Illustrator, Department of Entomological Sciences, University of California, Berkeley.

The photographs in Plate 1 were taken by Dr.

Edward S. Ross, California Academy of Sciences,

San Francisco, at 2 miles north of Rodeo, New

Mexico. Plates 2 and 3 are reproduced from

photographs supplied by Ronald F. Shunk, Spring

Valley, California, of a "sleeping" aggregation of

males on Acacia constricta located at the Bagwell

Ranch, 10 miles north of Rodeo, New Mexico.

We are indebted to Andrew R. Moldenke and

John L. Neff, University of California, Santa Cruz,

for providing us with much of the floral data pertaining to Notoxaea ferruginea that we have incorporated into Table 1.

We are grateful to Stanwyn G. Shetler, Department of Botany, Smithsonian Institution, Washington, D.C., for his advice, assistance, and review

of the botanical names recorded in Tables 1 and 2.

He is responsible for the arrangement of the plant

families, genera, and species in these tables and has

also compiled the information on botanical classifications embodied in this study. Accordingly he has

materially enhanced the overall value and usefulness of these tables. Herbert G. and Irene Baker

and Lincoln Constance, University of California,

Berkeley, read the section on flower preferences

and provided helpful comments.

Our investigations on, the systematics of these

bees have been most helpfully facilitated by a number of individuals who have generously given of

their time and accorded us their much appreciated

assistance. This has made it possible for us to examine the primary types of nearly all the North

American oxaeids as well as those of the South

American species described by Friese. Thus we are

indebted to E. Konigsmann, Zoologisches Museum,

Museum fur Naturkunde der Humboldt-Universitat

zu Berlin, for the loan of the holotype of Oxaea

rufa Friese as well as for providing us with specific

information about the Friese type of O. nigerrima

and for help in our attempts to locate the type repositories of O. texana Friese. In this latter regard

we are also grateful to Willem N. Ellis, Zoologisch

Museum, Universiteit van Amsterdam, Max

Fischer, Naturhistorishes Museum, Vienna, and

M. A. Lieftinck, Rhenen, the Netherlands, for their

efforts to locate the type specimens of O. texana

Friese. We are also grateful to Laszlo Moczar of the

Zoological Institute of the University, Szeged, Hungary, who arranged with J. Papp of the Hungarian

Natural History Museum, Budapest, for us to borrow the holotype of O. ferruginea Friese. Likewise

we gratefully acknowledge E. Tortonese, Director

of the Museo Civico di Storia Naturelle "Giacomo

Doria," Genoa, Italy, who kindly arranged with

Del fa Guiglia to compare our notes and a specimen

we sent of Protoxaea gloriosa (Fox) with the type

specimens of O. tristis Gribodo. The results of her

much appreciated assistance have contributed significantly to our studies. We are also indebted to

Karl V. Krombein, National Museum of Natural

History, Smithsonian Institution, Washington,

D.C., who kindly examined for us the label data on

the holotype of O. tachytiformis Cameron, and we

also thank G. Knerer, currently in residence at the

British Museum of Natural History, for additional

information that he supplied in this regard.

We would like to express our appreciation to the

following individuals who have helped to make

this study possible by the loan of more than 2500

specimens from their personal collections or from

collections in their care: Paul H. Arnaud, Jr.,

California Academy of Sciences, San Francisco

(CAS); G. E. Bohart, Bee Biology and Systematics

Laboratory, Agriculture Research Service, U.S. Department of Agriculture, Logan, Utah (GEB);

Horace R. Burke and Gordon A. Frankie, Texas

A & M University, College Station (TAM); M. A.

Cazier, Arizona State University, Tempe (ASU),

George E. Eickwort, Cornell University, Ithaca

(CU), Marjorie Favreau, American Museum of

Natural History, New York (AMNH); Saul Frommer, Entomological Teaching and Research Collection, University of California, Riverside (UCR);

James E. Gillaspy, Texas A & I University, Kingsville (TAI); David F. Hardwick, Canadian National Collections of Insects, Arachnids, and

Nematodes, Biosystematics Research Institute, Research Branch, Agriculture Canada, Ottawa

(CNC); Gary F. Hevel, National Museum of Natural History, Smithsonian Institution, Washington, D.C. (using herein the acronym USNM for

the former United States National Museum); Wallace E. LaBerge, Illinois Natural History Survey,

Urbana (INHS); U.N. Lanham, University of

Colorado Museum, Boulder (U. Colo.); John F.

Lawrence, Museum of Comparative Zoology, Harvard University, Cambridge (MCZ); C D. Mich-
NUMBER 220

ener, University of Kansas, Lawrence (KU); W.

Wayne Moss, Academy of Natural Sciences, Philadelphia (ANSP); J. A. Powell and J. A. Chemsak,

California Insect Survey, Essig Museum of Entomology, University of California, Berkeley (CIS);

Roy R. Snelling, Los Angeles County Museum of

Natural History (LACM); Robbin W. Thorp,

University of California, Davis (UCD); P. H.

Timberlake, University of California, Riverside

(PHT); Floyd G. Werner, University of Arizona,

Tucson (UA) and Thomas J. Zavortink, San Francisco, California (TJZ).

Historical Review

Klug (1807) proposed the genus Oxaea for a

new species from Brazil, O. flavescens, of which he

illustrated the male. He considered the genus

unique in having three-, rather than two-segmented

labial palpi, a term which he apparently applied

only to the short apical segments (in Oxaea and its

immediate relatives, the first segment of the palpus

is elongate and flattened, the last three segments

short, cylindrical, together not as long as the first).

He also noted that the maxillary palpi were absent,

a condition he believed to be the case in Melecta,

Melipona, and Acanthopus, although he did not

attempt to assign Oxaea to a position near these or

any other genus of bees.

Latreille (1810) was the first to attempt to place

Oxaea in a classification. He presumably knew only

the male, and included the genus among the parasitic bees near Melecta and Crocisa, as did Blanchard (1845). Many years later, Popov (1941, 1945,

1951) reported that the oxaeids, which he regarded

as a distinct family, and the parasitic anthophorids

of the genera Osiris, Pasites, Morgania, Schmiedeknechtia, and Neopasites (representing the tribes

Osirini, Ammobatini and Neopasitini), resemble

one another in having undifferentiated sagittae in

the male genitalia (penis valves not separated from

the penis). However, since this condition also occurs in the Andrenidae, Megachilidae, and Apidae

(Snodgrass, 1941), Popov considered that the causes

of morphological reduction in the sagittae were

different in various groups. Rozen (1964) also

found similarities between the third instar larvae

of Protoxaea gloriosa and those of Nomada-like

anthophorids. He wrote:

The similarities between the mature larva of P. gloriosa and

those of the Nomada-like parasitic anthophorids (Nomada,

Triepeolus, Oreopasites, etc.) are difficult to explain. Not

only do the larvae of these two groups share the generalized

features also encountered in the Colletidae, Andrenidae,

Halictidae, and Melittidae, but they both also exhibit certain

specialized characteristics, namely, the peculiar position of

the posterior tentorial pits and the absence of antennal

prominences and vertexal protuberances. Although, with the

mature larvae, the long mandible of P. gloriosa contrasts

with the short mandible of the parasitic bee, the long mandible of the first instar of P. gloriosa is somewhat similar to

the extremely long mandible of the first stage parasitic forms.

The mature larva of P. gloriosa is especially similar to that

of Triepeolus sp. (I have examined the same series used by

Michener, 1953). In addition to the characters referred to

above, these taxa share: spiculated hypopharynx; attenuate

mandibular apexes (though not blade-like in Triepeolus sp.);

body without tubercles; spiracles on pigmented, sclerotized

plates; and long atrial spines. Presumably the similarities

result from convergent evolution, for the adults of Oxaeinae

and the Nomada-like bees are too dissimilar to suggest a

close relationship.

A number of authors have considered oxaeids

(or at least Oxaea) related to the carpenter bees

(Anthophoridae: Xylocopinae). For example, F.

Smith (1854) placed them between Xenoglossa and

Melitturga near Xylocopa and Lestis, Sichel (1865)

considered their true position to be nearer the end

of the tribe Xylocopites of Lepeletier (1841), and

Blanchard (1845) and Ashmead (1899) assigned

them to the Xylocopidae, stating that Oxaea and

Lestis probably should be considered as a subfamily, the Oxaeinae.

Another relationship that has been suggested for

Oxaea is with the diphaglossine colletids. Dalla

Torre (1896) placed them in the Apidae, subfamily

Podaliriinae, between Diphaglossa and Lestis. Cockerell (1898) assigned the genus to the podaliriine

Anthophoridae between Centris and Megacilissa,

and Friese (1898) in his monograph of the genera

Megacilissa, Caupolicana, Diphaglossa, and Oxaea,

considered the genus to be a derivative of the

caupolicanine branch of the Diphaglossinae, related in particular to Megacilissa, which it resembles in habitus, coloration, and the extensive

pollen-collecting apparatus. He also derived the

Old World genus Melitturga from this group of

colletids. Schrottky (1902) placed Oxaea next to

Megacilissa (in the Andrenidae) but separated

both genera from Caupolicana and Diphaglossa

(Megacilissa is presently regarded as a synonym of

Caupolicana). Strand (1910) placed Oxaea next to
SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

Oxaea Ptiloglossa Diphaglossa

Caupolicana Pasiphae Stenocolletes

Lonchopria

Collates

FIGURE 1.—Phylogenetic relationships of Oxaeidae (Oxaea),

after Schrottky (1913).

Ptiloglossa, and Jorgensen (1912a, b) listed it at the

end of the Colletidae following Caupolicana and

Ptiloglossa. Ducke (1912) keyed out Oxaea next to

the Diphaglossinae, but did not include it in that

group. Schrottky (1913) discussed the interrelationships among the South American genera of

Colletidae, but also excluded Oxaea, regarding it

as a very aberrant genus, which to him apparently

originated in tropical Brazil and was only slightly

related to the colletids. His diagram of the relationships among the genera he discussed is reproduced

in Figure 1. The most recent statement regarding a

possible diphaglossine relationship is that of Roberts (1973) who comments:

The nests most similar architecturally to those of the Oxaeidae are those of the Diphaglossinae (Colletidae) such as

Ptiloglossa (Roberts, 1971). Although similar in some aspects

of adult morphology and behavior, fundamental differences

in both adult and pre-imaginal morphology proscribe attempts to link Oxaea and Protoxaea with the Diphaglossinae.

My observations on pupal morphology and nest architecture

of Oxaea flavescens corroborate the conclusions of Rozen

(1965) based on Protoxaea, and indicate that the Oxaeidae

have no close relatives within the Apoidea.

The more prevalent taxonomic view of the

group has been that they are andrenids. This possibility was first suggested by Gerstaecker (1867),

who rejected a close relationship with Lestis, as

proposed by Sichel (1865), on the basis of differences in wing venation, mouthparts, pollencollecting apparatus and form of the abdomen,

emphasizing similarities in labial structure to the

Andreninae, and to Andrena in particular in the

form of the cubital cell, the fringe on the sides and

apex of the abdomen and the strongly developed

coxal "curl." Friese (1898) agreed in general with

Gerstaecker and proposed assigning them to the

Melittinae behind Megacilissa and before Systropha. However, the strongest support for an andrenid relationship was the character analysis of

Bischoff (1934) which revealed a similarity between Oxaea and genera of andrenine and panurgine bees through the possession of subantennal

plates. This feature also impressed Michener who

characterized the subfamily Oxaeinae in the Andrenidae for the reception of Oxaea and Protoxaea,

noting some similarities to the Old World genus

Melitturga. His treatment was endorsed by Graf

(1966), who rejected separate family status for the

group, in particular the interpretation of Rozen

(see below). Graf, like Michener, relied heavily on

the subantennal sutures, but discussed other characters, including the reduction of the sting, a feature shared with most andrenine and panurgine

Andrenidae.

The most cogent arguments for regarding the

oxaeids as a distinct family has been provided by

Rozen (1965). We quote from him, in part, as

follows:

Michener (1944) divided the andrenid subfamily Panurginae

into two tribes, the Panurgini and Melitturgini, with the

latter containing the single Old World genus Melitturga.

This genus was relegated to tribal status apparently on the

grounds that the adults, unlike those of other panurgines,

bear certain striking resemblances to the essentially Neotropical Oxaeinae of the same family. In 1951 Rozen showed

that the male genitalia of Melitturga are unlike those of the

Oxaeinae and are not only typical of those of the Panurginae

in general but quite like those of the CamptopoeumPanurgus-Panurginus-Epimethea complex within the subfamily. On the basis of this information, Michener (1954)

abandoned the idea that the genus Melitturga represents a

distinct tribe of the Panurginae. Recently evidence in the

form of the larva of Protoxaea gloriosa Fox (Rozen, 1964)

suggested that the Oxaeinae were so unlike other Andrenidae

that they should be removed from the family unless some

form intermediate between the two subfamilies is found. In

spite of the structure of the male genitalia, Melitturga is

the only known possible intermediary ... . As an adult,

Af. clavicornis and presumably the other members of the

genus possess the following oxaeine-like features: body size

large (though not so large as that of the Oxaeinae); compound eyes of male large and strongly converging above;

ocelli, especially those of male, low on face first flagellar

segment as long as scape; other antennal segments short;

thorax robust and in male with dense pile; wings strongly

papillate.

However, the Oxaeinae differ from all the Panurginae,

including Melitturga, on the basis of numerous characters,

The features of the larvae and the genitalia have already

been mentioned. Michener (1944) gave the following signifi-
NUMBER 220

cant characteristics of adult oxaeines: apex of posterior

femur of female enlarged and forming a plate; scopa abundant from coxa to basitarsus; pterostigma absent; and marginal cell narrow and elongate. To these should be added:

clypeus strongly protuberant; labrum as long as, or longer

than, broad, and submentum fused into a single plate.

It is important to point out that the Oxaeinae differ from

the Andreninae on the basis of all the characters indicated

above except that the andrenine larvae lack conical tubercles

(though the transverse ones on the anterior end of the body

are more pronounced than those of the Oxaeinae). Also

females of the Andreninae have the scopa distributed from

the coxa to the basitarsus, as in the Oxaeinae. However, both

of these characters are almost certainly plesiomorphic, for

they are found in the Colletidae and are thus not significant

for a judgment of phylogenetic relationships.

On these bases, Rozen reaccorded the oxaeids

family status and we concur in this judgment.

Comparative Biology of the Family Oxaeidae

FLOWER PREFERENCES.—The oxaeids, in so far as

presently known, are narrowly polylectic, utilizing

related or counterpart pollen sources in the northern and southern hemispheres, especially species of

Cassia (Fabaceae) and Solarium (Solanaceae).

These plants produce an abundance of pollen but

no nectar, and are attractive only to pollengathering females which, like the males, must seek

nectar elsewhere. Species in these taxonomically

distant genera share convergent structural and

functional similarities in the method of pollen

presentation which are also approximated in such

remote groups as Tibouchina, Rhynchanthera, and

Tococa (Melastomataceae), (Laroca, 1970) and

Pedicularis (Scrophulariaceae) and Dodecatheon

(Primulaceae), vide Macior, 1971, 1974. Although

morphological details differ, they all shed their

pollen, which is dry, through terminal staminal

pores which must be "buzzed" and vibrated or

squeezed by the bee in order to release it. Perhaps

because of the abundance of the pollen, which

squirts out in clouds onto the venter of the abdomen and can be quickly gathered into the scopal

hairs, most bee visitors are large forms whose

weight inverts the flower, for example species of

Bombus, Xylocopa, Centris, Caupolicana, and Ptiloglossa (see Linsley,. 1962; Michener, 1962; Linsley and Cazier, 1963; Wille, 1963). In fact, Wille

states that in the Province of Guanacaste, Costa

Rica, Cassia biflora is called "abejon," meaning

"large bee," presumably because of the large conspicuous bees which visit it. However, inversion of

the flower, although facilitating the rapid capture

of large amounts of pollen, is not necessary for the

release of pollen, and these same authors have recorded small to medium-sized bees such as species of

Exomalopsis, Psaenythia, Augochloropsis, and Nomia, successfully gathering pollen. Not all bees,

however, vibrate or squeeze the anthers. In addition to the "buzzing bees," which include oxaeids,

Wille (1963) recorded "biting bees," which chew

into the anthers and "gleaning bees," which pick

up pollen left on the corolla by other bees. (For an

informative discussion of the behavior of bees at

flowers of Cassia, see Thorp and Estes, 1975).

Neither Cassia nor Solarium flowers produce nectar, and although Todd (1882) states that those of

Solarium have no odor, Harris and Kuchs (1902)

observed that "especially in the morning, the odor

was decidedly pronounced." Other distinctions

shared by at least some species of Cassia and

Solarium are the production of both "right-handed"

and "left-handed" flowers (Todd, 1882; Robertson,

1890; Harris and Kuchs, 1902) and two kinds of

stamens, a single large stamen in addition to the

four short ones in Solarium rostratum (but not all

species of Solarium) and two long stamens in addition to the shorter ones in some species of Cassia.

Although the functional interpretation of these

two kinds of stamens has been subject to different

interpretations (see Todd, 1882; Muller, 1883;

Meehan, 1886, Robertson, 1890; Halsted, 1890; and

Harris and Kuchs, 1902), the smaller anthers in

Cassia are now known to produce viable pollen, the

larger ones (the so-called "food anthers") smaller,

sterile pollen grains (Meeuse, 1961). However,

Bowers (1975) found that the dimorphic anthers

in Solarium rostratum perform a different function: the primary role of the small anther is to

provide pollen as an attractant to insect visitors

and the larger anther functions in pollination.

Linsley and Cazier (1972) reported that of a sample

of 2565 pollen grains of Cassia bauhinioides removed from the scopal hairs of Protoxaea gloriosa,

approximately 20 percent were large grains averaging about 80 microns in diameter and nearly 80

percent were small grains averaging about 27 microns, which, although not viable, contained protoplasm. Solarium also produces two sizes of pollen

grains, but Bowers (1975) found no detectable dif-
SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

ferences in the percentage of abortive grains between the large and small anthers within a flower,

and hand-pollination experiments demonstrated no

differences in pollen fertility between the two types

of anthers. Stow (1927) found a correlation in 5.

tuberosum between their presence and high temperatures, and Joergensen (1928) reported a high

proportion of "bad" pollen grains in experimentally produced euploid and aneuploid forms. However, since Heiser and Whitaker (1948) found that

specimens of S. elaeagnifolium from California had

the chromosome number n = 12, and were therefore diploid, it is possible that aborted pollen in

this species results from hybridization, as in the S.

nigrum complex (Stebbins and Paddock, 1949).

On the other hand, samples taken from bees gathering pollen from a pure stand of S. elaeagnifolium

near Portal and a stand mixed with S. rostratum at

Douglas, Cochise County, Arizona, each yielded

about 35 percent viable grains averaging about 35

microns in diameter and 65 percent lacking protoplasm and averaging about 26 microns in diameter

(Linsley, 1962; Linsley and Cazier, 1963). Solarium

rostratum growing in the mixed stand at Douglas

yielded about 85 percent viable pollen grains in

bee-collected samples. However, these particular

characteristics of pollen manufacture and presentation are not found in other plants known to be

utilized as pollen sources by oxaeids, all of which

also produce nectar.

Cazier and Linsley (1974) have commented on

the fact that Protoxaea gloriosa, although polylectic

as a species, either as individual females or as whole

populations may be more or less regular polyleges

constant to one of a variety of diverse plant species

(during the course of the nesting season the preferences may change in response to sequential

blooming of hosts). The species most commonly

selected not only represent an array of plant families, e.g., Fabaceae (Cassia, Prosopis, and Desmanthus), Zygophyllaceae (Kallstroemia and Larrea),

and Solanaceae (Solarium), but the flowers involved differ markedly in form, structure, and color

(at least to the human eye). Further the average

diameter of the pollen grains which they produce

ranges from about 20 microns to 80 microns, a

range which includes large numbers of plant species occurring in the territory of the bee which are

not known to be utilized by the bee.

Cazier and Linsley (1974) have described some

of the unique features of pollen and nectar presentation in Kallstroemia grandiflora and the following is extracted in part from their account:

[The flowers of Kallstroemia grandiflora] produce an abundance of pollen and nectar, which is exploited in three ways

by insects . . . including bees, wasps, flies and butterflies.

The plant is of special interest from the standpoint of

pollination ecology, as both the plant and some of the insects

have adaptations in behavior that are not always mutually

beneficial. The flowers of K. grandiflora are allogamous for

most of the daily flowering period and become autogamous

as the flowers close. Thus, they can be either cross- or selfpollinated, with the former evidently favored. The flowers

have little or no odor, a color range within "bee yellow"

(500-650 \L), and a strong ultraviolet reflective pattern on

both upper- and under-surfaces. Insects are apparently attracted by the color and ultraviolet reflection, with the latter

so arranged as to form non-reflective, dark target, "nectar

guide" areas on both upper- and under-surfaces.

One group of bees and wasps gather pollen and nectar

from the top of the flower, effecting both cross- and selfpollination in the process. A second group of smaller bees

and wasps gather nectar from within the flower but avoid

contact with the sexual portions and are therefore of no

importance in pollination. The third group of honeybees

and several other larger bees and wasps, extract nectar from

the underside of the flower completely avoiding the sexual

parts of the plant and therefore play no role in pollination.

Even though the plant gains no direct benefit from this last

group of insects it nevertheless provides them with a sepal

nectar guide that directs them to the nectaries. It is proposed that these species may be contributing to the economy

of the plant by reducing the quantity of available nectar so

that pollinators have to visit more flowers to get their full

nectar supply.

This conclusion is in line with that of Heinrich

and Raven (1972) that "a flower must provide

sufficient reward to attract foragers, but it must

limit this reward so that the animals will go on to

visit other plants of the same species," although the

presumed method for accomplishing this in Kallstroemia is different from any which they describe

or suggest.

Cazier and Linsley (1974) have reported on the

method utilized by Protoxaea gloriosa for the collection of pollen and nectar from the flowers of

Kallstroemia.

[Females] alight in the upturned flowers which droop downward with the bees weight, sometimes bending 180 degrees

so that the bee is working upside down. As they land in the

flowers, facing downward, they gather the style and all ten

stamens into the venter, holding them there with all six legs.

This places the anthers, which are abundantly covered with
NUMBER 220

Name

Oxaea festiva

Ducke, 1901

Oxaea flavescens

Schrottky, 1907, 1908;

Ducke, 1910

Oxaea austera

Schrottky, 1904, 1907;

Ducke, 1910

Notoxaea ferruginea

Jorgensen, 1909, 1912a, b

TABL E 1.—Flower preferences

Pollen sources *

SOLANAGEAE

Solatium grandiflorum

FABACEAE

Cassia bicapsularis

Cassia occidentalis

Cassia Sp.

Crotalaria paulina

FABACEAE

Cassia sericea

Cassia sp.

SOLANACEAE

Solanum grandiflorum

Solanum occarpum

Solanum atropurpureum

FABACEAE

Cassia aphylla

Cassia bicapsularis

Cassia occidentalis

Crotalaria paulina

ZYGOPHYLLACEAE

Larrea cuneifolia

Larrea divaricata

SOLANACEAE

Solanum elaeagnifolium

Solanum sp.

of South American oxaeids

Nectar sources?*

Unknown

Unknown

LAMIACEAE

Leonurus sibiricus

NYCTAGINACEAE

Allonia incarnata

CAPPARACEAE

Atamisquea emarginata

FABACEAE

Acacia furcatispina

POLYGALACEAE

Monnina lorentziana

ZYGOPHYLLACEAE

Tribulus terrestris

Larrea cuneifolia

BORAGINACEAE

Heliotropium mendocinum

SOLANACEAE

Grabowskia boerhavifolia

Nectar sources<f

Unknown

Unknown

LAMIACEAE

Leonurus sibiricus

PORTULACACEAE

Portulaca confertifolia

CAPPARACEAE

Atamisquea emarginata

FABACEAE

Acacia furcatispina

Prosopis torquata

Cercidium praecox

Melilotus alba

ZYGOPHYLLACEAE

Larrea cuneifolia

BORAGINACEAE

Heliotropium mendocinum

SOLANACEAE

Grabowskia boerhavifolia

ASTERACEAE

Verbesina encelioides

Verbesina microptera

Tessaria Sp.

*Not including pollen sources also used for nectar.

large orange pollen grains, and the stigma, which is covered

with stigmatic fluid, in contact with the abdominal venter

and hind legs.

As soon as the bees' hold on the style and stamens is secured, the mouthparts are inserted into the nearest nectary

for from a fraction of a second to about one second, presumably dependent on the amount of nectar present. From

one or two to all five nectaries are visited and on several

occasions six mouthpart insertions are made, the first nectary

being overlapped. After each nectary is sampled the bee

shifts its hold on the style and stamens, shaking them rather

violently, as its body moves around to a position from which

the mouthparts can contact the next nectary. This shifting

and shaking motion of the bee in rotating its position completely around the inside of the flower would account for

the large amounts of loose pollen found throughout much

of the body of the bee and occasionally in the bottom of

the corolla. In Protoxaea each stop in the rotary movements

is made at the juncture between two petals beneath which

is a nectary. These are small, elongate (rectangular) dark

green spongy-looking masses of tissue situated middorsal at

the base on the elongate, narrow, hairy sepals issuing from

beneath the area where the petals overlap.

By late morning the movements of females on exposed
SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

TABLE 2.—Flower preferences of North American oxaeids

Name

Protoxaea australis

Linsley & Michener, 1962

Protoxaea gloriosa

Crawford, 1915;

Linsley & Michener, 1962;

Linsley & Cazier, 1963, 1972;

Cazier & Linsley, 1974;

Hurd & Linsley, 1975

Pollen Sources ?

FABACEAE

Cassia Sp.

FABACEAE

Acacia angustissima

Desmanthus cooleyi

Prosopis glandulosa

Cassia bauhinioides

Cassia leptocarpa

Cassia wislizenii

ZYGOPHYLLACEAE

Kallstroemia grandiflora

Larrea tridentata

SOLANACEAE

Solanum elaeagnifolium

Solatium rostratum

Nectar Sources **

Unknown

CASUARINACEAE

Casuarina Sp.

CAPPARACEAE

Koeberlinia spinosa

BRASSICACEAE

Lepidium montanum

FABACEAE

Acacia angustissima

Prosopis glandulosa

Caesalpinia jamesii

Hoffmanseggia glauca

Dalea scoparia

Medicago sativa

Nelilotus alba

ZYGOPHYLLACEAE

Kallstroemia grandiflora

Larrea tridentata

LOASACEAE

Cevallia sinuata

OLEACEAE

Menodora scabra

ASCLEPIADACEAE

Asclepias subverticillata

VERBENACEAE

Aloysia gratissima

LAMIACEAE

Salvia azurea

ASTERACEAE

Baccharis glutinosa

Conyza coulteri

Guardiola tulocarpus

Verbesina encelioides

Gaillardia pulchella

Nectar Sources <f

Unknown

CASUARINACEAE

Casuarina Sp.

CAPPARACEAE

Koeberlinia spinosa

BRASSICACEAE

Lepidium lasiocarpum

FABACEAE

Acacia angustissima

Mimosa Sp.

Prosopis glandulosa

Caesalpinia jamesii

Hoffmanseggia glauca

Parkinsonia aculeata

Medicago sativa

Melilotus alba

ZYGOPHYLLACEAE

Kallstroemia grandiflora

EUPHORBIACEAE

Croton pottsii

Croton sp.

RHAMNACEAE

Condalia Sp.

MALVACEAE

Gossypium Sp.

Sphaeralcea

TAMARICACEAE

Tamarix gallica

Tamarix ramosissima

LOASACEAE

Cevallia sinuata

Mentzelia sp.

OLEACEAE

Menodora scabra

ASCLEPIADACEAE

Asclepias subverticillata
NUMBER 220

TABLE 2.—Flower preferences of North American oxaeids—(Continued)

Name

Protoxaea micheneri

Linsley & Michener, 1962

Mesoxaea tachytiformis

Mesoxaea rufescens

Mesoxaea vagans

Mesoxaea texana

Cockerel1, 1910,

1933, 1934a

Pollen Sources '

ZYGOPHYLLACEAE

Kallstroemia Sp.

Unknown

FABACEAE

Mimosa polyantha

SOLANACEAE

Solatium Sp.

Unknown

SOLANACEAE

Solanum rostra turn

Nectar Sources **

Unknown

Unknown

EUPHORBIACEAE

Croton culiacanensis

TURNERACEAE

Turners diffusa

LOGANIACEAE

Buddieja sessiliflora

ASTERACEAE

Verbesina encelioides

CAPPARACEAE

Wislizenia refracta

ASCLEPIADACEAE

Asclepias sp.

POLYGONACEAE

Polygonum Sp.

MALVACEAE

Gossypium sp.

Nectar Sources <>

VERBENACEAE

Aloysia gratissima

Aloysia wrightii

LAMIACEAE

Salvia azurea

ASTERACEAE

Baccharis glutinosa

Conyza coulteri

Solidago occidentalis

Guardiola tulocarpus

Helianthus annuus

Verbesina encelioides

Baileya multiradiata

Gaillardia pulchella

Psilostrophe tagetina

Lygodesmia juncea

Unknown

Unknown

FABACEAE

Acacia angustissima

Mimosa polyantha

ZYGOPHYLLACEAE

Kallstroemia grandiflora

EUPHORBIACEAE

Croton culiacanensis

TURNERACEAE

Turnera diffusa

LOGANIACEAE

Buddleja sessiliflora

CAPPARACEAE

Wislizenia refracta

ZYGOPHYLLACEAE

Larrea tridentata

ASCLEPIADACEAE

Asclepias Sp.

ASTERACEAE

Unidentified

POLYGONACEAE

Polygonum Sp.

EUPHORBIACEAE

Croton Sp.
10

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

TABLE 2.—Flower preferences of North American oxaeids—(Continued)

Name

Mesoxaea nigerrima

Mesoxaea arizonica

Mesoxaea clypeata

Pollen Sources ?

ZYGOPHYLLACEAE

Kallstroemia sp.

ZYGOPHYLLACEAE

Kallstroemia grandiflora

Unknown

Nectar Sources **

LAMIACEAE

Monarda punctata

PEDALIACEAE

Sesamum indicum

FABACEAE

Sesbania macrocarpa

TURNERACEAE

Turnera diffusa

LOGANIACEAE

Buddieja sessiliflora

FABACEAE

Sesbania macrocarpa

Nectar Sources <t

VERBENACEAE

Aloysia gratissima

Verbena halei

LAMIACEAE

Monarda sp.

ASTERACEAE

Vernonia texana

Eupatorium serotinum

FABACEAE

Cassia Sp.

ZYGOPHYLLACEAE

Kallstroemia sp.

VERBENACEAE

Vitex pyramidata

FABACEAE

Acacia angustissima

Tephrosia leiocarpa

ZYGOPHYLLACEAE

Kallstroemia grandiflora

TURNERACEAE

Turnera diffusa

LOGANIACEAE

Baddleja sessiliflora

Unknown

*Not Including pollen sources also used for nectar.

flowers are very rapid because most of these blossoms have

been already visited. Very often the bee then samples only

one, two, or three of the nectaries before flying to the next

flower. When an obscured, previously unvisited flower is

found, either inside the plant or beneath another plant in

the shade, females slow their flight and make a more complete visitation.

In southeastern Arizona, the principal pollen

competitors of Protoxaea at Cassia, Larrea, and

Solarium are large, fast-flying matinal colletids of

the genera Caupolicana and Ptiloglossa. However,

these bees start gathering pollen from this group of

plants at dawn and only partially overlap the activity period of Protoxaea, which on a clear day

starts about sunrise and continues for an hour or

two after the colletids have completed pollen collecting, although in the later period there may be

competition from species of Bombus, Xylocopa,

and Centris. The caupolicanine colletids do not

visit Kallstroemia, the flowers of which do not open

until an hour or more after sunrise, and we have

not seen them at Desmanthus. On Kallstroemia

Protoxaea is the largest pollen collector and dominates the flowers (Plate 1).

Protoxaea gloriosa is a matinal pollen gatherer,

and unlike some early morning bees of the families

Colletidae, Halictidae, and Anthophoridae, the

females do not return to the flowers for pollen in

the evening, even from plants such as Larrea tridentata, which make pollen available throughout

the day (Hurd and Linsley, 1975).

It is interesting to note that some of the South

American caupolicanine colletids which occur
NUMBER 220 11

within the range of oxaeids presumably also take

pollen from some of the same plant genera as those

in North America, for example, Caupolicana ruficollis (Friese) and C. bicolor (Friese) from Larrea

divaricata. Nectar plants from genera in common

include Hoffmannseggia for Caupolicana ruficollis

(Friese), C. lugubris (Friese), C. friesei Jorgensen,

and C. bicolor, and Baccharis for C. lugubris. Caupolicana is best represented in warm temperate and

subtropical regions of North and South America

but is poorly represented or absent in the intervening zones (Michener, 1966). The genus is apparently best represented in Chile where oxaeids

are absent.

On the other hand, although some Ptiloglossa in

South America take pollen from Solanum, according to Schrottky (1908), in Paraguay Pt. matutina

(Schrottky) visits Solanum balbisii and S. paniculatum, but not Cassia, which is visited by Oxaea

flavescens in the same area.

The known flower preferences of oxaeid bees as

interpreted from literature and supplemented with

previously unpublished records are summarized in

Tables 1 and 2. (For details of unpublished records

for North American oxaeids, see the full data from

specimens examined in the taxonomic accounts of

each species.) In Tables 1 and 2, the plant families

are arranged systematically according to the latest

Englerian classification as revised by Hans Melchior (1964, Engler's Syllabus der Pflanzenfamilien,

II, 666 pages, Berlin-Nikolassee: Gebriider Borntraeger). The generic arrangement within families

is alphabetical except in two cases. In the Fabaceae,

the genera are alphabetical within their subfamilies, which are ordered systematically as follows:

Mimosoideae, Caesalpinioideae, Faboideae. In the

Asteraceae, the genera are alphabetical within

their tribes, which are ordered systematically (according to Melchior, 1964) as follows: Vernonieae,

Eupatorieae, Astereae, Inuleae, Heliantheae, Heleniae, Anthemideae, Senecioneae, Calenduleae,

Arctoteae, Cardueae, Mutisieae, and Cichorieae.

Inasmuch as the practice of using only regular endings on the names of families is gaining among

plant taxonomists, the authors have elected to follow this trend and unlike Melchior, use the following regular names instead of the indicated irregular

ones: Asteraceae for Compositae, Brassicaceae for

Cruciferae, Fabaceae for Leguminosae, and Lamiaceae for Labiatae.

TERRITORIALITY OF MALES.—A behavioral activity

which appears to be a biological characteristic of

the Oxaeidae is male territoriality. Schrottky

(1905) mentions the hovering flight of males of

Oxaea in front of flowers in a manner suggestive of

tabanids or syrphids. Roberts (1973) has reported

that males of Oxaea flavescens Klug were aggressive

in the vicinity of flowers. Linsley and Michener

(1962) refer to territorial behavior in Protoxaea

[P. gloriosa (Fox)] and Mesoxaea [M. nigerrima

(Friese)] and fuller discussions are provided by

Cazier and Linsley (1963) and Linsley and Cazier

(1972). Essentially this behavior consists of guarding plants or groups of plants from all intruders

except their own pollen-collecting females. At trees

and shrubs the males hover in flight facing the

plant at various levels depending upon its size,

darting out to chase other insects and returning to

the same territory immediately. When guarding

annuals or low-growing plants they may poise above

them or between them. In any case, periodically

they leave the territory to obtain nectar to provide

the energy to sustain the activity, usually from the

plant or plants they are guarding if they are in

bloom (Plate 1)

Cazier and Linsley (1963) speculated that the

principal function of territoriality in these bees is

to assure the immediate mating of newly emerged

females through spacing and coverage of appropriate nectar plants. Because nesting females are

ignored by males, they suggested that territorial

belligerence toward other insects might reduce

competitive pressure on females, although they

noted that at least in some situations territorial

activity of males is out of phase with pollencollecting periods of females, especially at plants

whose pollen is exploited early in the morning.

MALE "SLEEPING" AGGREGATIONS.—Sleeping aggregations of males are well known for Protoxaea

gloriosa (see below) but do not appear to have

been reported for other oxaeids. Thus it remains to

be determined whether or not this is a family characteristic. Similar behavior has been recorded for

many genera of colletids (see Rayment, 1935;

Linsley and Cazier, 1972) and other bees.

NEST SITES.—In so far as known, oxaeids nest in

flat or slightly sloping ground. Roberts (1973)

found two nests of Oxaea flavescens in Colombia.

They were about 10 meters apart in a sparsely

grassy pasture a few hundred yards from an oxbow
12

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

lake, each entrance being concealed by a small

clump of grass. Bertoni (1911) also found the nest

entrances of Oxaea austera Gerstaecker hidden at

the bases of clumps of grass. On the other hand,

Mesoxaea texana (Friese) apparently nests in areas

of sparse vegetation with the entrances exposed

(Cockerell, 1933). Protoxaea gloriosa, although

sometimes nesting in grassy areas (Linsley and

Cazier, 1972), usually also has the entrance

exposed.

Roberts (1973) found Oxaea flavescens nesting

in homogeneous red, sandy clay devoid of rocks.

According to Birkmann (Cockerell, 1933) the surface soil at a nesting site of Mesoxaea texana was

sandy and deep grayish but the soil excavated from

the burrows (presumably where the cells were located) was of a brick red color. The only known

burrow of Mesoxaea nigerrima was in flat, hard

soil (Linsley and Michener, 1962). Protoxaea gloriosa was found nesting in sandy soil with gravelly

outcroppings (Linsley and Cazier, 1972).

Oxaea fuscescens Sichel is apparently a solitary

nester as is Mesoxaea nigerrima. Truxal (1962)

reported only seven burrows of an unidentified

species of Oxaea in an area 25 by 30 feet. Burrows

of Protoxaea gloriosa may be separated by less than

a meter but concentrations usually reflect population pressure on limited suitable soil resources and

not gregarious tendencies. However, Birkmann,

1932 (Cockerell, 1933), estimated that about 80,000

burrows of Mesoxaea texana were contained in an

area about 700 feet in length by 230 feet in width.

NEST ARCHITECTURE.—The nest architecture of

oxaeids is of the kind regarded by Michener and

Lange (1957) as the basic primitive type of subterranean bee nest. It is most similar to that of

diphaglossine Colletidae (Claude-Joseph, 1926;

Janvier, 1933; Roberts, 1971). Typically, the burrows are vertical, more or less straight, apparently

less so in Mesoxaea texana according to Birkmann

(Cockerell, 1933), with horizontal laterals which

are filled with soil when completed, each of which

terminates in a single vertical cell. Cell provisions

in Mesoxaea nigerrima were semiliquid and yellowish brown, filling the bottom one-third or onefourth of the cell (Linsley and Michener, 1962).

The burrows are deep, 58 cm in the single

known nest of Mesoxaea nigerrima, about 30-65

cm in several burrows of Protoxaea gloriosa (Cazier and Linsley, unpublished), and a remarkable

245 cm with laterals 20 to 50 cm long in a nest of

Oxaea flavescens excavated over a two-day period by

Roberts (1973). He speculated that the nest might

be perennial:

The wings and mandibles of the female captured leaving

the burrow showed very little wear. Thus, I cannot believe

she excavated the burrow herself. Although the volume of

soil removed from the nest amounts to 0.26 liters in compact

form (probably twice that as excavated granules) there was

no tumulus at the nest entrance. Oxaea flavescens is morphologically adapted for carrying especially small pollen grains.

It seems unlikely that the female could have carried away

the relatively large soil granules. Yet, it seems equally unlikely that wind and rain could have effectively dispersed so

much soil within the short life of the female occupying the

nest. Judging by her unworn wings and mandibles, she was

only 10-14 days old. Thus, she was probably the sister (not

the mother) of the viable prepupae and pupae associated

with the nest.

It would seem that the first, or one of the first, females to

emerge at the beginning of the dry season claims her natal

nest as her own. By so doing, she needs only to deepen the

burrow a little and excavate a lateral burrow. Most of the

soil excavated from each subsequent lateral is packed into

the previous lateral. The small excess of soil which must be

ejected from the nest is soon dispersed by wind and rain.

After a few years the nest could easily attain a depth of 2.5

meters. Of course, new nests founded by females forced to

leave their pre-empted natal nests should be much shallower.

If fluctuations of soil moisture or greater susceptibility to

parasitism increases pre-imaginal mortality in shallow nests,

then selection should favor the tendency to re-occupy and

deepen existing nests. Perhaps such a tendency explains why

this nest of O. flavescens is the deepest bee nest on record.

FEMALE AGGRESSIVENESS AT THE NEST SITE.—

Truxal (1962) reported aggressiveness at a nest

site of an unidentified species of Oxaea in eastern

Peru as follows:

Each individual had apparently established a territory in the

immediate vicinity of its burrow and was aggressively protecting this area. Individuals hovered about three feet above

the burrow openings, and all flying objects that approached

to within three or four feet would immediately be engaged

and chased away at a very high rate of speed. They also

engaged individuals of their own species in "combat." These

encounters never lasted for more than 10 to 15 seconds; then

the "owner" returned to its position above the burrow.

Occasionally, upon returning to their burrows, the bees

entered them and often remained below ground for several

minutes. Seven specimens were easily taken because of their

aggressive attitude toward the net. All specimens taken

proved to be females.

Roberts (1973) observed similar aggressive behavior in females of O. flavescens. Cockerell (1933)
NUMBER 220 13

records an experience of the Rev. G. Birkmann

(1932) at a large nesting site of Mesoxaea texana

where the "bees flew with such energy and speed

that they would not try to avoid anyone standing

in the way, but struck Mr. Birkmann's body and

hat." At nest sites of Protoxaea gloriosa, Linsley

and Cazier (1972) noted that females returning to

their burrows with loads of pollen enter quickly,

ignoring the presence of an observer, but when returning from midday nectar flights they commonly

strike the observer about the face and head and

enter the burrow less directly.

Since oxaeids have a reduced stinging apparatus,

"bumping" invaders of their nesting area may be

their principal means of defense.

NEST PARASITES.—Few positive data are available

on the parasites of Oxaeidae. Schrottky (1904) reported finding Thalestria smaragdina Smith flying

to flowers with males of Oxaea in Paraguay and

Brazil and speculated that this species of Thalestria

might be parasitic in the nests of Oxaea (he also

commented on the similarity in coloration of the

females). Subsequently, Bertoni (1911) observed

Thalestria smaragdina flying about the nests of

Oxaea austera in Paraguay.

A large and apparently undescribed species of

Triepeolus commonly lays its eggs in cells of Protoxaea gloriosa in the San Simon Valley of southeastern Arizona and western New Mexico. Rozen

(1966) has characterized the larva of this species

and that of a closely related species from the nests

of Ptiloglossa jonesi Timberlake at Portal, Cochise

County, Arizona.

IMMATURE STAGES.—Rozen (1964) described the

first stage and the post-defecating larva of Protoxaea gloriosa from a site one mile north of Rodeo,

Hidalgo County, New Mexico, as follows (format

slightly modified):

First stage larva: Head capsule and mouthparts

apparently without setae though sensilla may be

present, integument unpigmented except for raandibular apexes. Tentorium complete but very

weak; posterior tentorial pits situated below posterior thickening and hypostomal ridge; anterior

pits somewhat above epistomal groove; posterior

thickening of head capsule well defined, similar in

thickness to hypostomal ridge; pleurostomal ridge

moderately broad but very weak, scarcely noticeable; epistomal ridge not evident but external

groove present; parietal bands apparently absent.

Antennae moderately developed, perhaps a little

more pronounced than those of mature larva.

Labral tubercles large; labral apex cleft; labrum

bearing spicules laterally but not apically nor on

epipharyngeal surface. Mandibular corium nonspiculate; each mandible simple at apex, elongate,

bearing small, sharp-pointed teeth scattered along

upper and lower apical edges. Maxillae and labium

greatly fused, reduced, and recessed; maxillary

palpi evident but short; labial palpi evident but

even shorter than those of maxillae; salivary opening a short tranverse slit. Body form elongate,

cylindrical, without tubercles but with intrasegmental lines at least anteriorly. Integument nonsetose but minutely spiculate. Spiracular atrium not

pigmented and not projecting above body wall,

without elongate spines though perhaps with very

short teeth near primary tracheal opening; peritreme apparently missing; primary tracheal opening not on collar and circular in outline, that is,

not transverse as in mature larva; subatrium unpigmented, differing little in appearance from

trachea. Abdominal segments 9 and 10 normal,

without protuberances or ridges; anus apical.

Postdefecating larva: Head integument wrinkled,

with scattered sensilla; epipharyngeal surface of

labrum and hypopharynx spiculate; entire head

including mandibular corium pigmented; parietal

bands, all internal ridges, antennal rings and papillae, apex of labrum, apexes of mandibles, salivary

gland opening, and palpi more darkly pigmented

than other parts; coronal cleavage line and posterior part of labiomaxillary region less pigmented

than other parts. Tentorium complete but weak;

anterior and posterior arms darkly pigmented; remainder unpigmented; parietal bands, all internal

ridges, antennal rings and papillae, apex of labrum,

apexes of mandibles, salivary gland opening, and

palpi more darkly pigmented than other parts;

coronal cleavage line and posterior part of labiomaxillary region less pigmented than other parts.

Tentorium complete but weak; anterior and posterior arms darkly pigmented; remainder unpigmented; each posterior tentorial pit lying below

juncture of posterior thickening and hypostomal

ridge but joined to posterior thickening by spur of

former; anterior tentorial pits lying just above

epistomal thickening; posterior thickening of head

capsule moderately thin and resting slightly anterior to posterior margin of capsule; thickening
14 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

gradually bending forward immediately above juncture of spur to posterior tentorial pit and joining

hypostomal ridge; hypostomal ridges, pi euros tomal

ridges, and epistomal ridges of about the same

thickness as posterior thickening; epistomal ridge

briefly interrupted medially; longitudinal thickening of head capsule absent; parietal bands well

defined. Vertex nonprotuberant; clypeus somewhat

protuberant. Antennae not arising from prominences, small; each papilla small and lower than

basal diameter; each antenna bearing numerous

(perhaps 7) sensilla. Labrum cleft apically and

bearing two tubercles, which are beset with sensilla,

some of which are on small prominences. Each

mandible in adoral view extremely broad basally

but tapering rapidly to very thin, elongate, nearly

parallel-sided apex; when seen from above or below, mandible moderately narrow basally and

gradually tapering to moderately broad apex, consequently apex of mandible bladelike; extreme

apex simple when viewed adorally but, when

viewed from above or below with slender, sharppointed tooth on adoral side; adoral apical edge

beset with numerous sharp-pointed teeth (this edge

seems to be the dorsal inner apical edge); there is

no indication of a ventral inner edge); somewhat

smaller pointed teeth on both dorsal and ventral

surfaces in cuspal region. Labiomaxillary region

fused, reduced, and recessed, so that only notable

features are palpi and salivary gland opening;

maxillary palpi conspicuous, a little longer than

basal diameter, and with some sensilla on small

prominences; labial palpi much smaller than maxillary palpi; other sensilla on apex of labium on

small prominences; salivary opening a small curved

slit partly enclosing a low prominence. Body form

tapering and curving anteriorly to relatively small

head; posterior part nearly straight but blunt, due

at least in part to somewhat telescoped abdominal

segments 8, 9, and 10 (the compression results from

the larva's resting on the tip of its abdomen in a

vertical cell). Mesothorax and metathorax dorsally

divided into cephalic and caudal annulets; other

segments not noticeably divided on postdefecating

form; tubercles very faint, transverse, presumably

on anterior part of caudal annulations on predefecating form. Integument rigid, nonspiculate, finely

wrinkled, and yellowish, with intersegmental lines

more darkly pigmented. Spiracles resting on elevated, pigmented sclerites, peritreme flat; atrium

projecting above body wall, atrial wall darkly pigmented, beset with numerous hairlike spines, and

without rim; primary tracheal opening collared

and slitlike, with slit at right angles to long axis of

body; collar with numerous short spines; subatrium

pigmented just below atrium and near attachment

of trachea but with area in between unpigmented;

subatrial wall bearing spines immediately below

atrium. Except for telescoping, abdominal segments

9 and 10 normal, without protuberances, or ridges;

anus a transverse slit located apically.

Roberts (1973) who found the larva of Oxaea

flavescens comments about it as follows:

Rozen noted that the larva of Protoxea gloriosa differs

from all other bee larvae in possessing: (1) an apically cleft

labrum; (2) a reduced labiomaxillary region; (3) a long

mandible with a long blade-like apex; (4) a slit-like primary

tracheal opening. The larva of Oxaea flavescens possesses

these features and agrees in every detail with the description

of P. gloriosa. Thus, it seems that oxaeid larvae are as

similar morphologically as oxaeid adults.

The pupa of Oxaea flavescens has been described and figured by Roberts (1973). Pupae of

other oxaeids are as yet unknown. Roberts' description and comments follow:

Head: antenna with small tubercle (a) on outer surface

of pedicel. Mesosoma: pronotum with posterior lobes (b)

moderately produced; metanotum with small indistinct,

median tubercle (c). Each coxa and trochanter produced in

large apical spine (d); anterior femur with large basal spine

(e) and base of middle femur slightly produced; each femur

with rounded tubercle (f) anteriorly at apex, moderately

large on anterior femur to very large on posterior femur.

Metasoma: terga II-V each with small, irregular tubercles

(g) in sub-apical row; terminal spine (h) short and rounded

apically.

According to Yager and Rozen (1966), the pupae of the

Andrenidae possess mesoscutal tubercles, as do the pupae of

the Fideliidae (Rozen, 1970). The pupa of Oxaea flavescens

lacks mesoscutal tubercles. It may be distinguished from

pupae of the Colletidae by the presence of distinctive apical

tubercles on metasomal terga II-V, from the pupae of most

Megachilidae by its lack of setae, and from the pupae of the

Halictidae by the lack of tubercles on its wing. These differences, together with the unique apical tubercles on its

femora, support the decision to accord Oxaea and Protoxaea

familial rank.

Some Behavioral Characteristics of Adult

Protoxaea gloriosa (Fox)

Since the adult behavior of Protoxaea gloriosa is

better known at present than that of any other
NUMBER 220 15

oxaeid, a few aspects are elaborated upon for future

comparison with representatives of other genera as

data become available. For more details see Linsley

and Cazier (1972).

EMERGENCE OF MALES, ESTABLISHMENT OF TERRITORIES, AND OTHER PREMATING BEHAVIOR.—At least

in southeastern Arizona and western New Mexico,

moisture from the first summer rains, which usually

appear in the latter part of July or early August,

appears to be the key factor stimulating emergence.

As in most species of solitary bees, the males emerge

for the season several days before the females. They

immediately seek nectar, remaining at a single

flower for several minutes if the supply is adequate.

Favored nectar sources both for newly emerged

individuals and those in established territories are

Koeberlinia spinosa, Baccharis glutinosa, and Asclepias subverticillata, as well as most pollen plants

utilized by the female, including Kallstroemia

grandiflora and Larrea tridentata.

Male territorial behavior has been described by

Linsley and Michener (1962), Cazier and Linsley

(1963), Linsley and Cazier (1972), and Hurd and

Linsley (1975). The observations reported were

made mostly in the San Simon Valley near Portal,

Arizona, and Rodeo, New Mexico, or in the vicinity

of Douglas, Arizona. In the San Simon Valley, the

plant usually selected for the establishment of early

season territories is Koeberlinia spinosa, in those

areas where it is in bloom. This is a favored nectar

source for newly emerging females during this period. Subsequently, such shrubs as Baccharis glutinosa and Larrea tridentata are sought out by later

emerging females (the latter also an important pollen source for nesting females).

In open areas where females are active, territories are established near one or more low-growing

plants, such as Cassia bauhinioides and Kallstroemia grandiflora (both pollen plants for the female) and Ephedra trifurea, Lycium sp., Gutierrezia microcephala, and Cucurbita foetidissima

(which are not pollen sources for the female; at C.

foetidissima, territorial males keep away such regular pollen seekers as Peponapis pruinosa).

When guarding their territories, males attack

and often tumble to the ground with other male

Protoxaea but usually ignore pollen-collecting

(mated, nonreceptive) females when guarding a

pollen source. However, they fly out and pursue or

attack larger insects, including butterflies, and even

birds, but when small bees or wasps are abundant

about the flowers they are usually not molested.

MALE "SLEEPING" AGGREGATIONS.—Massed nocturnal aggregations of males have been reported by

Evans and Linsley (1960) and Cazier and Linsley

(1963) and fully described by Linsley and Cazier

(1972). The phenomenon is notable since the

males which are strongly antisocial during that

portion of the day in which they are attempting to

establish or are occupying territories become highly

social when congregating together for the night.

Male territorial behavior usually reaches a peak

near midmorning when most newly emerged, receptive females leave their natal burrows for their

first nectar flight. By midafternoon the males begin

to gather in large masses on selected individual

plants where they spend the night in a cluster

(Plates 2,3). The factors that influence the initial

selection of the cluster site are unknown, but they

include such diverse situations as the open empty

valves of the dried dehiscent pericarps of Datura

quercifolia, the upright stems of such weedy plants

as Heterotheca (telegraph-plant), and Amaranthus

(pig-weed), the slender, jointed branches of

Ephedra (Mormon tea), the horizontal leafy

branches of Condalia spathulata (squaw bush),

and Acacia constricta (white thorn) as well as

deserted bird nests, especially those of the verdin

(Auriparus flavipes).

Once the congregating site is selected, it is

stained with a yellow excrement and has a distinctly

noticeable odor. Linsley and Cazier (1972) reported

that of 344 males from one aggregation which were

marked with distinctive colors and released at various distances, five returned in the late afternoon

from a release site one mile away, four from three

miles distant, two from a distance of five miles, and

one from ten miles.

Most aggregations contain several hundred males

(Linsley and Cazier report one made up of more

than 2000 individuals), but in some cases smaller

separate clusters occur on the same plant near one

another. The numbers are largest when the males

first emerge and dwindle from attrition as the season progresses. Temperatures in the cluster fluctuate during the night (Linsley and Cazier report

ranges from 20.5° to 25.5°C with air temperatures

ranging downward from 23.5° to 18°C). When disturbed the cluster temperatures rise suddenly by 8°

to 10°C.
16 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

EMERGENCE OF FEMALES, FEEDING AND MATING.—

As mentioned above, as the season begins males

emerge several days before the females and are

established in territories by the time the first females appear. The females leave their natal burrows near midmorning (0830-1000 hrs.) and immediately seek nectar. Since on the initial flight

they sip at the flower for several minutes, they are

quickly found by males. Mating takes place on the

flowers and lasts for several minutes. Linsley and

Michener (1962) have described the action as

follows:

At 8:47 a.m. on July 26, a mating pair of P. gloriosa was

found hanging on the Baccharis blossoms, with a second

(smaller) male on top of the pair and apparently trying to

dislodge the first male to the accompaniment of a loud

buzzing. The female was taking nectar and was apparently

freshly emerged, since the wings were still soft and unfrayed

and her pubescence showed no signs of wear. On each of the

next two days, mating pairs were observed in the same

situation; in each case the female was freshly emerged and

in each there was an extra male atop the mating pair. During mating, the male forces the head between the propodeum

and first metasomal segment, grasping the base of the latter

with the mandibles. In one instance, mating was observed

for 5 minutes and 30 seconds from the time the pair was

discovered. When disturbed sufficiently, the female flies off

with the male hanging on, even though not in sexual contact. In no case was the second male successful in dislodging

the first, and in most instances there was a wide discrepancy

in size between the two.

Mating has been observed on the flowers of

Koeberlinia spinosa and Kallstroemia grandiflora

as well as those of Baccharis.

Systematics

In the accounts that follow we have presented

diagnoses of the family and genera with a revision

of the North American species belonging to the

genera Protoxaea and Mesoxaea which are centered

in Mexico, but extend into the southwestern United

States (Louisiana, Texas, New Mexico, and Arizona) and nearly to Central America (southwestern Chiapas, Mexico). The genus Notoxaea is

monotypic and is known to occur from the Province of Mendoza, Argentina, into south-central

Brazil (Mato Grosso). The species of the genus

Oxaea are chiefly centered in the Guiano-Brazilian

subregion, but one species {Oxaea fuscescens Sichel) evidently ranges from northern South AmerFICURE 2.—Wings of Oxaeidae, Protoxaea gloriosa (Fox).

ica well into Central America where it has been

recorded from Guatemala.

Most of the members of this family are moderately large, robust bees, often with thick, pale

pubescence on the thorax and in some species the

metasomal terga are brilliantly metallic, while in

others the terga are scarcely or not at all metallic.

The wings (Figure 2) are frequently infuscated

apically and in some species are entirely dark or

nearly so.

Family OXAEIDAE

OXAEINAE Ashmead, 1899:70.—Cockerell and Porter, 1899:

411.—Michener, 1944:241, 246; 1965:10.—Moure, 1950:

304.—Rozen, 1951:148.—Moure and Lucas de Oliviera,

1962:42.—Linsley and Michener, 1962:385.—Rozen, 1964:

223.—Michener, 1965:10.—Graf, 1966:137-138.

OXAEIDAE Cockerell, 1933:288; 1934a:8; 1936:6.—Rayment,

1935:610.—Popov, 1941:82; 1945:329, 336; 1951:895.—

Moure, 1944:4-5; 1946:12; 1947:226-227.—Rozen, 1965:

16.—Stephen, Bohart, and Torchio, 1969:45, 50.—Linsley

and Cazier, 1972:2-3.—Roberts, 1973:437.—Michener, 1974:

37.

Head: Integument without metallic luster; antennal sockets distinctly below middle of eyes; antennae short, elbowed; flagellum much shorter

than length of eye, first segment slender, elongated,

about as long as, or longer than, scape, remaining

segments enlarged broader than long except terminal segments longer than broad; eyes greatly en-
NUMBER 220 17

FIGURES 3-8.—Heads of Oxaeidae in frontal view, males: 3, Protoxaea gloriosa (Fox); 4, P.

australis Hurd and Linsley; 5, Mesoxaea clypeata Hurd and Linsley; 6, M. tachytiformis (Cameron); 7, Notoxaea ferruginea (Friese); 8, Oxaea flavescens Klug.
18 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

larged in males (Figures 3-8); inner orbits of males

converging strongly above (Figures 3-8); ocelli located low on face near antennal sockets, arranged

in a broad triangle about twice as broad as long;

epistomal suture biangularly arched, separated

from antennal sockets by less than diameter of

sockets; subantennal areas defined by an arched

outer suture which is longer than inner suture and

meets antennal socket near middle of side, and an

inner nearly straight suture which meets antennal

socket at lower inner margin; facial foveae absent;

face above hind ocelli strongly differentiated (especially in female) from adjacent lateral areas by a

vertical sulcus extending dorsally from each hind

ocellus and face between sulci densely and confluently punctured to vertex or nearly so; clypeus

strongly protuberant, projecting about as far as

lower width of eye when seen in profile, lateral

margins below tentorial pits, more or less feebly

concave, true dorsolateral angles situated at anterior tentorial pits; lower lateral portions of clypeus

strongly bent to rear, forming, as seen from below,

an emargination for reception of labrum in repose;

malar area not present; rear angle of mandibles

behind posterior eye margin; labrum about as long

as broad, or longer; mentum and submentum not

represented by distinct sclerites; glossa short, slender, several times as long as broad, acutely pointed;

flabellum absent; labial palpi elbowed, basal segment elongated and broadly flattened, longer than

remaining three subcylindrical, subequal segments

together; stipites with posterior margins entire,

without a row or comb of spines.

Thorax (including propodeum): Integument

without metallic luster; preepisternal suture not

evident below scrobal suture; scutellum gently

rounded or nearly flat, almost horizontal in profile;

propodeum declivous, without a well-defined basal

area; wings (Figure 2) with pterostigma scarcely

evident; marginal cell long and narrow, considerably longer than distance from apex to wing tip,

apex bent away from wing margin; three submarginal cells, third largest and longest, first and second subequal in length along posterior margins,

second not distinctly narrowed anteriorly; first recurrent vein interstitial with second transverse cubital vein; second recurrent vein received well beyond middle of third submarginal cell; wing surface

with only a few large patches of hairs in basal area,

distal area strongly papillate; jugal lobe of posterior wing long, more than three-fourths length of

vannal lobe; intermediate coxae elongate but appearing short because basal portions are concealed

by pleurae at least externally, much shorter than

distance from summits to posterior wing bases;

pollen-collecting hairs of female (see also abdomen)

present on propodeum laterally, coxae, trochanters,

femora, tibiae, and basitarsi; middle femora with a

subtriangular, densely compacted brush of short

orange bristles, basally on ventral surface and with

a similar brush of matching bristles also at apex of

middle trochanters ventrally in females; posterior

femora of female enlarged and flattened to form a

plate associated with, and similar to, basitibial

plate; basitibial plates located at bases of posterior

tibiae; posterior basitarsi shorter than tibiae; inner

posterior tibial spurs longest; tarsal claws cleft,

inner and outer rami of subequal length in the

males, inner rami about half as long as outer ones

in females; tarsal arolia present but reduced in

males, scarcely evident in females.

Abdomen (metasoma): Tergal integument usually metallic (scarcely if at all in Notoxaea and

without metallic luster in Protoxaea micheneri and

Oxaea rufa), apical margins impunctate, glabrous

and shining on terga I-V in males and I-IV in females; graduli present, laterally strongly bent posteriorly and passing above and behind spiracles;

first sternum angularly emarginate at middle;

pollen-collecting hairs of females present on ventral surfaces of first and second terga and medially

on sterna I—III, although that on III very much

smaller; pygidial plate present in female; male

genitalia with capsule tapering basally so that

gonobase is small, gonostyli incompletely differentiated from gonocoxites or not evident; penis valves

not separated from penis (Figures 12, 16, 28, 60).

DISCUSSION.—It is readily evident from the foregoing diagnosis that this family is distinguished

from the other families of bees by a number of

unique characteristics. Among these are the structure of the antennae, the position of the ocelli of

the males in relation to the antennal sockets, the

strongly differentiated face above the hind ocelli,

the venation, the brushes of orange bristles located

basally on the ventral surface of the middle femora,

the apical modification of the hind femora of the

female, and the structure of the male genitalia. On

the other hand there are a number of features

which are shared with one or more families of bees,
NUMBER 220

not the least of which is the presence of a subantennal area defined by two subantennal sutures.

However, to emphasize this condition or any other

single character believed to be of fundamental importance in determining phylogeny, and hence the

classification of the oxaeids, is to ignore the evidence obtained by viewing the constellation of all

characters. Thus in retrospect it is quite apparent

that in many previous classifications, as we have

discussed elsewhere in this study, seemingly undue

reliance upon one or only a few anatomical features has resulted in classifications in which the

oxaeids have at one time or another been assigned

to all the families of the Apoidea except the Halictidae and the Megachilidae.

Our analyses of the characters presented above

reveal that apart from those unique to the oxaeids,

the majority of characters are shared with the family Colletidae. It would thus appear to us from this

19

indication as well as from the known overall similarities in their biologies that the oxaeids and the

colletids were derived from a common ancestry.

It may be of some phylogenetic significance that

among all the Apoidea only males of the colletid

genus Ctenocolletes (Stenotritinae) and those of

the Oxaeidae share an eighth metasomal sternum

which has a large, exposed, hairy disc. These

groups also share such other features as two subantennal sutures below each antennal socket, elongated first flagellar segments of the antennae which

are about as long or longer than the scape, the lack

of preepisternal sutures below the scrobal sutures,

and the ocelli are situated low on the face, much

nearer the antennal sockets than to the posterior

margin of the vertex.

The family is composed of four genera, which

may be distinguished in the following key.

Key to the Genera of the Family Oxaeidae

1. Maxillary palpi present, six-segmented; gonostyli evident, partially differentiated from

gonocoxites (Figures 16, 20, 28, 40, 44, etc.) 2

Maxillary palpi absent; gonostyli not evident (Figure 12) Oxaea

2. Mandibles simple apically (Figures 3-6, 8); metasomal terga brownish-black or black, not

partly or largely reddish 3

Mandibles bidentate apically (Figure 7); metasomal terga I-III and sometimes IV at least

partly or largely reddish Notoxaea

3. Eighth metasomal sternum of male entire, not emarginate medially on apical margin (Figures

19, 23, 27, 31, 35, 39); metasomal terga VI (males) and V (females) without long, conspicuous tufts of white hairs at sides Protoxaea

Eighth metasomal sternum of male deeply omarginate medially on apical margin (Figures

43, 47, 51, 55, 59, 67); metasomal terga VI (males) and V (females) with long, conspicuous

tufts of white hairs at sides Mesoxaea

Genus Oxaea Klug

FIGURES 8-12

Oxaea Klug, 1807:261.—Latreille, 1809: 172; 1810:338; 1811:

588; 1825:464.—Olivier, 1809:588— Blanchard, 1845:411.—

Sichel, 1865:331.—Gerstaecker, 1867:318.—Dalla Torre,

1896:297.—Cockerell, 1898:42.—Friese, 1898:78, 1899:239.—

Ashmead, 1899:70-71.—Cockerell and Porter, 1899:411.—

Schrottky, 1902a:413; 1907:13; 1913:238—Ducke, 1912:60,

82.—Bischoff, 1934:330.—Popov, 1941:83.—Michener, 1944:

241.—Moure, 1944:4-5; 1946:7, 8, 12-13, fig. 1.—Moure and

Oliviera, 1962:4— Moure and Seabra, 1962:235.—Graf,

1966:137.—Linsley and Cazier, 1972:2, 3.—Roberts 1973:

437. [Type-species: Oxaea flavescens Klug, monotypic]

Dasyglossa Illiger, 1807:217; 1810:44.—Sandhouse, 1943:544.

[Type-species: Oxaea flavescens Klug, monotypic]

DIAGNOSIS.—Males usually with pale maculations

on clypeus, labrum, mandibles, and basal segments

of antennae; mandibles simple apically (Figure 8);

maxillary palpi absent; metasomal terga brownish

black, black or partly or largely reddish; metasomal

terga VI (males) and V (females) with long, conspicuous tufts of white or black hairs at sides;

eighth metasomal sternum of male deeply emarginate medially on apical margin (Figure 11);

gonobase greatly reduced, ringlike, much wider

than long, not tapering basally and gonostylus not

evident (Figure 12).

DISCUSSION.—The lack of maxillary palpi and the

absence of partially differentiated gonostyli in the

male (Figure 12) uniquely serve to distinguish the

genus Oxaea from the other genera of the Oxaeidae. These features and others clearly indicate that
20

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

10

FIGURES 9-12.—Oxaea flavescens Klug, male (S. Ben to, Duque de Caxias, Rio, Brazil): 9, seventh

metasomal tcrgutn (left half ventral); 10, 11, seventh and eighth metasomal sterna (left halves

dorsal); 12, genitalia (left half ventral).

Oxaea has departed more from the ancestral archetype than have the genera Mesoxaea, Notoxaea,

and Protoxaea, all of which have retained the sixsegmented maxillary palpi and have the gonostyli

of the males partially differentiated from the gonocoxites (Figures 16, 28, 60). It may be significant

that while Oxaea is centered in the GuianoBrazilian subregion (Moure and Seabra, 1962:235)

and occurs as far north as Central America (Moure

and Urban, 1963:361), the other genera of the

family occupy peripheral ranges either well to the

north of Oxaea in North America {Mesoxaea and

Protoxaea) or chiefly to the south of it in South

America {Notoxaea). Since these genera, as is dis-
NUMBER 220 21

cussed elsewhere in this study, are more nearly related to one another than they are to Oxaea, it may

be that the ranges of their precursors were ecologically preempted by the evolutionary events which

have led to the formation of Oxaea. If so, this could

explain why Mesoxaea, Notoxaea, and Protoxaea

occupy ranges peripheral to that of the genus

Oxaea.

In coloration, the integument of several metasomal terga in some species of Oxaea is largely reddish and nearly without metallic green reflections

(e.g., O. rufa Friese), while in other species the

terga of the females are largely metallic green and

their males have only the marginal depressions of

the terga either brilliantly metallic green (e.g., O.

fiavescens Klug) or black and nearly without metallic reflections (e.g., O. schwarzi Moure and Seabra). In some species the eyes of the males approximate closely on the vertex (e.g., O. alvarengai

Moure and Urban), but in other species (e.g., O.

fiavescens) the eyes are widely separated dorsally

(Figure 8). Unlike the females of Mesoxaea, Notoxaea, and Protoxaea which have the surfaces of

the mesoscutum, scutellum, and metanotum closely

punctured and dull, these sclerites in the females of

several species of Oxaea are broadly impunctate

and shining.

As mentioned previously, only one of the eight

species now included in the genus Oxaea is known

to be present in Central America. This species, O.

fuscescens Sichel (1865:342), was originally described from Caracas, Venezuela, and has subsequently been reported from the Guianas (Cockerell,, 1917a: 135) and Guatemala (Moure and Urban,

1963). It is easily recognized from the original

description and the color illustration provided by

Sichel (1865:342).

NAMES CURRENTLY APPLIED IN Oxaea.—Although

the studies of Moure (1944, 1947, et seq.) have contributed substantially to an understanding of the

systematics of oxaeids in general and Oxaea in particular, the genus has not been revised and there is

no key available for the identification of the included species. We have listed below the names

currently applied in the genus. All were proposed'

in Oxaea and at present all are considered to represent valid species.

alvarengai Moure and Urban (1963:362)

austera Gerstaecker (1867:318)

festiva Smith (1854:316)

fiavescens Klug (1807:262)

fuscescens Sichel (1865:342)

rufa Friese (1899:244)

schwarzi Moure and Seabra (1962:236)

stenocoryphe Moure (1947:223)

Notoxaea, new genus

FIGURES 7, 13-16

Oxaea.—Friese, 1898:78-85 [in part; not Klug, 1807:261].

Protoxaea.—Moure, 1944:4-5; 1947:226. [in part; not Cockerell and Porter, 1899:410, 411].—Moure and Seabra, 1962:

235.

TYPE-SPECIES.—Oxaea ferruginea Friese, 1898,

present designation.

DIAGNOSIS.—Males with pale maculations on clypeus and mandibles; mandibles bidentate apically

(Figure 7); maxillary palpi present, six-segmented;

metasomal terga I—III and sometimes IV at least

partly or largely reddish; metasomal terga VI

(males) and V (females) with long, conspicuous

tufts of white hairs at sides; eighth metasomal

sternum of male deeply emarginate medially on

apical margin (Figure 15); gonobase greatly reduced, ringlike, much wider than long, not tapering

basally and gonostyli evident, partially differentiated from gonocoxites (Figure 16).

DISCUSSION.—The correct name for the single included species has presented some problems because

although Schrottky (1907:19) placed his earlier

described Oxaea versicolor Schrottky (1903:45) as

a junior synonym of Oxaea ferruginea Friese

(1898:83), he did so believing that these were the

sexes of the species as it occurred in Asuncion,

Paraguay (ferruginea, the male and versicolor, the

female). However, this association has been open to

some question since ferruginea was described from

northeast Brazil (Piauhy) by Friese (1898:83)

while versicolor was named from specimens (12 9)

which Schrottky (1903:45) believed had been collected in northern Argentina or in Brazil. Since

Moure and Seabra (1962:235) state that ferruginea

is ".. . known from Mendoza, Argentina to southern Mato Grosso, Brazil," and since the type locality

of ferruginea given by Friese (1898:83) lies well to

the northeast of this, then three possibilities exist:

(1) the species ranges or occurs farther northward

in Brazil than Moure and Seabra (1962:235) indicate; (2) the species described by Friese (1898:83)

from the Brazilian State of Piaui (Piauhy) is dif-
22

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

13 16

14 15

FIGURES 15-16.—Notoxaea ferruginea (Friese), male (La Rioja, Argentina): 13, seventh metasomal

tergum (left half ventral); 14, 15, seventh and eighth metasomal sterna (left halves dorsal); 16,

genitalia (left half ventral).

ferent from that which occurs in Argentina, Paraguay, and northward into Mato Grosso, Brazil, and

(3) the type locality cited by Friese (1898:83) for

ferruginea is erroneous.

Our examination of the Friese type specimen of

ferruginea reveals unequivocally that it represents

the same species that Schrottky (1907:17) and subsequent investigators believed it to be. However,

the type specimen bears a locality label (hand

printed in black ink) which reads: "Tianhy, Brasilia 1889." In addition, there are three other labels,

a Friese identification label and two labels which

contain hand-printed, black ink arabic numbers:

856/22 and 9, respectively. The identification label
NUMBER 220 23

is lowermost on the pin and reads: Oxaea ferruginea $ (in Friese's handwriting in black ink) followed by det Friese 1897 (machine printed) and

below which Friese has written in black ink:

"Piauhy-Brasilia." Clearly Friese is responsible for

making the decision that "Tianhy, Brasilia" equals

"Piauhy-Brasilia." We have been unable to discover

the reason which prompted Friese to make this

change and likewise we have been unable to locate

"Tianhy" in any of the geographic place name

sources available to us. It may be of some significance that among the 25 other specimens of oxaeid

bees loaned to us for study from the collections of

the Hungarian Natural History Museum by Dr. J.

Papp, there are three specimens bearing identical

Tianhy, Brasilia, 1889 locality labels. These have

been identified by Friese as: Oxaea austera Gerstaecker (1 $ ) and O. flavescens Klug (2 $ ). There

is also another specimen of ferruginea ($ ) in that

collection which has the same locality label as the

type specimen (Tianhy, Brasilia), but it lacks a

Friese identification label. Presumably this specimen, which is only about 13 mm in length, was not

seen by Friese since he cites only a single male, 15

mm in length.

CONCLUSIONS.—From the foregoing discussion we

conclude that while Schrottky (1907:17) correctly

associated the sexes of ferruginea, it is still doubtful

whether or not the species occurs in northeast

Brazil. It may be that there existed in 1889 a

"Tianhy" or equivalent geographic place name in

that region, but it also appears equally, if not more

probable, that such a place name may have existed

in southern Brazil. If so then the type specimen

could well have been collected within the currently

known range of the species as defined by Moure

and Seabra (1962:235).

The single known specimen ( 9) of Oxaea rufa

Friese (1899:244) from northeast Brazil (Para) has

the metasoma almost entirely reddened and thus

bears a somewhat similar appearance to ferruginea.

However, our examination of this specimen not

only confirms the conclusion of Schrottky (1907:17)

that it is a larger and different species, but unlike

ferruginea it lacks maxillary palpi and belongs to

the genus Oxaea.

The synonymy of Notoxaea ferruginea has been

reviewed by Moure (1944:4-5; 1947:226), who assigned the species to the North American genus

Protoxaea primarily because the maxillary palpi

are six-segmented. Although Notoxaea shares a

number of characteristics with the North American genera Mesoxaea and Protoxaea, including the

six-segmented maxillary palpi, it is readily separated from these, as well as from the genus Oxaea,

by its apically bidentate mandibles.

NAMES CURRENTLY APPLIED IN Notoxaea.—Listed

below are the names applied in the genus Notoxaea and, except for argentina which was first assigned to the genus Ptiloglossa, were originally

proposed in Oxaea. Only the name ferruginea is

currently considered valid since the others have

been shown to be junior synonyms.

argentina (Jorgensen, 1909:221), new combination

ferruginea (Friese, 1898:83), new combination

haematura (Cockerell, 1918:252), new combination

versicolor (Schrottky, 1903:45), new combination

Genus Protoxaea Cockerell and Porter

FIGURES 2-4, 17-40; MAP 1; PLATES 1, 2

Megacilissa.—Fox, 1893:421 [in part; not Smith, 1854:123].

Oxaea.—Authors [not Klug, 1807:261].—Cockerell, 1898:42.

Protoxaea Cockerell and Porter, 1899:410, 411.—Cockerell,

1933:288; 1934b: 152.—Michener, 1944:246.—Linsley, 1951:

1086.—Mitchell, 1960:20, 21.—Linsley and Michener, 1962:

385-388.—Moure and Seabra, 1962:235.—Linsley and Cazier,

1972:2, 3.

TYPE-SPECIES.—Megacilissa gloriosa Fox, monotypic and original designation.

DIAGNOSIS.—Males without pale maculations on

clypeus, labrum, mandibles, and antennae; mandibles simple apically (Figures 3, 4); maxillary

palpi present, six-segmented; metasomal terga

brownish black or black, not partly or largely reddish; metasomal terga VI (males) and V (females)

without long, conspicuous tufts of white hairs at

sides; eighth metasomal sternum of male entire, not

emarginate medially on apical margin (Figures

19, 27, 39); gonobase elongate, not ringlike, much

longer than wide, tapering basally and gonostyli

evident, partially differentiated from gonocoxites

(Figures 20, 28, 40).

DISCUSSION.—Unlike the genus Oxaea which lacks

maxillary palpi, the genus Protoxaea, like the

genera Mesoxaea and Notoxaea, possesses sixsegmented maxillary palpi. Also the males of Protoxaea, like those of Mesoxaea and Notoxaea, have

the gonostyli partially differentiated from the gonocoxites while they are not evident in Oxaea (Figure
24 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

12). Thus in these and other characters Protoxaea

appears to be more closely related to Mesoxaea and

Notoxaea than it is to Oxaea. However, since the

mandibles of Notoxaea are uniquely bidentate apically, we conclude that although Protoxaea, Mesoxaea, and Notoxaea were derived from the same

ancestry, Protoxaea is more closely related to Mesoxaea. It is noteworthy that while Protoxaea and

Mesoxaea occupy partly sympatric ranges, these

genera occur in North America, well to the north

of both Oxaea and Notoxaea.

As now understood, Protoxaea is composed of

three species, two of which (P. australis and P.

micheneri) occur in Mexico below the elevation of

Mexico City and well to the south of the disjunct

from the third (P. gloriosa), which is widely distributed across northern Mexico and adjacent

southwestern United States (Map 1).

Key to the Species of the Genus Protoxaea

1. Males 2

Females 4

2(1). Sixth metasomal tergum entirely pale pubescent, chiefly yellow or yellowish white, hairs

at sides frequently white; discal pubescence of metasomal terga III-V mostly or

entirely pale 3

Sixth metasomal tergum mostly or entirely dark pubescent, reddish to brownish black,

occasionally a few hairs at extreme sides white; discal pubescence of metasomal terga

III-V mostly or entirely dark 1. P. australis

3 (2). Eyes less strongly convergent above, separated on vertex by more than distance between

rims of antennal sockets; middle and hind femora bright reddish yellow apical ly

2. P. gloriosa

Eyes more strongly convergent above, separated on vertex by less than distance between

rims of antennal sockets; middle and hind femora black 3. P. micheneri

4(1). Fifth and sixth metasomal terga chiefly pale pubescent, mostly yellow or yellowish

white 5

Fifth and sixth metasomal terga dark pubescent, reddish brown to brownish black, at

most with a few white hairs on extreme sides 1. P. australis

5(4). Vertex evenly rounded, not modified as below; pronotal collar emarginate medially

2. P. gloriosa

Vertex carinate posteriorly, medially emarginate, and as seen from in front bilobate in

outline; pronotal collar elevated medially 3. P. micheneri

1. Protoxaea australis, new species

FIGURES 4, 17-24; MAP 1

Protoxaea sp. nr. gloriosa.—Linsley and Michener, 1962:388

[Mexico: Yautepec, Morelos: males hovering about Cassia].—Michener, 1962:169 [Morelos].

GEOGRAPHIC RANGE (Map 1).—South-central and

southwestern Mexico (Guerrero, Morelos, and

Puebla).

MALE.—Length 15-18 mm. Length of forewing,

including tegula, 14-17 mm. Head and body black;

metasomal terga faintly metallic blue green in

bright light, most evident on impunctate, apical

margins; antennae brownish black above, reddish

brown on some flagellar segments below; mandibles

black, reddened apically; clypeus black, narrowly

reddened on medioapical margin; wings dissimilar

in coloration, forewings hyaline basally, rather contrastingly infuscated with brown black apically and

weakly violaceous in bright light, hind wings nearly

hyaline throughout, faintly stained with brownish

black infuscation narrowly along costal margin;

wing veins dark reddish brown, nearly black apically; tegulae reddish brown to dark mahogany,

darkening basally to black; legs dark reddish brown

to nearly black basally, tarsi usually somewhat redder; tibial spurs pale reddish brown; claws deep

mahogany red, banded medially with black.

Vestiture of head chiefly ochraceous to nearly

white on hind margins and face below antennal

sockets; pubescence on face above antennal sockets

fulvo-ochraceous to fulvous, paling to ochraceous

or white on vertex behind; thorax thickly clothed

dorsally with fulvous pubescence, paling posteriorly

to nearly white on propodeum and to fulvoochraceous on lower sides and ventrally; legs chiefly
NUMBER 220 25

• gloriosa

• micheneri

O australis

MAP 1.—Known occurrence of the species of Protoxaea.

pale pubescent, mostly whitish or cinereous basally,

darkening on tibiae and tarsi to reddish brown;

pubescence of first metasomal tergum consisting of

long, erect white plumose hairs, thickest laterally

and on ventrally exposed surfaces; metasomal terga

II-V without apical bands of pale pubescence,

mostly clothed with short, posteriorly directed,

brownish black pubescence, longest and thickest at

sides, and basal discal surface of tergum II with

erect whitish or cinereous pubescence; metasomal

tergum VI with pubescence much longer than that

on preceding terga, longest at sides and entirely

brownish black; metasomal tergum VII with pubescence disposed laterally as posteriorly projected

tufts of plumose hairs, similar in size and color to

that on tergum VI; metasomal sterna thickly

clothed with rather long plumose hairs, usually

mostly or entirely white on sterna I—III and reddish

brown on succeeding sterna, which in some specimens may display variable amounts of pale pubescence apically and usually laterally at extreme

sides.

Eyes moderately converging above, separated on

vertex by considerably more than minimum distance between hind ocelli and also by more than

minimum distance between rims of antennal sockets, but by much less than distance across hind

ocelli; clypeus densely and confluently punctate,
26

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

17

18 19

FIGURES 17-20.—Protoxaea australis Hurd and Linsley, holotype male (Mexcala, Guerrero,

Mexico): 17, seventh metasomal tergum (left half ventral); 18, 19, seventh and eighth metasomal

sterna (left halves dorsal); 20, genitalia (left half ventral).

punctures on sides much smaller than those on

median surface; supraclypeal area about as finely

and densely punctate as on upper sides of clypeus;

frontal carina short, narrowly triangular in outline, about as long as maximum diameter of median ocellus, weakly excavated basally; middle tibiae and basitarsi of equal length; punctation of

metasomal terga I and II dense, crowded, about

equal size, that on tergum II more crowded, usually

separated by less than their diameters, and extending well posteriorly onto marginal depression;

punctation on terga III-V of similar size to that on

tergum II, but becoming sparser on each tergum

and with larger impunctate areas medially, al-
NUMBER 220 27

though punctures extend well onto marginal depressions; structure of seventh metasomal tergum

(Figures 17, 21), seventh and eighth metasomal

sterna (Figures 18, 19, 22, 23), and genitalia (figures 20, 24) as illustrated.

FEMALE.—Length 16-18 mm. Length of forewing,

including tegula, 15-16 mm. Head and body black,

somewhat paler on metasomal sterna; metasomal

terga faintly metallic blue green in bright light,

most evident on impunctate, apical margins; antennae brownish black above, somewhat paler below and scape reddened at apex; mandibles black,

reddened apically and often medially; clypeus

black; wings dissimilar in coloration, forewings

hyaline basally, apically lightly infuscated with

brownish black, hind wings nearly entirely hyaline,

scarcely infuscated apically; wing veins dark reddish brown; tegulae reddish brown, darker basally;

legs dark reddish brown, blacker on coxae, and

trochanters, redder on tarsi; tibial spurs pale reddish brown, nearly testaceous; claws dark mahogany

red, blackish medially.

Vestiture of head moderately long and pale

ochraceous or white on clypeus, becoming fulvoochraceous about antennal sockets and above posterior ocelli; hind margins of head thickly clothed

with long pubescence, mostly white or pale ochraceous, becoming intermixed dorsally near vertex

with fulvo-ochraceous hairs; thorax thickly clothed

with fulvous pubescence dorsally, paling to ochraceous or white on scutellum, metanotum propodeum

and on lower sides and ventrally; legs chiefly white

pubescent, but external surfaces of fore and middle

tibiae and tarsi reddish brown pubescent; scopal

hairs of hind legs entirely white or nearly so; discal

surface of first metasomal tergum with a few long,

erect, whitish or ochraceous hairs basally, and apically with some similarly colored, but shorter and

posteriorly projecting hairs situated laterally and

forming a medially incomplete transverse band;

similar medially interrupted bands of pale pubescence present on discs of metasomal terga II-V,

which at extreme sides consist of longer hairs;

otherwise discal surfaces clothed only basally at

sides with very short, inconspicuous brownish hairs;

scopal hairs of terga I and II white or nearly so;

metasomal tergum V apically transversed by a thick

fringe of long, posteriorly directed, plumose hairs

characteristically reddish brown, but at extreme

sides sometimes with a few intermixed white hairs;

metasomal tergum VI densely clothed on either

side of pygidial plate with moderately long reddish

brown hairs; scopal hairs of metasomal sterna I—III

white, other surfaces of metasomal sterna II-VI

thickly clothed with rather short to long, posteriorly directed, reddish brown hairs; extreme sides

of metasomal sterna II-V with long tufts of white

plumose hairs projecting posteriorly from under

overlapping tergal surfaces.

Eyes scarcely diverging below (48:52); clypeus

densely, coarsely, and confluently punctate on sides,

punctures on median surface larger and nearly confluent, not separated by their diameters; supraclypeal area with punctures about as coarse and

crowded as on upper surface of clypeus; frontal

carina narrowly triangular in outline, deeply excavated, longer than maximum diameter of median ocellus; metasomal tergum I with discal surface basal to marginal depression nearly impunctate

medially, but with large punctures laterally, separated by less than their diameters at extreme sides;

discal surfaces of metasomal terga II-IV sparsely

impunctate medially, more crowded laterally, but

mostly separated by their diameters or more; fifth

metasomal tergum rather densely punctured

throughout, medially punctures separated by about

or less than their diameters.

TYPES.—Holotype male and allotype female were

collected at Mexcala, Guerrero, Mexico, on 29

June 1951 by P. D. Hurd, Jr., and are deposited in

the collections of the California Academy of

Sciences, San Francisco. Additional specimens

(paratypes) were collected at the type locality on

the same date by H. E. Evans (1 $ , 1 9) and P. D.

Hurd, Jr. (I $) and are, respectively, on deposit in

the collections of the California Insect Survey,

Essig Museum of Entomology, University of California, Berkeley, and the Snow Entomological Museum, University of Kansas. Other specimens examined (paratypes) include: 1 $ , 3.7 miles north

of the Rio Balsas (Highway 95), Guerrero, Mexico,

5 August 1963 (G. W. Byers, and party, KU);

9 ,$ , 2 2 , 4 miles southwest of Yautepec (3800 feet),

Morelos, Mexico, 2 July 1961, from flowers of an

unidentified species of Cassia (C. D. Michener,

KU); 1 $ , 10 miles west of Cuautla, Morelos, Mexico, 22 July 1962 (H. E. Milliron, CNC); and 2 $ ,

Petlalcingo, Puebla, Mexico, 23 July 1962 (H. E.

Milliron, CNC).

DISCUSSION.—Like P. micheneri, this species ap-
28

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

22

24

23

FIGURES 21-24.—Protoxaea australis Hurd and Linsley, paratype male (Petlalcingo, Puebla,

Mexico): 21, seventh metasomal tergum (left half ventral); 22, 23, seventh and eighth metasomal

sterna (left halves dorsal); 24, genitalia (left half ventral).

pears to inhabit the upland areas of south-central

Mexico below the lower limits of the pine forest. As

will be noted in the accompanying illustrations, the

paratype male from Petlalcingo, Puebla, Mexico,

has a somewhat broader eighth metasomal sternum

with a more slender apex (Figure 23) than that

exhibited by the holotype male from Mexcala,

Guerrero, Mexico (Figure 19). In addition, the

pubescence on the seventh metasomal tergum of

the Puebla specimen (Figure 21) is noticeably

longer than that of the holotype male (Figure 17).

In spite of these differences and some others evident in these structures, as well as the genitalia

(Figures 20, 24), we interpret them as individual

variations and regard the specimens as conspecific.

To judge by the overall similarities in structural

features, P. australis is more nearly related to P.

micheneri than it is to P. gloriosa. Doubtless these

species were derived from a common ancestry since

they share so many characters.
NUMBER 220 29

2. Protoxaea gloriosa (Fox)

FIGURES 2, 3, 25-36; MAP 1; PLATES 1-3

Megacilissa gloriosa Fox, 1893:421:422 $ [New Mexico: Las

Cruces].

Oxaea gloriosa.—Cockerell, 1898:70 [New Mexico: Mesilla

Valley].—Friese 1898:85, 86, $ $ [New Mexico: Las Cruces,

Mesilla].

Protoxaea gloriosa.—Cockerell and Porter, 1899:411.—Crawford, 1915:578, $ $ [Texas: Sabinal, females at flowers of

Salvia sp. and S. pitcheri; males, Barstow],—Cockerell,

1926:80, $ [Arizona: Baboquivari Mountains]; 1934b: 153

[Texas: Bexar County].—Rozen, 1951:142, 148, pi. 2: fig.

5.—Linsley and Michener, 1962:386-388, figs., $ $ [Southern Arizona and western New Mexico; females taking pollen from flowers of Kallstroemia californica, Solarium

elaeagntfolium, and S. rostratum; also present at flowers of

Baccharis glutinosa, Larrea tridentata, Asclepias subverticillata, Hoffmanseggia jamesii, Melilotus alba, and Medicago

sativa; Mexico: Chihuahua, at flowers of Tamarix gallica].—Linsley and Cazier, 1963:8, figs. 3-5 [Arizona: 1 mile

east of Douglas, females taking pollen from the flowers

of Solarium and nectar from flowers of Asclepias, Verbesina,

and Larrea; males taking nectar from the flowers of

Verbesina, Asclepias, and Lepidium].—Rozen, 1964:223-230,

12 figs.—Linsley and Cazier, 1972:1-25 [Arizona: Douglas,

Portal; females taking pollen from the flowers of Cassia

bauhinioides, Kallstroemia grandiflora, Solarium elaeagnifolium, S. rostratum, and Calliandra schottii].—Cazier

and Linsley, 1974:1:20, $ $ [New Mexico: 2 miles north

of Rodeo, females taking pollen and nectar from the

flowers of Kallstroemia grandiflora; males taking nectar

from the same plant].—Hurd and Linsley, 1975:27, $ $

[Southern Arizona, New Mexico and Sonora, females taking pollen from the flowers of Larrea tridentata, Solanum

elaeagnifolium, S. rostratum, Cassia bauhinioides, Calliandra schottii, and Kallstroemia grandiflora].

Protoxaea gloriosa pallida Cockerell, 1934b: 153, 9 $ [Texas:

Bexar County].—Linsley, 1951:1086, $ $ [Texas]. [New

synonymy.]

Oxaea tristis Gribodo, 1894:278-280, $ [Mexico: Durango].—

Friese, 1898:83, $ [Mexico: Durango]. [New synonymy.]

LOCATION OF TYPES.—Protoxaea gloriosa, Academy of Natural Sciences, Philadelphia; P. gloriosa

pallida, American Museum of Natural History,

New York; P. tristis, Museo Civico di Storia Naturale, "Giacomo Doria," Genoa, Italy.

GEOGRAPHIC RANGE (Map 1).—Southwestern

United States (Arizona, New Mexico, and Texas)

and adjacent northern Mexico (Baja California,

Chihuahua, Coahuila, Durango, Nuevo Leon, Sinaloa, Sonora, and Tamaulipas).

MALE.—Length 14-18. Length of forewing, including tegula, 13-16 mm. Head and body black,

metasomal terga feebly metallic blue green, most

evident on impunctate apical margins; antennae

dark reddish brown above, redder below on apical

flagellar segments; mandibles mahogany red, nearly

black apically; clypeus black, narrowly edged with

red apically; wings dissimilar in coloration, forewings hyaline basally, heavily infuscated apically

with brownish black, which reflect violaceous tints

in bright light, hind wing hyaline scarcely infuscated apically; wing veins reddish brown, darker

apically; tegulae testaceous; legs black on coxae,

trochanters, femora, and much of tibiae; tibiae

apically and tarsal segments yellow or reddish

yellow; tibial spurs testaceous; claws dark mahogany red, blackened medially.

Vestiture of head chiefly ochraceous, paler and

whiter on hind margins, turning to fulvoochraceous on face above ocelli and on vertex

(older specimens chiefly or entirely whitish or

white); thorax thickly clothed dorsally with fulvoochraceous to fulvous pubescence (whiter in older

specimens), paling to ochraceous or white on

thorax behind and to ochraceous on sides and ventrally (whiter in older specimens); legs entirely

pale pubescent, mostly ochraceous basally turning

to fulvous on tarsi; pubescence of first metasomal

tergum consisting of very long, erect, ochraceous to

whitish plumose hairs, thickest laterally and on

ventrally exposed surfaces; pubescence on second

metasomal tergum similar in color to that on first

tergum, but shorter, chiefly erect, and pale ochraceous or whitish on basal surface of disc medially,

much shorter, mostly decumbent, directed posteriorly and situated laterally, forming a subapical

transverse band of ochraceous pubescence; metasomal terga III-V clothed with short, ochraceous to

yellowish pubescence (whitish or white in older

specimens), chiefly disposed laterally, but forming

subapical band, which at sides consists of much

longer and frequently whitish hairs; pubescence on

metasomal tergum VI much longer than that on

preceding terga, chiefly yellow, ochraceous, or yellowish white (white in older specimens), hairs at

sides becoming white or nearly so; metasomal tergum VII with pubescence disposed laterally as

posteriorly projecting tufts of plumose hairs, similar in size aud color to that on tergum VI; metasomal sterna thickly clothed with moderately long

ochraceous or yellowish hairs (whitish in older

specimens), those situated apically on sterna longer
30

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

and disposed mainly as transverse bands of

pubescence.

Eyes moderately converging above, separated on

vertex by much more than minimum distance between hind ocelli and also by more than minimum

distance between rims of antennal sockets, but less

than distance across hind ocelli; clypeus densely

and confluently punctured on sides, punctures on

median surface larger and coarser, and well separated; supraclypeal area densely, finely, and confluently punctured, punctures much smaller and

more crowded than those on adjacent median surface of clypeus; frontal carina short, narrowly triangular in outline, nearly as long as maximum

diameter of median ocellus, deeply excavated

throughout its length; middle tibiae longer than

basitarsi (65:60); punctation on metasomal terga

I-VI dense, crowded, separated by about puncture

width or less, extending posteriorly well onto marginal depressions with punctures on tergum I

larger than those on succeeding terga; structure of

seventh metasomal tergum (Figures 25, 29, 33),

seventh and eighth metasomal sterna (Figures 26,

27, 30, 31, 34, 35), and genitalia (Figures 28, 32,

36) as illustrated.

FEMALE.—Length 17-22 mm. Length of forewing,

including tegula, 15-17 mm. Head and body black;

metasomal terga scarcely to feebly metallic blue

green with some purplish tints evident in bright

light and most evident on impunctate apical margins; antennae dark reddish brown to brownish

black, somewhat paler below, and apex of scape

narrowly reddened; mandibles chiefly mahogany

red to nearly red basally, darkening to black apically; clypeus black, occasionally reddened somewhat apically; wings dissimilar in coloration, forewings hyaline tinged with yellowish on basal third,

heavily infuscated with brownish black on apical

two-thirds and rather strongly violaceous in bright

light, hind wings mainly hyaline except for some

slight infuscation apically; wing veins reddish

brown to black; tegulae testaceous; legs mostly

reddish yellow, but coxae and trochanters black and

femora stained or marked in part with black; tibial

spurs reddish yellow; claws dark mahogany red,

medially marked with black.

Vestiture of head chiefly moderately long and

mostly ochraceous on face, sometimes paler about

antennal sockets and above ocelli near vertex: hind

margins of head thickly clothed with ochraceous

pubescence, paling to white on propodeum; legs

entirely pale pubescent, chiefly ochraceous, somewhat paler on hind femora; scopal hairs of hind

legs mostly ochraceous, usually intermixed with

some paler, nearly white hairs; discal surface of

first metasomal tergum clothed with long, erect

pale (mostly white) plumose hairs; discal surfaces

of metasomal terga II-IV before glabrous margins,

clothed with very short, erect, ochraceous hairs,

thickest and longest at sides, which are replaced on

extreme sides and ventrally exposed surfaces of

terga I and II by scopae consisting of long, densely

plumose, mainly whitish hairs and ventroapically

on terga III and IV with long tufts of ochraceous

plumose hairs; metasomal tergum V apically traversed by a thick fringe of long, posteriorly directed, plumose hairs, characteristically and uniformly ochraceous or yellowish, but occasionally

turning to white on extreme sides; metasomal tergum VI densely clothed on either side of pygidial

plate with moderately long, reddish yellow hairs,

rarely turning to white at extreme sides; scopal

hairs of metasomal sterna I—III ochraceous, other

surfaces of metasomal sterna II-VI thickly clothed

with rather short to long, posteriorly directed,

ochraceous pubescence; extreme sides of metasomal

sterna II-V with long tufts of ochraceous to whitish

plumose hairs projecting posteriorly from under

overlapping tergal surfaces.

Eyes scarcely diverging below (55:57), clypeus

closely and finely punctate on sides, punctures on

median surface larger and somewhat less crowded;

supraclypeal area densely and confluently punctured except at sides and above; frontal carina narrowly triangular in outline, short, longer than

diameter of median ocellus, deeply excavated

throughout length; discal surfaces before marginal

depressions of metasomal terga I-V densely and

finely punctured, those on tergum I separated by

mostly twice or more their diameters and those on

succeeding terga by less or about their diameters.

SPECIMENS EXAMINED.—United States. Arizona: Cochise

County: Apache, 2 $, VIII-8-72, on flowers of Kallstroemia

sp. (G. and K. Eickwort, CU); $, VIII-10-72 (G. and K.

Eickwort, CU). Apache, 4 mi SE, 3$, 9, VIII-18-64, on

flowers of Hoffmanseggia densiflora (C. D. Michener, KU).

Apache, 2 mi S, $, VIII-3-74 (S. Szerlip, CIS). Apache, 3-5

mi SW, $, VIII-&-59, on flowers of Baccharis glutinosa (H.

E. Evans, CU). Benson, £ , VII-27-54, on flowers of Prosopis

sp. (G. D. Butler, UA). Bernardino, $, VII-24-62, on flowers

of Verbesina encelioides, 1717 MST (M. A. Cazier, CIS).
NUMBER 220 31

28

26

FIGURES 25-28.—Protoxaea gloriosa (Fox), male (Douglas, Arizona): 25, seventh metasomal tergum

(left half ventral); 26, 27, seventh and eighth metasomal sterna (left halves dorsal); 28, genitalia

(left half ventral).

Bisbee, 6 mi SW, $, VII-26-73, on flowers of Menodora

scabra, 0915-0930 MST (T. J. Zavortink, TJZ); 47$, VIII5-75, most females collecting pollen from flowers of Cassia

xvislizenii, 0545-1030 MST (T. J. Zavortink, TJZ). Bowie,

5 $, VII-19-56, on flowers of Medicago sativa (A. Telford,

UA). Bowie, 8 mi S, $, VIII-22-67 (E. I. Schlinger, UCR).

Bowie Junction, $, VIII-6-54 (F. G. Werner, UA). Carr

Canyon, Huachuca Mountains, $, VIII-6-24 (E. P. Van

Duzee, CAS). Cascabel, 3 mi NW, $, VII-17-60, on flowers

of Prosopis glandulosa var. torreyanum, (J. E. Gillaspy, CAS).

Cave Creek Ranch, 2 $, $, VIII-2 to 4-69 (E. G. Linsley, CIS).

Double Adobe, 17 $, 26$, VIII-9-75, on flowers of Kallstroemia grandiflora, 0830-0900 MST (E. G. and J. M. Linsley, CIS). Cochise Stronghold, Dragoon Mountains, $, VIII5-55 (G. D. Butler, UA). Douglas, $, VII-22-26 (W. W.

Jones, PHT); $, VIII-25-26 (W. W. Jones, PHT); $, VIII15-40 (W. W. Jones, CIS); $, VIII-26-64 (J. G. Rozen,

AMNH); $, VIII-9-75, on flowers of Acacia angustissima,

1300-1330'MST (P. D. Hurd, A. E. and M. M. Michelbacher,

CIS); 2 $, VIII-10-75, on flowers of Acacia angustissima,
32

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

29

30

32

31

FIGURES 29-32.—Protoxaea gloriosa (Fox), male (Las Delicias, Chihuahua, Mexico): 29, seventh

metasomal tergum (left half ventral); 30, 31, seventh and eighth metasomal sterna (left halves

dorsal); 32, genitalia (left half ventral).

08S0-O900 MST (P. D. Hurd, A. E. and M. M. Michelbacher,

CIS). Douglas, 3 mi E, 2 9, VIII-5-75, collecting pollen from

flowers of Cassia wislizenii, 0520-0700 MST (E. G. and J. M.

Linsley, CIS). Douglas, 3.6-3.8 mi E, 46 $, VIII-3-75, most

females collecting pollen from flowers of Cassia wislizenii,

0530-1000 MST (T. J. Zavordnk, TJS); 47 9, VIII-4-75,

same data, except 0515-0530 (T. J. Zavortink, TJS). Douglas,

4.0-4.1 mi E, $, VIII-2-75, on flowers of Menodora scabra,

0730-0745 (T. J. Zavortink, TJZ). Douglas (about A3. mi

SW of road to Rucker Canyon Recreation Area), 2 $, VIII19-69, on flowers of Kallstroemia grandiflora, 0800-0830 MST

(T. J. Zavortink, TJZ). Douglas, 8 mi NE, VIII-11-40,

freshly dead by roadside (P. H. Timberlake, PHT). Douglas,

16 mi NE, 2$, VIII-23-62 (J. G. Rozen, M. Statham and

S. J. Hessel, AMNH). Douglas, 1 mi E, $, VII-16-62 (M. A.

Cazier, CIS); $, VH-20-62, sleeping in flowers of Conyza
NUMBER 220 33

coulteri, 1845 MST (M. A. Cazier, CIS); 2$, VII-21-62, on

flowers of Asclepias subverticillata, 1315 MST (M. A. Cazier,

CIS); $, VII-23-62, on flowers of Verbesina encelioides

(M. A. Cazier, CIS); 4$, VII-26-62 (M. A. Cazier, CIS); 2$,

VII-26-62, on flowers of Asclepias subverticillata (M. A.

Cazier, CIS); 12 4 , 2$ , VII-26-62, on flowers of Asclepias

subverticillata, 0935-0940 MST (M. A. Cazier, CIS); $, VII26-62, on flowers of Lepidium lasiocarpum, 0937 MST (M. A.

Cazier, CIS); $, 9, VII-26-62, in copulo, 0947 MST (M. A.

Cazier, CIS); 11 $, $, VII-26-62, on flowers of Verbesina

encelioides, 0942-1025 MST (M. A. Cazier, CIS); $, VII-26-

62, on flowers of Lepidium lasiocarpum, 1031 MST (M. A.

Cazier, CIS); 5$, 9, VII-26-62, on flowers of Asclepias

subverticillata, 1030-1037, MST (M. A. Cazier, CIS); 2$,

VII-26-62, 1040-1125 MST (M. A. Cazier, CIS); $, VII-26-

62, on flowers of Verbesina encelioides, 1040-1125 MST

(M. A. Cazier, CIS); 6$, VII-26-62, on flowers of Asclepias

subverticillata, 1125-1135 MST (M. A. Cazier, CIS); 7$, VII26-62, on flowers of Verbesina encelioides, 1130-1138 (M. A.

Cazier, CIS); 8$, VII-26-62, on flowers of Lepidium lasiocarpum, 1130-1138 MST (M. A. Cazier, CIS); 129, VII-26-62,

1140 MST (M. A. Cazier, CIS); $, VII-26-62, on flowers of

Lepidium lasiocarpum, 1540 MST (M. A. Cazier, CIS); \2$,

VII-26-62, on flowers of Conyza coulteri, 1542 (M. A. Cazier,

CIS); 19,$, VII-26-62, sleeping duster on Conyza coulteri,

1547 MST (M. A. Cazier, CIS); 2$, VII-26-62, sleeping on

Conyza coulteri, 1550 MST (M. A. Cazier, CIS); 4 $, VII26-62, on flowers of Conyza coulteri, 1623 MST (M. A. Cazier,

CIS); $, VII-26-62, on flowers of Ascelpias subverticillata,

1648 MST (M. A. Cazier, CIS); 2 $, VII-26-62, on flowers of

Lepidium lasiocarpum, 1655-1700 MST (M. A. Cazier, CIS);

$, VII-26-62, 1755 MST (M. A. Cazier, CIS); $. VII-26-62,

sleeping in flower of Argemone platyceras, 1755 MST (M. A.

Cazier, CIS); $, VII-27-62, hovering about one foot above

ground in open area and knocking scarabs down as they flew

over and also a Ptiloglossa was knocked off flower of Larrea

(No holes in ground), 1215-1230 MST (M. A. Cazier, CIS);

3$, VII-27-62, on flowers of Lepidium lasiocarpum, 1249

MST (M. A. Cazier, CIS); 63 , VII-27-62, sleeping cluster on

Conyza coulteri, 1645 MST (M. A. Cazier, CIS); 10 $, VII27-62, on flowers of Lepidium lasiocarpum, 1700-1730 MST

(M. A. Cazier, CIS); S3 , VII-27-62, inside seed pod of

Datura quercifolia, 1900 MST (M. A. Cazier, CIS); 5 $, VII27-62, inside seed pod of Datura quercifolia, 1910 MST (M.

A. Cazier, CIS); 73 $, VII-27-62, sleeping cluster on seed pod

of Datura quercifolia (started clustering at 1730 MST), 1915

MST (M. A. Cazier, CIS); 39 £, VII-27-62, sleeping cluster

on seed pod of Datura quercifolia (started clustering

at 1735 MST), 1920 MST (M. A. Cazier, CIS); 9, VIII^t-62,

on flowers of Solanum elaeagnifolium 0705-0715 (M. A.

Cazier, CIS); 9, VIII-4-62, on flowers of Solanum rostratum,

0952 MST (M. A. Cazier, CIS); 3 9, VIII-7-62, on flowers of

Solanum rostratum, 0905-1117 MST (M. A. Cazier, CIS);

2 9 , VIII-8-62, on flowers of Solanum elaeagnifolium, 0622

and 0652 MST (M. A. Cazier, CIS); 9, VIII-9-62, on flowers

of Solanum elaeagnifolium, 0705-0715 MST (M. A. Cazier,

CIS); $, VIII-9-62, on flowers of Condalia sp., 0952 MST

(M. A. Cazier, CIS); $, VIII-10-62, on flowers of Condalia

sp. (M. A. Cazier, CIS) 9, VIII-11-62, on flowers of Solanum

elaeagnifolium, 0632-0645 MST (M. A. Cazier, CIS); $, VIII11-62, on flowers of Hoffmanseggia densiflora, 0710 MST

(M. A. Cazier, CIS), $, VIII-11-62, on flowers of Solanum

rostratum, 0714 MST (M. A. Cazier, CIS); 3,9 , VIII-15-62,

mating on flowers of Conyza coulteri, 0907 MST (M. A.

Cazier, CIS); 15 9, VIII-17-62, on flowers of Solanum

elaeagnifolium, 0600-0930 MST (M. A. Cazier and E. G.

Linsley, CIS); 59 , VIII-17-62, on flowers of Solanum rostratum, 0600-0900 MST (M. A. Cazier and E. G. Linsley,

CIS); 5 9, VIII-18-62, on flowers of Solanum elaeagnifolium,

0540-0740 MST (M. A. Cazier and E. G. Linsley, CIS); 229,

VIII-18-62, on flowers of Solanum rostratum, 0650-0900 MST

(M. A. Cazier and E. G. Linsley, CIS); $, VIII-19-62, on

flowers of Verbesina encelioides (M. A. Cazier, CIS); 39 ,

VIII-20-62, on flowers of Solanum rostratum (M. A. Cazier,

CIS); 89 , VIII-17-67, on flowers of Solanum rostratum,

0640-0940 MST (M. A. Cazier and E. G. Linsley, CIS). Douglas, 3.7 mi E, 4,400 ft, 9, IX-2-74, on flowers of Kallstroemia grandiflora 1130-1145 MST (T. J. Zavortink, TJZ).

Douglas, 8 mi E, 9, VIII-8-58 (P. D. Hurd, CIS). Douglas

Station, 9, VII-27 to 31-70 (W. F. Brewton, GEB). Dry

Canyon Sands Ranch, SE and Whetstone Mountains, 3£,

VIII-10-52 (H. B. Leech and J. W. Green, CAS); 2$, same

data, but at flowers of Aloysia wrightii (H. B. Leech and

J. W. Green, CAS). Elfrida, 2$, VII-27-54, on flowers of

Medicago sativa (G. D. Butler, UA). Hereford, 9, 1954

(W. M. Mann, USNM). Huachuca Mountains, 9, VIII-28-03

(Oslar CU). Mescal, $, VIII-28-27 (R. H. Beamer, KU); 9,

VIII-30-53, on flowers of Kallstroemia grandiflora (G. D.

Butler, UA). Pearce, 2$, VII-27-54, on flowers of Kallstroemia grandiflora (G. D. Butler, UA). Pearce, 6 mi N, 9,

VIII-6-55, on flowers of Lepidium sp. (G. D. Butler and

Z. Noon, UA). Pearce (east), $, 2 9, VIII-7-63 (J. G. and B. L.

Rozen, AMNH). Pearce, 6 mi E, 16 $, VII-28-55 (G. Butler

and F. Werner, UA); 10 $, VIII-28-55, on flowers of

Mimosa sp. (G. Butler and F. Werner, UA). Pearce, 5 mi S,

7^, VII-28-55 (G. Butler and F. Werner, UA); $$, VII-28-

55, on flowers of Acacia angustissima (G. Butler and F.

Werner, UA). Portal, $, VII-24-54 (R. M. Bohart. UCD);

$, I-I3-61 (A. E. Michelbacher, CIS). Portal, 1 mi N, 22$,

279, VIII-2 to 5-72, on flowers of Larrea tridentata, 0630-

1529 MST (E. G. and J. M. Linsley, CIS). Portal, 2 mi N,

8 9 , VIII-16-70, on flowers of Cassia bauhinioides, 0520-

0640 MST (E. G. Linsley, CIS); 38 9, VIII-17-70, on flowers

of Cassia bauhinioides, 0640-0820 MST (E. G. Linsley. CIS);

469, VIII-19-70, on flowers of Cassia bauhinoides, 0600-

0740 (E. G. Linsley, CIS); 349, VIII-21-70, on flowers of

Cassia bauhinioides, 0540-0730 MST (E. G. Linsley, CIS).

Portal, 25 mi N, $, VII-25-61, 0923 MST (E. G. Linsley,

CIS); 3,9 , VII-26-61, in copulo, 0845 MST (E. G. Linsley,

CIS); 3,9 , VII-27-61 (E. G. Linsley, CIS); 9£ , 9, VII-27-

61, on flowers of Baccharis sp. (E. G. Linsley, CIS); 7 $, 9.

VII-31-61 (E. G. Linsley, CIS). Portal. 13 mi NE, $, VIII8-59 (E. G. Linsley, CIS). Portal, 2 mi NE, 2$, VIII-1-59.

on flowers of Baccharis sp. (M. Statham, AMNH); 9, VIII4-59, oq flowers of Baccharis glutinosa (M. A. Cazier,

AMNH). Portal, 2.5 mi NE, $, 9, VIII-9-59 (M. A. Cazier,
34 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

M. Statham, AMNH). Portal, 29 , VIII-21-70 (J. G. and

K. C. Rozen, AMNH); 3$ , VIII-21-70, on flowers of Cassia

sp., 0510-0630 MST (J. G. and K. C. Rozen, AMNH). Portal,

2.6 mi. E., 53 , VIII-15-72, on flowers of Prosopis glandulosa

var. torreyana, 0945-1045 MST (T. J. Zavortink TJZ). Portal,

4 mi E, $, VIII-28-71 (J. Doyen, CIS). Portal, 0.7 mi W,

2 3 , VIII-3-72, on flowers of Baileya multiradiata, 0850-

0950 MST (T. J. Zavortink, TJZ); $, VII-27-73, on flowers

of Menodora scabra, 0900-0915 MST (T. J. Zavortink, TJZ).

Portal, 1.7 mi NW, 23 , VIII-8-72, on flowers of Baccharis

sp., 0745-0830 MST (T. J. Zavortink, TJZ). Portal, 2 mi

NW, 29 , VIII-11-75, on flowers of Menodora scabra, 0900-

0930 MST (T. J. Zavortink, TJZ). Portal, 2.9 mi NW, 5,300

ft., 8, VIII-6-72, on flowers of Aloysia wrightii, 1530-1630

MST (T. J. Zavortink, TJZ). Portal, 12.9 mi NW, 5,300 ft,

3 , VIII-6-72, on flowers of Mentzelia sp., 1530-1630 MST

(T. J. Zavortink, TJZ). Portal, 6.2 mi NNW, 3 3,49 . VIII3-72, on flowers of Larrea tridentata, 0510-0710 MST (T. J.

Zavortink, TJZ). San Bernardino Ranch 3750 ft, $ (F. T.

Snow, CAS). San Simon (21 mi N Portal), 14 $, VIII-20-69,

ex bird nest in Yucca data, 1250-1255 MST (T. J. Zavortink,

TJZ). Sierra Vista, 4 mi SE, 3$, 29 , VIII-11-75, on flowers

of Kallstrocmia grandiflora, 1130-1200 MST (P. D. Hurd,

A. E. and M. M. Michelbacher, CIS). Skeleton Canyon, 6 mi

SE Apache, 9, IX-4-58, on flowers of Solarium sp. (P. D.

Hurd, CIS). Southwestern Research Station, 2 3,9 , VII-24-

56, on flowers of Melilotus alba (C. and M. Cazier, AMNH);

VII-24-56, on flowers of Melilotus alba (E. Ordway, CIS); 3 $ ,

VII-26-56, on flowers of Meliotus alba (E. Ordway, PHT);

2 3 , VII-27-56 (E. Ordway, PHT); 3 , VIII-28-56, on flowers

of Melilotus alba (E. Ordway, AMNH); 3 , VII-30-56, on

flowers of Melilotus alba (E. Ordway, PHT); 4 3 , 2 9, VII22-66 (R. G. Beard, CU); 5 3,39 , VIII-3-66 (C. R. Kovacic

and D. R. Miller,, UCD); 3,9 , VII-20-69 (L. L. Pechuman,

CU). Tombstone, 9, VIII-13-40, on flowers of Larrea tridentata (P. H. Timberlake, PHT). Willcox, 17 mi NW, 23 ,

9. VII-19-69 (A. E. and M. M. Michelbacher, CIS). Willcox,

21 mi SSW, 4500 ft, 39 . IX-1-74, on flowers of Cassia leptocarpa, 0945-1000 MST (T. J. Zavortink, TJZ). Willcox Dry

Lake. 4200 ft, 9, IX-2-74, on flowers of Dalea scoparia,

0730-0745 MST (T. J. Zavortink, TJZ). Gila County: Calva

Junction, 33 , 109. VIII-15-58, on flowers of Gaillardia

pulchella (E. G. Linsley, CIS). Globe, 23 , 389, VIII-29-64,

on flowers of Kallstroemia grandiflora (C. D. Michener, KU).

Graham County: Graham Mountains, 3500-4500 ft, 3 , 9,

VIII-18-54 (R. M. Bohart, UCD). Safford. 29 , VII-29-54,

on flowers of Medicago saliva (G. D. Butler, UA). Safford,

17 mi S, 49 , VIII-17-72, on flowers of Prosopis glandulosa

var. torreyana, 0840-0940 MST and 10 9, on flowers of

Kallstroemia grandiflora 0840-0940 MST (T. J. Zavortink,

TJZ). Snowline Ranch, Mt. Graham, 3 , VIII-21-52 (E. J.

Taylor. UCDV Maricopa County: Aguila, 2 mi W, 3 , on

flowers of Kallstroemia grandiflora (P. H. Timberlake, PHT).

Pima County: Arivaca, 7.5 mi W, 3 , VII-31-52, on flowers

of Verbes'ma encelioides (H. B. Leech and J. W. Green CAS).

Ajo, 30 mi SE. 3 . VH-30-66 (D. R. Miller, UCD). Baboquivari Mountains, 9, VIII-15-24 (O. C. Poling, PHT). Continental, 73 , 5 9, VIII-2 to 4-59, on flowers of Kallstroemia

grandiflora (K. V. Krombein, USNM); $, VII-31-66 (D. R.

Miller, UCD); 9, VIII-1-72, on flowers of Cassia sp., 0715-

0815 MST (T. J. Zavortink, TJZ). Continental, 5.2 mi N,

3 9 , VIII-11-69, on flowers of Kallstroemia grandiflora, 1200-

1230 MST (T. J. Zavortink, TJZ). Continental, 6 mi ESE,

2 9 , VIII-24-64, on flowers of Kallstroemia sp. (J. C. Bequaert and F. Werner, UA). Continental, 6.7 mi SE, 83 ,

6 9 , VIII-12-69, on flowers of Kallstroemia grandiflora 0945-

1045, MST (T. J. Zavortink, TJZ). Continental, 7 mi SE,

9, VIII-17-66 (J. G. and B. L. Rozen, AMNH). Continental,

8 mi E, 43 , VIII-13-55, sleeping on Thurberia sp. (R. A.

Flash, PHT); 3 , VIH-14-55, on flowers of Kallstroemia

grandiflora (R. A. Flash, PHT). Kits Peak (near), Baboquivari Mountains, 3600 ft, $, VIII-7 to 9-16 (AMNH).

Marana, 3 , 3 9, VIII-14-53, on flowers of Kallstroemia

grandiflora (F. E. Todd, UA); 9, VIH-25-53, on flowers of

Kallstroemia grandiflora (F. E. Todd, CIS). Marana, 3 mi

W, 3 3 , 29 , IX-13-62, on flowers of Kallstroemia grandiflora

(J. C. Bequaert, UA). Sabino Canyon, Santa Catalina Mountains, 3 , IX-1-53, on flowers of Acacia angustissima (G. D.

Butler, UA). Sahuarita, 0.8 mi E, 11 9, VIII-11-69, on flowers

of Larrea tridentata, 0700-0930 MST (T. J. Zavortink TJZ).

Sahuarita, 1.9 mi SW, 33 , VIII-11-69, on flowers of Larrea

tridentata, 1000-1130 MST (T. J. Zavortink, TJZ). Sahuarita,

3 mi E, 3 , VII-31-63 (V. L. Vesterby, UCD). Santa Rita IBP

Site, 9, (UA). Santa Rita Ranger Res., $, 149, VIII-29-64,

on flowers of Kallstroemia grandiflora (J. C. Bequaert, UA).

Santa Rita Mountains, 9 , VII-26-25 (USNM); 3 , VIII-18-

68 (R. M. Bohart, UCD). Three Points, 5 mi W, 3 , VIII18-63 (F. Werner UA). Tucson, 29 , VIII-14 to 17-16

(AMNH); $.9, VIII-16-33 (J. Russell, KU); 3 , IX-4-39

(R. H. Crandall, UA); 3 , 9, VIII-28-51, on flowers of

Kallstroemia grandiflora (C. D. MacNeill, CIS); $, VIII10-63 (V. L. Vesterby, UCD); 16 3 , 13 9, VIII-5-11-73, on

flowers of Larrea tridentata, 0600-1259 MST (E. G. and

J. M. Linsley, CIS). Tuscon, 6 mi SW, 3 , VIII-26-68, on

flowers of Kallstroemia sp. (R. W. Thorp, UCD). Tucson,

10 mi N, 3,29 , VIII-8-54, on flowers of Prosopis glandulosa var. torreyana (F. G. Werner, UA). Tucson Mountains,

3 , IV-VIII-55 (G. D. Butler, UA). Pinal County: Eloy, 4

mi S, 3 , VII-14-53 (T. R. Haig, UCD). Eloy, 11 mi SE, 9.

VII-16-53 (T. R. Haig, UCD). Oracle, 14 mi E, $, VH-27-

24 (J. O. Martin, CAS). Picacho Pass, 23 , 49 , VIII-27-54,

on flowers of Kallstroemia grandiflora (P. H. Timberlake,

PHT). Santa Cruz County: Amado, 3 , VII-23-57 (G. D.

Butler, UA). Amado, 4 mi S, 13 3 , 129, VHI-10-75, on

flowers of Kallstroemia grandiflora, 0900-0930 MST (E. G.

and J. M. Linsley, CIS). Madera Canyon, 43 , VII-31-66

(D. R. Miller, UCD); 9, VII-31-66 (C. R. Kovacic, UCD).

Nogales, 3 , VII-21-03 (Oslar, CAS). Nogales, 30 mi N, 49,

VII-31-72, on flowers of Kallstroemia grandiflora, 0730-0930

MST (T. J. Zavortink, TJZ). Nogales International Airport,

269, VIII-7-75, collecting pollen from flowers of Desmanthus

cooleyi, 0730-0830 MST (P. D. Hurd, A. E. and M. M.

Michelbacher, CIS); 29 , same data, except 1100 MST (CIS).

Patagonia, 3 , VIII-24-53 (G. D. Butler and F. G. Werner,

UA); 129, VIII-29-54, on flowers of Solanum rostratum

(P. H. Timberlake, PHT). Patagonia, 9 mi NE, 3 .

 vm '
NUMBER 220 35

27-55 (H. A. Scullen, GEB). Patagonia 5 mi W, #,3$ ,

VIII-25-55, on flowers of Acacia angustissima (UA). Pena

Blanca Lake, $, VIII-17-74, on flowers of Acacia angustissima, 0930-0945 MST (T. J. Zavortink, TJZ); 6 $, 8 $, VIII18-74, on flowers of Kallstroemia grandiflora, 0845-1130 MST

(T. J. Zavortink, TJZ); 4$, 3$ , VIII-20-74, on flowers of

Kallstroemia grandiflora, 0830-1145, MST (T. J. Zavortink,

TJZ). Pena Blanca Lake, 5.3 mi W, 4700 ft, VII-29-72, on

flowers of Acacia angustissima, 1115-1215 MST (T. J. Zavortink, TJZ); 4$ , VIII-10-74, on flowers of Cassia leptocarpa,

0700-0730 MST (E. G. and J. M. Linsley, CIS). Ruby, $,

VIII-15-61 (J. C. Bequaert, UA); 8$, VIII-16-61 (J. C.

Bequaert, UA). Sonoita, 7.6 mi E, $, VIII-15-55. on

flowers of Croton corymbulosus (P. H. Timberlake, PHT).

New Mexico. Bernalillo County: Albuquerque, 2 $, VII23-50 (R. H. Beamer, KU); $, VII-27-50 (P. P. Cook, KU).

Isleta, $, VH-17-52, on flowers of Melilotus alba (R. H.

and L. D. Beamer, W. E. LaBerge, and C. Liang, KU).

Chaves County: Roswell, 5 mi S, $, VII-22-52 (R. H. and

L. D. Beamer, W. E. La Berge, and C. Liang, KU). Roswell,

10 mi W, $, $, VII-22-52, on flowers of Hoffmanseggia

jamesii (R. H. and L. D. Beamer, W. E. LaBerge, and C.

Liang, KU); 9, VII-22-52, on flowers of Larrea tridentata

(R. H. and L. D. Beamer, W. E. La Berge, and C. Liang,

KU). Dona Ana County: Hatch, 8$, VII-16-52, on flowers

of Medicago sativa (R. H. and L. D. Beamer, W. E. La Berge,

and C. Liang, KU). Las Cruces, $, VIII-21 (T. Townsend,

ANSP); 13 $, VIII-1927, on flowers of Gossypium sp. (R. C.

Stockdale, USNM); 2$, VII-15-52, on flowers of Lygodesmia

juncea (R. H. and L. D. Beamer, W. E. La Berge, and C.

Liang, KU); $, VIII-4-66 (R. L. Brumley, UCD); $, 9 .

VIII-15-67 (R. M. Bohart, UCD). Las Cruces, 5 mi NE, $,

VIII^l-66 (R. L. Brumley, UCE). Las Cruces, 5 mi E, 2J ,

VII-15-52 (R. H. & L. D. Beamer, W. E. La Berge, and

C. Liang, KU). Mesilla, 5 mi S, $, IX-4-57, on flowers of

Sphaeralcea sp. (P. Gerhardt, PHT). Mesilla Valley $,

(AMNH). Eddy County: Carlsbad, $$, VII-11-42 (E. C.

Van Dyke, CAS). Carlsbad, 5 mi N, $, VIII-14-50 (J. W.

Mac Swain, CIS); $, IX-21-56 (J. W. MacSwain, CIS). Hwy.

137, 7 mi W Jet Hwy 285 at Rocky Arroyo, $, VII-30-72

(S. L. Szerlip, CIS). Hidalgo County: Las Animas, 17.5 mi

NNE, 3 $, 29 , VIII-22-72, on flowers of Larrea tridentata,

0545-1045 MST (R. J. Zavortink, TJZ). Animas 1.4 mi E,

2 9 , VIII-11-72, on flowers of Hoffmanseggia densiflora 0630-

0730 MST (T. J. Zavortink, TJZ); 119, VIII-11-72, on

flowers of Solanum elaeagnifolium, 0630-0750 MST (T. J.

Zavortink, TJZ). Animas, 5.5 mi E, 49 , VIII-11-72, on

flowers of Solanum elaeagnifolium, 0800-0830 MST (T. J.

Zavortink, TJZ). Animas, 10 mi SW, 10 9, VIII-11-72, on

flowers of Larrea tridentata, 0530-0600 MST (T. J. Zavortink,

TJZ). Antelope Pass, 123 9, VIII-8 to 10-72, on flowers of

Larrea tridentata, 0630-1359 MST (E. G. Linsley, CIS).

Cienega Lake, 13 mi N Rodeo, 14,$, 39 , VIII-3-71, on

flowers of Tamarix sp. (J. T. Doyen, CIS). Granite Pass,

24,5, 22 9, VIII-19 to 20-72, on flowers of Larrea tridentata,

0700-1329 MST (E. G. and L. M. Linsley, M. A. Cazier,

and T. J. Zavortink, CIS). Lordsburg, 3 mi NE, 2 $, VIII21-71 (R. F. Denno and E. E. Grissell, UCD). Lordsburg,

5 mi NE, $, VIII-21-71 (R. F. Denno and E. E. Grissell,

UCD). Road Forks, 4miS,2j , 9, VIII-20-67 (J. G. Rozen

and G. Kreuger, AMNH). Rodeo, 9, VIII-1-58 (P. M.

Marsh, UCD); 2$, VIII-3-59 (H. E. Evans, CU); 3^, 39 ,

VIII-5-59 (H. E. Evans, CU); 9, VII-23-60 (M. A. Cazier,

AMNH); 5 $, VIII-12-69 (J. G. and K. C. Rozen, AMNH);

2$, VIII-6-72 (G. Eickwort, CU). Rodeo, 1 mi N, 9, VIII18-70 (J. G. Rozen, AMNH); 9, VIII-23-70 (J. G. Rozen,

AMNH); 29 , VIH-25-70 (J. G. Rozen, AMNH); 9, VIII30-70 (J. G. Rozen, AMNH); Rodeo, 2 mi N, $, VII-29-59,

on flowers of Asclepias subverticillata (E. G. Linsley, CIS);

115 3 , 1469, VIII-14-74, on flowers of Kallstroemia grandiflora, females 0800-1130 MST, males 0800-1400 MST (E. G.

and J. M. Linsley, CIS). Rodeo, 8 mi N, $, VIII-13-69

(J. G. and K. C. Rozen, AMNH). Rodeo, 11 mi N, 9, VIII19-68 (J. G. Rozen and M. Favreau, AMNH). Rodeo, 16-20

mi N, $, VIII-18-71 (J. G. Rozen and M. Favreau, AMNH).

Rodeo, 20 mi N, 3^, 49 , VIII-3-66 (R. L. Brumley and

D. R. Miller, UCD). Rodeo, 1 mi S, 18 $, VIII-3-59, sleeping in cluster, 1550 MST (E. G. Linsley, CIS). Rodeo, 1.5

mi S, 7 $ , 4 9, VII-24-59, on flowers of Asclepias subverticillata (E. G. Linsley, CIS); 3£ , 9, VIII-6-59, on flowers of

Asclepias subverticillata (E. G. Linsley, CIS). Rodeo, 1 mi

SW, $, VIII-4-72, on flowers of Psilostrophe tagetina, 1200-

1230 MST (T. J. Zavortink, TJZ). Rodeo, 1 mi W, $, VIII10-60 (J. E. Gillaspy, CAS). Luna County: Deming, 9, IX3-27 (J. C. Bradley, CU). Deming, 5 mi E, $, 9, VIII-13-63,

on flowers of Asclepias sp (J. G. Rozen, AMNH). Deming,

7 mi E, 5$, VIII-4-66 (D. R. Miller, UCD). Rockhound

State Park, 9, VIII-1-72 (G. C. Eickwort, CU). Otero

County: High Rolls, 6 mi W, $, VI-11-70 (W Apperson,

GEB). Organ, 3 mi E, 3<J, VII-26-57 (C. W. O'Brien, UA).

Sierra County: Elephant Butte, 5 mi W, $, 11 9, VIII-8-

73, on flowers of Larrea tridentata, 0730-0859 MST (E. G.

and J. M. Linsley, CIS). Hot Springs, $, VII-22-50, on

flowers of Solidago occidentalis (W. J. Arnold, KU). Socorro

County: Bernardo, $, VH-24-70 (W. Apperson, GEB); 9,

VIII-1-70 (W. Apperson, GEB). San Antonio, 20 mi SSW,

20$, 59 , VIII-18-62 (J. G. and B. L. Rozen, AMNH).

Texas: Pecos River, 2$, 9 (J. C. Bequaert, MCZ). Bexar

County: $, VI-18-30 (H. B. Parks, USNM); $, VI-19-30

(H. B. Parks, USNM); $, VI-21-30 (H. B. Parks, TAM); $,

VI-24-30 (H. B. Parks, TAM); 2$, VI-26-30 (H. B. Parks,

TAM); $, IX-25-30 (H. B. Parks, TAM); $ VI-4-31 (H. B.

Parks, TAM); 39, VII-1-31 (H. B. Parks, MCZ, TAM, U

Colo); 3,79 , VII-10-31 (H. B. Parks, TAM); $, VII-20-31

(H. B. Parks, TAM); $, IX-10-31 (H. B. Parks, TAM); 29 .

IX-15-31 (H. B. Parks, TAM); 9, VI-26-32 (H. B. Parks,

TAM); 3,9 , IX-21-32 (H. B. Parks, TAM); 9, IX-16-36

(TAM). Ft. Sam Houston, 9, VII-19-52 (M. S. Wasbauer,

AMNH). San Antonio, 9, (USNM). Brewster County: 29 .

VI-27-55, on flowers of Larrea tridentata (A. H. Alex. TAM).

Alpine, <J, VI-3-42 (E. C. Van Dyke, CAS); $, VII-1-42

(E. C. Van Dyke, CAS). Big Bend National Park, $, VII16-50 (R. F. Smith, AMNH). Chisos Mountains, Big Bend

National Park, 3$, 29 , on flowers of Aloysia lycioides (J. C.

Bequaert, MCZ). Cov. Springs (6 mi W) Big Bend National

Park, $, VII-16-50 (R. F. Smith, AMNH). Lajitas, 3 mi W.
36

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

36

34

FIGURES 33-36.—Protoxaea gloriosa (Fox), male (8 mi S Canutillo, Durango, Mexico): 33, seventh

metasomal tergum (left half ventral); 34, 35, seventh and eighth metasomal sterna (left halves

dorsal); 36, genitalia (left half ventral).

$, X-2-60, on flowers of Tamarix sp. (J. E. Gillaspy, CAS).

Santa Elena Canyon, Big Bend National Park, $, VIII-25-

67 (R. C. Gardner and C. R. Kovadc, UCD); £ , VIII-26-67

(R. C. Gardner and C. R. Kovacic, UCD). Terlingua, $,

VH-19-50 (R. F. Smith, AMNH). Dimmit County: 5$,

VII-7-34, "swarmed on mesquite" (S. E. Jones, TAM); 9,

X-2-34 (TAM). Caurina, 9, VI-10-48 (M. A. Cazier,

AMNH). El Paso County: El Paso, $, VII-14-42 (E. C. Van

Dyke, CAS); 2$ , 49 , VIII-15-51 (H. E. Evans, KU). Frio

County: 9, VII-19-01 (G. P. Engelhardt, AMNH). Frio

State Park, 5 $, VI-28-48 (H. E. Evans, CU). Harris County:

Balesville, $, VI-28-10 (USNM). Hidalgo County: 2$, V-1S33 (J. W. Monk, TAM). Jeff Davis County: Ft. Davis, 21

mi W, 2$, VII-22- to 25-56 (E. G. Matthews, CU). Jim

Wells County: Alice, &, VI-22-30 (J. O. Martin, CAS).

Kleberg County: Kingsville, 2$, (C. T. Reed, MCZ); $, X20-72 (J. M. Bogart, GEB). Medina County: 33 , VI-26-55,

on flowers of Helianthus annum (A. H. Alex, TAM). Pecos
NUMBER 220 37

County: Ft. Stockton, $, VII-5- (J. C. Bequaert, MCZ).

Presidio County: Chinati Mountains, 25 mi NW Presidio,

4400 ft, $, VI-25-68 (J. E. Hafernik, TAM). Plata, 3 $, 3 $,

VIII-24-69 (Board and J. E. Hafernik, TAM). Presidio, 3 mi

N, $, IX-1-66 (C. L. Cole, TAM). Presidio, 58 mi E, 2$ ,

IX-8-71 (R. F. Denno and E. E. Grissell, UCD). Shatter,

4$, VH-18-65 (J. C. Schaffner, TAM); $, VI-3-68 (J. E.

Hafernik, TAM); 2$ , VIII-21-69 (Board and J. E. Hafernik,

TAM). Reeves County: Pecos, 16 3 , VII-1938 (H. Sellers

USNM). Uvalde County: Garner State Park, $, VI-29-56

(H. E. Evans and E. G. Matthews, CU). Sabinal, $, VI-10-

10, on flowers of Salvia sp. (F. C. Pratt, USNM); $, VI13-10, on flowers of Salvia pitcheri (F. C. Pratt, USNM).

Uvalde, $, VI-15-30 (J. O. Martin, CAS); $, VIII-6-33

(TAM). Val Verde County: Del Rio, $, VII-7-38 (J. Russell,

KU); $, $, VII-8-38 (L. W. Hepner, KU). Devil's River

near Del Rio, 9, VI-16-32 (J. O. Martin, CAS). Ward

County: Barstow, $, VIII-11-05 (A. W. Morrill, USNM);

3 $ , VI-28-72, on flowers of Medicago sativa (GEB).

Mexico. Baja California; Canipole, 9, X-2--11 (E. S. Ross

and G. E. Bohart, CAS).

Chihuahua: Casas Grandes, $, VII-24-71 (D. W. Davis,

GEB). Chihuahua, 2$ , VIII-12-51 (H. E. Evans, KU). Chihuahua, 16 mi SE, $, VIII-11-47 (W. J. Gertsch, David

Rockefeller Mexican Expedition, AMNH). Chihuahua, 20

mi SE, $, VII-13-64, on flowers of Parkimonia aculeata

(J. A. Chemsak, CIS). Chihuahua, 3$, 9, VII-9-54 (J. W.

MacSwain, CIS). 6 $, same data (E. I. Schlinger, CIS). Ciudad

Camargo, 4 mi N, $, VII-29-67 (K. C. Gardner, C. R.

Kovadc and K. Lorenzen, UCD). Ciudad Camargo, 42 mi

SW, 4900 ft, 2$, VII-15-47 (C. D. Michener, David Rockefeller Mexican Expedition, AMNH). Hildalgo del Parral,

2$, 9, VIII-10-67 (R. C. Gardner, C. R. Kovacic, and K.

Lorenzen, UCD); $, VIII-15-67 (R. C. Gardner, C. R.

Kovacic, and K. Lorenzen, UCD). Jimenez, 10 mi N, 5 $,

9 $ , IX-21-70, on flowers of Solarium sp. (G. E. and R. M.

Bohart, GEB, UCD). Jimenez, 18 mi W, $, VIII-10-51, on

flowers of Baccharis sp. (P. D. Hurd, CIS). Las Delidas, 14 $,

9, VII-1S-47, one male at flowers of Cevallia sinuata (C. D.

Michener, David Rockefeller Mexican Expedition, AMNH);

3 ^, 29 , VII-11-51 (H. E. Evans, KU); 2$, 69 , same date,

on flowers of Casuarina sp. (P. D. Hurd, CIS). Moctezuma,

22^, 39 . VII-4-54 (E. I. Schlinger, CIS). Meoqui, 6 mi NE,

15$, 49 , IX-2-50 (R. F. Smith, AMNH). Ricardo Flores

Magon, ^,9 , VIII-15-52 (J. D. Lattin, CIS). Rio Conchos,

Ciudad Camargo, 4^, 9, VII-17-52 (J. D. Lattin, CIS).

Coahuila: La Gloria (South of Mondova), 3300 ft, 9, VII24-47 (M. A. Cazier, David Rockefeller Mexican Expedition,

AMNH). San Juan de Sabinas, 1160 ft, 9, V-22-52 (P. S.

Chraphwy, KU). San Pedro, 5 mi W, A$, VII-20-55 (J.

Connors, CAS).

Durango: Canutillo, 8 mi S, &, VIII-9-51 (H. E. Evans,

KU); 3$, 9, VIII-9-51, on flowers of Guardiola tulocarpa

(P. D. Hurd, CIS, $, Genova). La Quebrada, 29 , VII-20-47

(C. D. Michener, David Rockefeller Mexican Expedition,

AMNH). San Juan del Rio, 5200 ft, 2$ , 9, VIII-30-47 (C. D.

Michener, David Rockefeller Mexican Expedition, AMNH.

KU). Yerbanis, Cuencame Dist. 6700 ft, $, VIII-19-42, on

flowers of Lippia ligustrina (C. D. Michener, David Rockefeller Mexican Expedition, AMNH).

Nuevo Leon: Valledllo, 4# , VI-2 to 5-51 (H. E. Evans,

KU); 13 $, same date (P. D. Hurd, CIS).

Sinaloa: Los Mochis, 2$, VIII-9-22 (C. T. Dodds, CIS,

KU).

Sonora: Agua Prieta, $, V-17-44 (W. W. Jones, CIS); $,

VIII-13-46 (W. W. Jones, PHT). Guaymas, 5£ , VI-12-62

(D. H. Janzen, CIS); 9, VIII-31-68 (R. W. Thorp, UCD).

Hermosillo, 3.5 mi S, 2$, VIII-3-73, on flowers of Larrea

tridentata, 0930-0959 MST (E. G. and J. M. Linsley, A. E.

and M. M. Michelbacher, and J. A. Chemsak, CIS). Hermosillo, 10 mi S, 5^, VII-18-54 (M. A. Cazier, W. J. Gertsch,

and Bradts, AMNH). Hermosillo, 20 mi N, 9, VIII-13-59

(W. L. Nutting and E. G. Werner, UA).

Tamaulipas: Padilla, $, 9, VI-15-53, on flowers of

Cevallia sinuata (University of Kansas Mexican Expedition,

KU).

DISCUSSION.—Of the three species of the genus

Protoxaea as now understood, P. gloriosa is the

most widely distributed (Map 1). It occurs disjunctly well to the north of its congeners and is the

only species of the family Oxaeidae known to inhabit the Mexican Plateau. However, in the northeastern part of its range (Texas and Nuevo Leon,

Mexico) and also in a portion of its northwestern

range (Arizona, Sonora, and Sinaloa, Mexico) it

has been found at a number of localities with several species of the genus Mesoxaea (viz., M. texana,

M. arizonica, and M. rufescens). It has also been

collected on one occasion in Baja California (Canipole) somewhat to the north of the Cape Region

where M. vagans is evidently endemic.

Protoxaea gloriosa was first described by Fox

(1894:421-422) from Las Cruces, New Mexico,

from a single female specimen. In the following

year, Gribodo (1894:278-280) described the same

species as Oxaea tristis, n. sp., on the basis of two

males collected in Durango, Mexico. Apart from

the independent views of these authors whose aesthetic impressions differed on the appearance of

this species, our initial studies revealed that only

one species of the family Oxaeidae is found on the

Mexican Plateau, namely P. gloriosa. With this in

mind we reexamined the original descriptions provided by Fox and Gribodo and concluded that these

authors had characterized the same species even

though the names they had used suggested otherwise (gloriosa versus tristis). To confirm the suspected synonymy, a male specimen of P. gloriosa

from Durango (8 miles south of Canutillo) was

compared for us with the type of the Gribodo spe-
38

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

cies by Dott. Delfa Guiglia who communicated

that the last visible metasomal sternum (VIII) is,

as we suspected, not deeply emarginate (cf. Figures 27, 31, and 35 with the same structure in the

genus Mesoxaea, e.g., Figures 43, 47, 51, etc.). Accordingly, we have placed Oxaea tristis Gribodo as

a synonym of P. gloriosa. A reexamination of the

specimens recorded by Cockerell (1906:314; 1934b:

153) as tristis from Arizona (Douglas and San

Bernardino Ranch) reveals that these represent

Mesoxaea rufescens, which is described as a new

species elsewhere in this study.

There is a great deal of individual variation in

the size of the body as well as in the coloration of

the vestiture of this species. Almost certainly much

of the variation in body size is directly attributable

to the amount of food available to the larva during

development. Presumably small-sized adults result

from larvae that have fed upon less than the average amounts of pollen and nectar stores, although

doubtless other factors are involved and should be

investigated experimentally. Older adult specimens

(e.g., wings markedly frayed, mandibles of females

worn apically) tend to possess paler and sometimes

largely whitish pubescence. This suggests that certain nesting colonies, such as those observed by

H. B. Parks in Bexar County, Texas (Cockerell

1934b: 153), likely differ in their relative ages. Perhaps this sort of variation prompted Cockerell

(1934b: 153) to characterize P. gloriosa pallida as a

subspecies. As he points out the observed differences among nesting colonies are interesting and

deserve further study. However, since we have been

unable to detect geographic, size or colorational

differences within the species, we have relegated P.

gloriosa pallida to synonymy.

There is, as would be expected, a certain degree

of individual variation in the structure of the male

genitalia as well as in the associated metasomal

terga and sterna. To indicate the extent and nature

of this variation, we have selected and illustrated

these structures in males from Douglas, Arizona

(Figures 25-28), Las Delicias, Chihuahua, Mexico

(Figures 29-32), and Canutillo, Durango, Mexico

(Figures 33-36).

Protoxaea gloriosa is the most well-known species

of the family and much of our current knowledge

about behavior and comparative biology of these

bees is based upon studies of this species.

3. Protoxaea micheneri, new species

FIGURES 37-40

Protoxaea sp.—Linsley and Michener, 1962:388.

MALE.—Length 17 mm. Length of forewing, including tegula, 15 mm. Head and body black;

metasomal terga without metallic luster; antennae

dark reddish brown above, somewhat paler on apical flagellar segments below; mandibles black,

stained above apically and near middle with dark

mahogany red; clypeus black; wings similar in

coloration, hyaline basally, lightly infuscated with

brownish black apically, more noticeably so on forewings; wing veins chiefly dark reddish brown, somewhat darker apically and along costal margin;

tegulae testaceous; legs chiefly brownish black, apical tarsal segments reddish brown; tibial spurs pale

reddish brown; claws deep mahogany red, banded

medially with black.

Vestiture of head chiefly ochraceous, paler, and

whiter on hind margins, turning to fulvoochraceous on face above ocelli and on vertex;

thorax thickly clothed dorsally with mostly fulvoochraceous pubescence, paling to nearly white on

thorax behind and to ochraceous on lower sides and

ventrally; legs entirely pale pubescent, chiefly fulvoochraceous, paling to nearly white especially on

hind trochanters and coxae; pubescence of first

metasomal tergum consisting of long, erect white

plumose hairs, thickest laterally and on ventrally

exposed surfaces; metasomal terga II-V clothed

(except on tergum II with long, erect whitish hairs

medially at base) with golden, posteriorly directed,

simple hairs, shortest basally, much longer and

denser over marginal depressions imparting the

appearance of apical bands, which on ventrally exposed sides pale to white; pubescence on metasomal tergum VI much longer than that on preceding terga, mostly pale yellow, turning to white

on sides; metasomal tergum VII with pubescence

disposed laterally as posteriorly projecting tufts of

plumose hairs, similar in size and color to that on

tergum VI; metasomal sterna thickly clothed with

rather long, white plumose hairs.

Eyes rather strongly converging above, separated

on vertex by more than minimum distance between hind ocelli, but by much less than minimum

distance between rims of antennal sockets; clypeus

densely and confluently punctate, punctures on
NUMBER 220 39

38

40

FIGURES 37-40.—Protoxaea micheneri Hurd and linsley, holotype male (3 mi NW Tequesquitengo, Morelos, Mexico): 37, seventh metasomal tergum (left half ventral); 38, 39, seventh and

eighth metasomal sterna (left halves dorsal); 40, genitalia (left half ventral).

sides much smaller than those on median surface;

supraclypeal area about as finely and densely punctate as on upper sides of clypeus; frontal carina

short, scarcely evident, about as long as maximum

diameter of median ocellus, scarcely excavated basally; middle tibiae shorter than basitarsi; punctation of metasomal terga I and II extending well

posteriorly onto marginal depressions, rather dense,

and fine laterally, larger and sparser medially, and

separated there by much more than their diameters; punctation of terga III-V similar in size to

that on tergum II but much sparser and with some

impunctate areas medially, although punctures extend well onto marginal depressions; structure of

seventh metasomal tergum (Figure 37), seventh

and eighth metasomal sterna (Figures 38, 39), and

genitalia (Figure 40) as illustrated.

FEMALE.—Length 16-19 mm. Length of forewing, including tegula, 13-14 mm. Head and body

black, metasomal terga scarcely with metallic blue

green reflections in bright light, observable only in

certain positions on impunctate areas; antennae

dark brownish black above, somewhat paler below,

and scape reddened at apex; mandibles black, reddened somewhat above and apically; clypeus black;

wings dissimilar in coloration, forewings hyaline
40 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

basally, apically lightly infuscated with brownish

black, hind wings nearly entirely hyaline, scarcely

infuscated apically; wing veins mostly reddish

brown, darker on costal and outer veins; tegulae

testaceous; legs dark reddish brown, blacker on

coxae, trochanters, and femora; tibial spurs pale

reddish brown; claws dark mahogany red, irregularly mottled with black.

Vestiture of head moderately long and ochraceous or white on clypeus, becoming fulvo-ochraceous

about antennal sockets and above posterior ocelli;

hind margins of head thickly clothed with long pubescence, mostly pale ochraceous to white, becoming intermixed dorsally near vertex with fulvoochraceous hairs; thorax thickly clothed with

bright fulvous, pubescence dorsally, becoming ochraceous or whitish posteriorly on propodeum and

on lower sides and ventrally; legs chiefly white pubescent, but external surfaces of fore and middle

tibiae ochraceous to fulvo-ochraceous pubescent;

scopal hairs of hind legs entirely white pubescent

or nearly so; discal surface of first metasomal tergum with a few long, erect whitish or ochraceous

hairs dorsally, otherwise nearly devoid of pubescence; discal surfaces of metasomal terga II-IV before glabrous margins, clothed with very short,

mostly decumbent, posteriorly directed whitish

hairs mostly or entirely situated laterally, which at

extreme sides and on ventrally exposed surfaces are

replaced on terga I and II by scopae consisting of

long, densely plumose, mainly whitish hairs and

ventroapically on terga III and IV with long tufts

of chiefly whitish plumose hairs; metasomal tergum V apically traversed by a thick fringe of long,

posteriorly directed, plumose hairs characteristically white at sides and ochraceous or yellowish on

medial tergal surface; metasomal tergum VI densely

clothed on either side of pygidial plate with moderately long, yellowish hairs, which at sides are replaced by small tufts of whitish pubescence; scopal

hairs of metasomal sterna I—III white, other surfaces of metasomal sterna II-VI thickly clothed

with rather short to long, posteriorly directed,

whitish or yellowish hairs; extreme sides of metasomal sterna II-V with long tufts of white plumose

hairs projecting posteriorly from under overlapping tergal surfaces.

Eyes scarcely diverging below (43:45); clypeus

densely, coarsely, and confluently punctate on sides,

punctures on median surface larger and separated

by about their diameters; supraclypeal area with

punctures smaller and more crowded than those on

adjacent median surface of clypeus; frontal carina

narrowly triangular in outline, deeply excavated,

longer than maximum diameter of median ocellus;

vertex carinate posteriorly, medially emarginate,

and as seen from in front bilobate in outline; pronotal collar tuberculately elevated medially; discal

surface basal to marginal depression of metasomal

tergum I nearly impunctate except for a few wellseparated punctures basally and at extreme sides;

discal surface of metasomal tergum II medially

nearly impunctate, with only a few widely scattered

punctures, which laterally become denser and

crowded; discal surfaces of terga III and IV with

punctation similar to that on tergum II, but impunctate medial area much reduced; fifth metasomal tergum densely and confluently punctured

throughout.

GEOGRAPHIC , RANGE (Map 1).—South-central

Mexico (Morelos and Puebla).

TYPES.—Holotype male, allotype female, and 3

paratypes (3 $) were collected at 3 miles northwest

of Tequesquitengo, Morelos, on 16 August 1962,

from the flowers of an unidentified species of Kallstroemia (Zygophyllaceae) by Radclyffe B. Roberts

and Norman B. Marston. These specimens are the

property of the University of Kansas and are deposited in the Snow Entomological Museum. One

additional paratype (9) was collected at 13 miles

southeast of Acatlan, Puebla, Mexico, on 10 July

1952, by E. E. Gilbert and C. Don MacNeill and is

located in the collection of the California Insect

Survey, Essig Museum of Entomology, University

of California, Berkeley.

DISCUSSION.—Protoxaea micheneri, named in

honor of Dr. C. D. Michener for his outstanding

contributions to the classification of bees, is one of

the most distinctive species not only of this genus,

but also of the family Oxaeidae. The peculiarly

carinate and medially emarginate vertex of the female is unique among the oxaeids and one can

only speculate on the possible adaptive significance

of this unusually modified head. It seems unlikely

that it represents an adaptive response stemming

from its intrafloral relationships and equally unlikely that it is specialized for some unusual nesting

requirements. Protoxaea micheneri is more closely

related to P. gloriosa than it is to P. australis, but

differs from both in many characteristics.
NUMBER 220

41

Mesoxaea, new genus

FIGURES 5, 6, 41-68; MAPS 2, 3

Oxaea.—Authors [in part; not Klug, 1807:26].

Protoxaea.—Authors [in part; not Cockerell and Porter, 1899:

410].

TYPE-SPECIES.—Oxaea nigerrima Friese, present

designation.

DIAGNOSIS.—Males without pale maculations on

clypeus, labrum, mandibles, or antennae; mandibles simple apically (Figures 5, 6); maxillary palpi

present, six-segmented; metasomal terga brownish

black or black, not partly or largely reddish; metasomal terga VI (males) and V (females) with long,

conspicuous tufts of white hairs at sides; eighth

metasomal sternum of male deeply emarginate medially on apical margin (Figures 43, 47, 51, 55, 59,

67); gonobase greatly reduced, ringlike, much

wider than long, not tapering basally and gonostyli evident, partially differentiated from gonocoxites (Figures 44, 48, 52, 56, 60, 64, 68).

DISCUSSION.—As presently understood the genus

Mesoxaea is composed of seven species which occupy much of the Mexican mainland to the northeast, south, and west of the Mexican Plateau.

Although none of the species is known to inhabit

MAP 2.—Known occurrence of Mesoxaea vagans (Fox), 'M. rufescens Hurd and Linsley,

Af. tachytiformis (Cameron), and M. texana (Friese).
42

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

the Mexican Plateau north of the Valley of Mexico

(Maps 2, 3), it is possible that one or more may

eventually be found in some of the deep barrancas

that originate in the Plateau and drain into the

Pacific. Two of the species occupy isolated ranges

with M. vagans apparently restricted to the Cape

Region of Baja California (Map 2) and M. texana

confined to western Louisiana, southern Texas, and

adjacent northeastern Mexico (Map 3). The other

species of the genus are centered primarily in

southwestern and western Mexico with two of the

species, Af. arizonica and M. rufescens, ranging

northward along the west coast of Mexico into extreme southern Arizona (Maps 2, 3). Mesoxaea

tachytiformis is known from the Valley of Mexico

and westward from several localities in Morelos,

Guerrero and Colima (Map 2). Mesoxaea clypeata

thus far has been collected only at a few localities

in the State of Nayarit, Mexico (Map 3), and

Mesoxaea nigerrima occupies the southernmost

range of the genus from Veracruz and Chiapas

northwestwardly to Colima and Jalisco (Map 3).

No species of the genus has been found in the Gulf

Coast region of Mexico between Vera Cruz and

Nuevo Leon. Almost surely one or more species of

Mesoxaea occur in this area and we interpret the

apparent absence of the genus to the lack of

collecting.

O arizonica

• clypeata

• nigerrima

MAP 3.—Known occurrence of Mesoxaea arizonica (Cockerell), M. clypeata Hurd and Linsley,

and M. nigerrima (Friese).
NUMBER 220 43

Key to the Species of the Genus Mesoxaea

1. Males 2

Females 8

2(1). Metasomal terga II-V black pubescent apically, without transverse bands of white

pubescence; eyes moderately to strongly convergent above, separated on vertex by

less than distance between rims of antennal sockets 3

Metasomal terga II-V pale pubescent apically, provided with transverse bands of white

pubescence; eyes less strongly convergent above, separated on vertex by more than

distance between rims of antennal sockets 1. M. tachytiformis

3(2). Clypeus nearly punctate throughout 4

Clypeus, except on sides, nearly impunctate, at most with a few widely scattered

punctures medially 7. M. clypeata

4(3). Forewings heavily infuscated with brownish black, at most with cells R and first Cu

hyaline or nearly so 5

Forewings hyaline or nearly so on basal half or more, only apical third or so infuscated

with brownish black 6

5(4). Forewings heavily and nearly uniformly brownish black, somewhat paler near base,

but not hyaline or nearly so; eyes moderately convergent above, separated on vertex

by about or more than distance between hind ocelli 5. M. rdgerritna

Forewings heavily infuscated with brownish black, but hyaline in cells R and first Cu;

eyes more strongly convergent above, separated on vertex by much less than distance

between hind ocelli 6. M. arixonka

6(4). Face above antennal sockets to hind margin of head at least in part and often chiefly

pale pubescent; hind margins of head mostly pale pubescent 7

Face above antennal sockets to hind margin of head entirely dark, pubescent; hind

margins of head, except pale pubescence on cheeks below, dark reddish-brown to

nearly black pubescent 4. M. texana

7(6). Eyes strongly convergent above, separated on vertex by less than distance between

hind ocelli 3. M. vagans

Eyes less strongly convergent above, separated on vertex by at least and usually much

more than distance between hind ocelli 2. M. rufescens

8(1). Dorsum of thorax pale pubescent, chiefly ochraceous, fulvo-ochraceous or fulvous, often

paler posteriorly 9

Thorax wholly dark pubescent, dark brown to brownish black pubescent 12

9(8). Second metasomal tergum punctate across much of discal surface before marginal

depression; clypeus flat or somewhat depressed on median surface, not furrowed

mediolongitudinally 10

Second metasomal tergum impunctate except at extreme sides; clypeus conspicuously

furrowed mediolongitudinally 1. M. tachytiformis

10(9). Hind margins of head and thorax below chiefly or entirely pale pubescent; second

metasomal tergum basal to marginal depression not densely punctate, with large

impunctate areas extending across discal surface 11

Hind margins of head and thorax below chiefly or entirely dark brown or brownish

black pubescent; second metasomal tergum basal to marginal depression densely

punctate, without large impunctate areas extending across discal surface ...4. M. texana

11(10). Punctures on second metasomal tergum just before marginal depression well scattered

and much of surface impunctate 2. M. rufescens

Punctures on second metasomal tergum just before marginal depression more densely

punctate, surface punctured more orless continuously across disc 3. M. vagans

12(8). Clypeus nearly punctate throughout 13

Clypeus, except on sides, nearly impunctate, at most with a few widely scattered

puctures medially 7. M. clypeata

13(12). Scopal hairs of hind femora chiefly or entirely white 5. M. mgerrhna

Scopal hairs of hind femora brownish black 6. M. arixonka
44 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

Mesoxaea is more closely related to Protoxaea

and Notoxaea than it is to Oxaea as indicated by a

number of structural features including the presence of six-segmented maxillary palpi and the

partly differentiated gonostyli of the males. Since

among other features, the mandibles of Notoxaea

are bidentate apically, it would appear that because of the simple mandibles Mesoxaea is more

nearly related to Protoxaea, although it may well

have been derived from the same ancestry that also

led to the formation of Notoxaea. Curiously these

three genera occupy allopatric ranges to the south

(Notoxaea) and north (Mesoxaea, Protoxaea) of

the genus Oxaea.

It should be noted that while the eyes in the

males of Mesoxaea converge above and sometimes

markedly so, there is individual variation in the

expression of this character so that some caution

must be exercised when using the degree of convergence for identification purposes. Thus, for example, individual variation in the minimum upper

interorbital distance ranges in 42 specimens of M.

vagans from 7 to 13 units with the mode at 8, while

in 53 specimens of M. arizonica this distance ranges

from 5 to 10 units with the mode at 7.

1. Mesoxaea tachytiformis (Cameron),

new combination

FIGURES 6, 41-44, MAP 2

Oxaea tachytiformis Cameron, 1901:315-316, $ [New Mexico:

Santa Fe Mountains].—Cockerell, 1920:177, $ [Mexico:

Distrito Federal; Guerrero: Venta Zopilote].

Protoxaea tachytiformis.—Graf, 1966:138.

Protoxaea impunctata Cockerell, 1920:178, $ [Mexico: Distrito Federal]. [New synonymy.]

LOCATION OF TYPES.—Mesoxaea tachytiformis,

British Museum of Natural History, London; M.

impunctata, U.S. National Museum of Natural

History, Washington, D.C.

GEOGRAPHIC RANGE (Map 2).—South-central and

southwestern Mexico (Colima, Distrito Federal,

Guerrero, and Morelos).

MALE.—Length 20-22 mm. Length of forewing,

including tegula, 16-18 mm. Head and body black;

metasomal terga faintly metallic blue green with

some purplish reflections in bright light, most evident on impunctate apical marginal depressions;

antennae dark brownish black above, reddened on

some flagellar segments below; mandibles deep

mahogany red with some black usually at base and

subapically; clypeus black or noticeably reddened

on medial surface; wings similar in coloration,

hyaline basally, infuscated with brownish black apically, mostly beyond closed cells; wing veins reddish

brown, somewhat darker apically; tegulae testaceous externally, darkening to brownish black basally; legs chiefly dark reddish brown or brownish

black, tarsi not appreciably darker than tibiae;

tibial spurs reddish brown; claws deep mahogany

red, usually marked irregularly with black.

Vestiture of head chiefly pale ochraceous, paler

and whiter on face below antennal sockets and on

hind margins of head; pubescence on face above

antennal sockets fulvo-ochraceous, darkening to

brownish black on upper part of face and vertex;

thorax thickly clothed with mostly ochraceous to

fulvo-ochraceous pubescence, paling to nearly

white on thorax behind, at sides, and below; legs

pale pubescent, chiefly ochraceous to white basally,

turning to bright fulvous on exterior surfaces of

tibiae and tarsi; pubescence of first metasomal tergum consisting of long, erect ochraceous to whitish

plumose hairs, thickest at sides and on ventrally

exposed surfaces; metasomal terga II-V with apical

bands of long, white plumose hairs, which extend

onto ventroapical margins, otherwise clothed with

short, posteriorly directed, brownish black pubescence except on basal discal area of tergum II,

which is densely clothed with erect, moderately

long white pubescence; metasomal tergum VI with

numerous long, posteriorly directed, white plumose

hairs arising just before impunctate apical margin,

which are longest and somewhat tuftlike laterally

and usually are intermixed on median tergal surface with brownish black hairs; metasomal tergum

VII with brownish black pubescence becoming

white at extreme sides, disposed laterally as posteriorly projecting tufts of plumose hairs; metasomal sterna thickly clothed with rather long plumose hairs, usually white on sterna I and II and

sometimes also on apical margins of sterna III and

IV, especially laterally, otherwise chiefly or entirely

reddish brown pubescent.

Eyes moderately convergent above, separated on

vertex by considerably more than minimum distance between hind ocelli and also by more than

minimum distance between rims of antennal sockets, but by much less than distance across hind

ocelli; clypeus depressed mediolongitudinally,
NUMBER 220 45

44

42

FIGURES 41-44.—Mesoxaea tachytiformis (Cameron), male (Venta de Zopilote, Guerrero, Mexico):

41, seventh raetasomal tergum (left half ventral); 42, 43, seventh and eighth metasomal sterna

(left halves dorsal); 44, genitalia (left half ventral).

coarsely and nearly confluently punctured on sides

and above, punctures becoming much less crowded

medially and nearly impunctate on somewhat

smoothed and reddened apical margin; supraclypeal area about as coarsely and densely punctate as

on upper sides of clypeus; frontal carina triangular

in outline, very short, much less than diameter of

median ocellus, deeply excavated basally; middle

tibiae and basitarsi of equal length; punctation on

discs of metasomal terga I and II extending well

posteriorly onto apical depressions, dense, rather

fine with punctures separated by mostly less than

their diameters, on terga III and IV similarly extending well onto apical depressions, punctures

somewhat larger and less crowded, especially medially, and with some impunctate or sparsely punctate areas extending across terga on and basal to

raised submarginal areas; fifth metasomal tergum
46

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

with discal punctures similar in size and disposition, but larger and less densely crowded medially;

structure of seventh metasomal tergum (Figure

41), seventh and eighth metasomal sterna (Figures

42, 43), and genitalia (Figure 44) as illustrated.

FEMALE.—Length 20-22 mm. Length of forewing,

including tegula, 16-18 mm. Head and body black,

reddened on metasomal terga and sterna; metasomal terga feebly metallic blue green with some

purplish tints in bright light and most noticeable

on impunctate surfaces; antennae dark brownish

black, reddened below and at apex of scape and

apex of pedicel; mandibles deep mahogany red,

black subapically; clypeus black, somewhat reddened apically; wings dissimilar in coloration, forewings rather uniformly infuscated throughout and

feebly violaceous in bright light, hind wings mainly

hyaline except for some blackish infuscation apically and posteriorly; wing veins dark reddish

brown to black; tegulae testaceous, somewhat darkened basally; legs dark reddish brown dorsally, extensively reddened below; tibial spurs nearly testaceous, apically darker; claws dark mahogany red

basally, black apically.

Vestiture of head chiefly moderately long and

ochraceous or white on face from clypeus nearly to

vertex where it becomes brownish black; hind margins of head thickly clothed with long pubescence,

mostly pale ochraceous to white, becoming brownish black on top of head; thorax thickly clothed

with bright fulvous pubescence dorsally, becoming

ochraceous or whitish posteriorly on metanotum,

propodeum, and on lower sides and ventrally; legs

chiefly white pubescent basally, darker pubescent,

usually reddish brown on exterior surfaces and on

tarsi; scopal hairs of hind legs entirely pale, mostly

white or nearly so; discal surfaces of metasomal

terga I-IV devoid of pubescence except for very

short, posteriorly directed brownish black hairs

situated laterally, which on terga I and II at extreme sides and on ventrally exposed surfaces are

replaced by scopae consisting of long, densely plumose, mainly whitish hairs and ventroapically on

terga III and IV with long tufts of chiefly whitish

plumose hairs; metasomal tergum V apically traversed by a thick fringe of long, posteriorly directed, plumose hairs characteristically white at sides

and brownish black on medial tergal surface; metasomal tergum VI densely clothed on either side of

pygidial plate with moderately long, brownish black

hairs, which at extreme sides are replaced by small

tufts of white pubescence; scopal hairs of metasomal sterna I—II white, other surfaces of metasomal

sterna II-V (except at extreme sides) and VI

thickly clothed with rather short to moderately

long, posteriorly directed, reddish brown to brownish black hairs; extreme sides of metasomal sterna

II-V with long tufts of white plumose hairs projecting from under overlapping tergal surfaces.

Eyes weakly diverging below (61:66); clypeus

mediolongitudinally depressed, closely, coarsely,

and rugosely punctate, except narrowly impunctate

mediolateral apical corners and roughened apical

margin, supraclypeal area about as coarsely and

rugosely punctured as adjacent clypeal areas; frontal carina narrowly triangular in outline, deeply

excavated, slightly longer than maximum diameter

of median ocellus; discal surfaces basal to marginal

depressions of metasomal terga I-IV impunctate

except for a few large punctures at sides; fifth

metasomal tergum densely and confluently

punctured.

SPECIMENS EXAMINED.—Mexico. Colima: Colima, 1 $, X (T.

Conradt, USNM).

Distrito Federal: Mexico, $, $ (J. R. Inda, USNM).

Morelos: Amacuzac, 6 mi S, $, X-8-63 (A. E. and M. M.

Michelbacher, CIS).

Guerrero: Venta de Zopilote, 2800 ft, $, X (H. H. Smith,

USNM).

DISCUSSION.—Although this species is known only

from a few specimens, we have no hesitancy in associating the sexes which were described as different

species. The male, which was named first by

Cameron (1901), is indeed reminiscent of a large

wasp of the genus Tachytes, but as Cockerell

(1920:177) has pointed out the type locality given

by Cameron (Santa Fe Mountains, New Mexico) is

erroneous. The holotype was examined for us by

Dr. Karl V. Krombein, who reports that there are

three labels affixed to the pin—a handwritten label:

"(17a547) Oxaea tachytiformis Cam. type New

Mexico," a small machine-printed label: "Cam.

Coll. 1902-105," and a third label: "B. M. TYPE

HYM. (machine printed), 17.2.547 (handwritten)."

Dr. G. Knerer subsequently looked further into

this matter for us and writes:

The records for this number (Cam. Coll. 1902-105) reveal

that the insects in this shipment came from "diverse localities." Then I had a look at the original description of Oxaea

tachytiformis and checked the other types of new species
NUMBER 220 47

described in the same article. These have labels similar to

Oxaea (Cam. Coll. 1902-105, and a handwritten label by

Cameron with the species name) but seem to have lost their

original collection labels. Cameron did none of the collecting himself and never saw any of the places where these

insects came. Cockerell was probably right when he thought

that the Santa Fe locality in New Mexico was erroneous.

The same strange locality records (or lack of) is seen for the

other new species described in the same paper: Paramasaris

fuscipennis, Zanysson aureobalteatus, Alyson flavomaculatus

and Osmia foxi. Based upon our current knowledge of the

distribution of these species it looks as if they came from a

more southern locality than the one given by Cameron.

None of these species has any syntypes in the British Museum of Natural History, but several O. tachytiformis were

collected later at Venta de Zopilote, Guerrero, Mexico.

In a report on the bees from San Miguel County,

New Mexico, Cockerell (1904:8,9) writes:

It seems advisable here to make some statement regarding

the species of Oxaea, Nomia, &c, which Mr. Cameron has

described (Trans. Am. Ent. Soc), purporting to come from

the region about Santa Fe, New Mexico. The character of

the species is Mexican, and I am quite confident that the

locality assigned is entirely wrong. I wrote Mr. Cameron

about it, and he kindly informed me that the material was

collected years ago by a person who was known to have

visited the Santa Fe region, but who might very well have

obtained the insects elsewhere. The collection included some

species of Bombus which might have come from near Santa

Fe.

The female of M. tachytiformis was described by

Cockerell (1920:178) as a new species (Protoxaea

impunctata) from the Federal District of Mexico.

The collections of the National Museum of Natural History, Smithsonian Institution, contain

both sexes from this locality. Consequently, there is

scarcely any doubt that these specimens represent

the sexes of the same species because this is the

only Mesoxaea known to occur in the Valley of

Mexico.

Mesoxaea tachytiformis is structurally a uniquely

distinctive species as is evident from the key to

species. To judge from the structure of male genitalia (Figure 44) it occupies an annectant position

between those species represented by M. arizonica,

M. clypeata, M. nigerrima, and M. texana on the

one hand and by M. rufescens and M. vagans on

the other. Conceivably M. tachytiformis was derived from the same ancestry that led to the formation of these two groups of species and is the sole

representative of its group.

2. Mesoxaea rufescens, new species

FIGURES 45-48; MAP 2

Oxaea tristis.—Authors [not Gribodo, 1894].—Cockerell,

1906a:72 [Arizona: San Bernardino Ranch]; 1906b:314

[Arizona: Douglas].

Protoxaea tristis.—Authors [not Gribodo, 1894].—Cockerell,

1934b: 153 [Arizona: San Bernardino Ranch and Douglas].—

Linsley, 1951:1086, $ [Mexico and Arizona].

GEOGRAPHIC RANGE (Map 2).—Southwestern

United States (Arizona) and adjacent northwestern

Mexico (Nayarit, Sinaloa, and Sonora).

MALE.—Length 15-19 mm. Length of forewing,

including tegula, 13-15 mm. Head and body chiefly

black; metasomal terga feebly metallic blue green,

most evident on apical marginal depressions in

bright light; antennae dark reddish brown to

brownish black, except apex of scape narrowly, and

pedicel and first flagellar segment nearly always red

or reddened; mandibles black, usually reddened on

basal half; clypeus black, normally narrowly reddened apically and sometimes all or much of median surface also reddened; wings dissimilar in

coloration, forewings hyaline on about basal twothirds, apical third darkly infuscated with brownish black, hind wings very nearly hyaline throughout, scarcely and only faintly infuscated apically;

wing veins mostly reddish brown, apical veins beyond closed cells darker; tegulae testaceous, darkened to brownish black basally; legs dark brownish

black basally, tarsi reddish brown; tibial spurs reddish brown; claws deep mahogany red, banded medially with black.

Vestiture of head chiefly ochraceous (nearly

white in older specimens), usually becoming pale

fulvo-ochraceous above antennal sockets and often

turning to brownish black on upper part of face

and vertex; hind margins of head chiefly or entirely ochraceous (paler in older specimens), sometimes intermixed dorsally with a few brownish

black hairs; thorax thickly clothed with fulvoochraceous pubescence throughout (paler in older

specimens), usually paling to ochraceous on sides

and ventrally; legs chiefly ochraceous to reddish

brown pubescent, palest basally, darker on tarsi and

nearly brownish on hind femora ventrally; pubescence of first metasomal tergum consisting of

long, whitish plumose hairs, thickest at sides and

ventrally, not darkened to black on raised submarginal areas laterally; metasomal terga II-TV
48

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

without apical bands of pale pubescence, mostly

clothed with short, posteriorly directed, brownish

black pubescence, longest and thickest at sides; tergum II tufted with white pubescence on ventrally

exposed sides and with shorter and less conspicuous, erect white hairs on discal surface medially at

base; metasomal terga III and IV tufted with long,

whitish pubescence on extreme ventrally exposed

sides; metasomal tergum V entirely brownish black

or black pubescent, shortest on punctate surface

basally, much longer and forming a transverse band

of simple hairs arising just before impunctate apical margin, projecting posteriorly and terminating

in thick tufts of plumose hairs laterally; metasomal

tergum VI with numerous long, posteriorly directed,

white plumose hairs arising just before impunctate

apical margin, longest and somewhat tuftlike laterally, usually intermixed on median tergal surface

with brownish black or black hairs; metasomal

tergum VII with pubescence chiefly white and disposed laterally as posteriorly projected tufts of

plumose hairs; metasomal sterna thickly clothed

with rather long, plumose hairs, usually white or

cinereous on sterna I—III, and on terga IV—VI

brownish pubescent basally, but apical hairs white

or nearly so.

Eyes moderately convergent above, separated on

vertex by about or more than minimum distance

between hind ocelli, but always by much less than

minimum distance between rims of antennal sockets; clypeus densely, finely and confluently punctured on sides, punctures on median surface larger,

coarser, and separated by shining interspaces on

lower half, except medial margin apically almost

impunctate, smooth, and reddened; supraclypeal

area, except above, densely and confluently punctured, without shining interspaces; frontal carina

somewhat triangular in outline, deeply excavated

basally, nearly as long as diameter of median ocellus; middle tibiae and basitarsi of equal length;

punctation on discs of metasomal terga I and II

dense, rather fine with punctures separated by

mostly less than their diameters, that on terga III

and IV similar, but larger and much less densely

disposed, especially medially, and with large impunctate or sparsely punctate areas extending

across terga and basal to submarginal depressions;

fifth metasomal tergum with discal punctures similar in size and disposition to those on tergum II,

but larger and less densely crowded especially medially; structure of seventh metasomal tergum (Figure 45), seventh and eighth metasomal sterna (Figures 46, 47), and genitalia (Figure 48) as

illustrated.

FEMALE.—Length 16-20 mm. Length of forewing,

including tegula, 14-15 mm. Head and body black,

not much paler on metasoma below; metasomal

terga feebly metallic blue green with some purplish

tints in bright light and most noticeably so on impunctate surfaces; antennae dark brownish black,

somewhat reddened at apex of scape, pedicel, and

variably below on usually apical flagellar segments;

mandibles black, reddened usually at base and occasionally apically; clypeus brownish black to black,

not reddened on medial surface; wings dissimilar in

coloration, forewings hyaline on about basal twothirds, apical third darkly infuscated with brownish black, hind wings nearly hyaline throughout,

scarcely and only faintly infuscated apically; wing

veins mostly reddish brown, apical veins beyond

closed cells darker; tegulae testaceous externally,

darkening to brownish black basally; legs mostly

brownish black to black, somewhat reddened on

tarsi and below; tibial spurs dark reddish brown;

claws deep mahogany red, banded medially with

black.

Vestiture of head chiefly long and white on face,

especially that about ocelli antennal sockets, clypeus at sides, becoming brownish black on upper

part of face and on vertex; hind margins of head

rather thickly and mostly clothed with long whitish

pubescence, sparsest and shortest on cheeks, intermixed with and sometimes replaced by dark brownish black pubescence dorsally near and on vertex;

thorax thickly clothed with fulvous pubescence

dorsally, paling to ochraceous on metanotum, propodeum, and on lower sides and ventrally; legs

chiefly white pubescent basally, darker pubescent,

usually reddish brown on exterior surfaces and on

tarsi; scopal hairs of hind legs entirely pale, mostly

white or nearly so; discal surfaces of metasomal

terga I-IV devoid of pubescence except for very

short, posteriorly directed, brownish black hairs

situated laterally, which at extreme sides and on

ventrally exposed surfaces are replaced on terga I

and II by scopae consisting of long, densely plumose, mainly whitish hairs and ventroapically on

terga III and IV with long tufts of chiefly whitish

plumose hairs; metasomal tergum V apically traversed by a thick fringe of long, posteriorly di-
NUMBER 220 49

46

FIGURES 45-48.—Mesoxaea rufescens Hurd and Linsley, male (5.3 mi W Pena Blanca Lake,

Santa Cruz County, Arizona): 45, seventh metasomal tergum (left half ventral); 46, 47, seventh

and eighth metasomal sterna (left halves dorsal); 48, genitalia (left half ventral).

rected, plumose hairs characteristically white at

sides and brownish black on median tergal surface;

metasomal tergum VI densely clothed on either

side of pygidial plate with moderately long, mostly

brownish black plumose hairs, sometimes at extreme sides with a few white hairs; scopal hairs of

metasomal sterna I—III white; other surfaces of

metasomal sterna II-V (except at extreme sides)

and VI thickly clothed with rather short to moderately long, posteriorly directed, reddish brown to

brownish black hairs; extreme sides of metasomal

sterna II-V with long tufts of white plumose hairs

projecting posteriorly from under overlapping tergal surfaces.

Eyes weakly diverging below (46:51); clypeus,

except narrowly impunctate mediolateral corners,

coarsely, rugosely, and nearly confluently punctate

throughout; supraclypeal area, except narrowly
50

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

impunctate sides, about as coarsely punctate as

clypeus; frontal carina narrowly triangular in outline, deeply excavated, slightly longer than maximum diameter of median ocellus; discal surfaces of

metasomal terga I-V basal to marginal depressions

deeply and coarsely punctured, punctures well separated medially, crowded laterally, sparsest on tergum I and closest and densest on tergum V.

TYPES.—Holotype male was collected 5.3 miles

west of Pena Blanca Lake, Santa Cruz County,

Arizona, on 29 July 1972 from the flowers of

Acacia angustissima (Fabaceae) between 1000-1100

(Mountain Standard Time) by T. J. Zavortink,

and is deposited in the collections of the California

Academy of Sciences, San Francisco. The allotype

female was collected one mile east of Douglas,

Cochise County, Arizona, and 20 August 1962,

from the flowers of Verbesina encelioides by M. A.

Cazier and is deposited in the collections of the

California Academy of Sciences.

SPECIMENS EXAMINED, (paratypes).—United States. Arizona.

Cochise County: San Bernardino Ranch, Douglas, 3700 ft,

$, 9 , August (F. H. Snow, CU, PHT); $, same data (N.

Banks, MCZ). Hereford, $, VII-4-58 (Diane Luther, UCD).

Santa Cruz County: Pena Blanca Lake, 9.5 mi WNW Nogales, 4000 ft, $, VIH-18-74, from the flowers of Kallstroemia grandiflora, 0915-0930 MST (T. J. Zavortink, TJZ); $,

same data, except VIII-20-74, 0945-1000 MST (T. J. Zavortink, TJZ); $$, same data, except 1015-1030 MST (T. J.

Zavortink, TJZ).

Mexico. Nayarit: Acaponete, 4 mi S, 2 $, 5 $, IX-2-62

(D. H. Janzen, CIS). Arroyo Santiago, near Jesus Maria, 2$,

VII-5-55 (B. Malkin, CIS). Jesus Maria, $, $, VI-26-55 (B.

Malkin, CIS). San Bias, 17 mi E, $, VII-20-53 (University

of Kansas Mexican Expedition, K.U). Tepic, 18 mi N, 9,

VIII-16-60 (P. H. Arnaud, E. S. Ross, and D. C. Rentz,

CAS). Sinaloa: Campana, 16 mi NW on Hwy 15, 9, IX-13-

64 (A. E. Michelbacher, CIS). Choix, 3.5 mi NW, 9 VII25-69 (T. A. Sears, R. C. Gardner, and C. S. Glaser, UCD).

Choix, 5.5 mi NW, 29 , VII-5-68 (T. A. Sears, R. C. Gardner,

and C. S. Glaser, UCD); 9, VII-25-69 (same collectors, UCD).

Concordia, 9 mi E. 9, VII-7-64 (J. F. McAlpine, CNC).

Culiacan, 49 , VIII-22-60 (R. L. Westcott, LACM). Culiacan,

50 mi N, 9, IX-9-72 (W. Hanson, GEB). Culiacan, 6 mi S,

$, VII-22-54 (M. Cazier, W. Gertsch and Bradts, AMNH).

Culiacan, 30 mi S, $ , 129, VII-24-73, on flowers of Mimosa

polyantha (E. G. and J. M. Linsley, A. E. and M. M. Michelbacher, and J. A. Chemsak, CIS). Culiacancito, 3 mi E, 9,

IX-3-64 (G. R. Noonan, LACM). Elota, 13 mi N, 9, VIII14-60 (P. H. Arnaud, E. S. Ross, and D. C. Rentz, CAS).

Elota, 18 mi S. $, 9, VHI-7-64, on flowers of Croton

culiacanesis (J. A. Chemsak, CIS). Espinal, 14 mi SE, 3 9,

IX-14-64 (A. E. Michelbacher, CIS). Guamuchil, 4 mi S, 9,

VII-7-64 (J. A. Chemsak. CIS). Guamuchil, 18 mi S, 5 $, 39 ,

VIII-7-64 (J. A. Chemsak, CIS); 5$, 29 , same data (J. A.

Powell, CIS). Guamuchil, 19 mi SE, 9, IX-13-64 (A. E.

Michelbacher, CIS). Loreto, 9, VII-17-68 (T. A. Sears, R. C.

Gardner, and C. S. Glaser, UCD). Los Mochis, $, VIII-9-22

(C. T. Dodds, CAS). Los Mochis, 15 mi W, 9 , VHI-28-66

(E. G. and J. M. Linsley and J. A. Chemsak, CIS). Mazatlan,

$, 9, VIII-6-64 (W. R. M. Mason, CNC); $, VII-24-65

(W. J. Gertsch and R. Hastings, AMNH); 9, VIII-9-70

(M. S. and J. S. Wasbauer, CIS); 29 , IX-11-70 (G. E. and

R. M. Bohart, GEB); 9, VII-26-72 (A. E. and M. M. Michelbacher and J. A. Chemsak, (CIS). Mazatlan area, $, IX-5-

71 (W. J. Hanson, GEB). Mazatlan, 2.5 mi N, 9, VIII-10 to

11-70 (J. A. Chemsak, CIS). Mazatlan, 5 mi N, 9, VII-27-64,

on flowers of Buddleja wrightii (J. Chemsak, CIS); 4$, same

data (J. A. Powell, CIS); 21 £, 29 , VII-29-73, on flowers of

Turnera diffusa (E. G. and J. M. Linsley, A. E. and M. M.

Michelbacher, and J. A. Chemsak, CIS). Mazatlan, 9 mi N,

$, 99 , VII-25 to 28-72, on flowers of Solarium sp. (J. A.

and M. A. Chemsak and A. E. and M. M. Michelbacher,

CIS); 5$, 9, VII-26-73 (same collectors). Mazatlan, 35 mi

N, 9, IX-14-64 (A. E. Michelbacher, CIS). Mazatlan, 46 mi

NW, 9, IX-5-71 (W. J. Hanson, GEB). Villa Union, 100 ft,

9, VII-20-53 (University of Kansas Mexican Expedition,

KU). Villa Union, 3 mi E, $, 9, VII-24-72 (J. A. and M. A.

Chemsak and A. E. and M. M. Michelbacher, CIS). Sonora:

Alamos, 9, VIII-10-52 (P. and C. Vaurie, AMNH); $, IX1-60 (R. L. Westcott, LACM); 2 9, IX-7-70 (G. E. and R. M.

Bohart, GEB); 9, IX-5-70 (R. M. Bohart, UCD). Alamos,

W, 63 , IX-4-70 (W. J. Hanson and T. L. Whitworth,

GEB). Alamos, SE, 3$, IX-5-70 (W. J. Hanson and T. L.

Whitworth, GEB). Ciudad Obregon, $, IX-15-70 (E. M.

Fisher, LACM). Estacion Llano, 20 mi N, $, VIII-17-64

(UCR). Estacion Llano, 19.4 mi S, 700 m, 2$, 9, VIII-25-

64 (M. E. Irwin, UCR); $, 9, same data (P. A. Rauch,

UCR); 12 £, same data (E. I. Schlinger, UCR). Guaymas, $,

VIII-5-40 (R. P. Allen, CAS); $, VI-12-62 (D. H. Janzen,

CIS); 2$, VIII-31-68 (R. W. Thorp, UCD). Hermosillo, 29

mi N, 9, VIII-9-60 (D. C. Rentz, LACM). Hermosillo, 40

mi N, $, VIII-21-54 (R. E. Ryckman, C. P. Christianson,

and D. Spencer, USNM). Hermosillo, 15 mi S, $, VIII-21-

65 (W. J. Gertsch and R. Hastings, AMNH). Hermosillo, 30

mi S, 9, IX-3-70, on flowers of Solarium sp. (G. E. and

R. M. Bohart, GEB). Navajoa, 39 , IX-9-70, on flowers of

Solarium sp. (G. E. and R. M. Bohart, GEB). Navajoa, 5 mi

E, 29 . IX-9-70 (R. M. Bohart, UCD). Navajoa, 8 mi E, $,

VIII-28-68 (R. W. and J. T. Thorp. UCD). Navajoa, 13 mi

E, 9, IX-12-64, on flowers of Solarium sp. (A. E. Michelbacher, CIS). Rio Chuchuhagui, 7 mi S, 2$, VII-25 to

VIII-7-53 (F. S. Truxal, LACM). Rio Mayo, 9 , VIII-10-35

(CIS). San Bernardo, Rio Mayo, i$, 9, VIII-16-35 (GEB);

$, VIII-25-35 (UA); 5 $, VIH-25-35 (H. S. Gentry, LACM).

DISCUSSION.—This species inhabits the northwestern Mexican mainland from Nayarit to southern Arizona (Map 2) and has been collected on a

number of occasions with M. arizonica, which occupies a similar geographic range (Map 3). While
NUMBER 220 51

the female of M. rufescens is readily distinguished

by the pale pubescent thorax from that of M.

arizonica, whose thorax is darkly pubescent, the

males of these species are quite similar in general

appearance. However, they may be easily separated

owing to the less heavily infuscated forewings of M.

rufescens as noted in the accompanying key. Mesoxaea rufescens is most closely related to M. vagans,

which is apparently endemic in the Cape Region

of Baja California.

3. Mesoxaea vagans (Fox), new combination

FIGURES 49-52; MAP 2

Oxaea vagans Fox, 1894:119-120, 9 $ [Mexico: Baja California: El Taste, west side and at 3400 ft, San Jose del

Cabo; Texas record applies to Mesoxaea texana, q. v.).—

Friese, 1898:84.—Cockerell and Porter, 1899:411.

Protoxaea vagans.—Linsley and Michener, 1962:385.

LOCATION OF TYPE.—California Academy of

Sciences, San Francisco.

GEOGRAPHIC RANGE (Map 2).—Mexico (Cape Region of Baja California).

MALE.—Length 13-19 mm. Length of forewing,

including tegula, 12-16 mm. Head and body chiefly

black; metasomal terga feebly metallic blue green,

most evident on apical marginal depressions in

bright light; antennae brownish black above, mahogany red below; mandibles dark brownish black

or black, usually reddened on basal half; clypeus

dark brownish black, sometimes somewhat reddened

apically; wings similar in coloration, chiefly or entirely hyaline basally, infuscated with black apically; wing veins dark reddish brown; tegulae testaceous externally, darkening to brownish black

basally; legs chiefly dark brownish black, tarsi dark

reddish brown; tibial spurs reddish brown; claws

deep mahogany red, banded medially with black.

Vestiture of head chiefly ochraceous, somewhat

paler, and often turning to white on hind margins

of head and genae below; pubescence on face usually pale ochraceous or white below antennal sockets, darkening to brownish black on upper part of

face and vertex; thorax thickly clothed with fulvoochraceous pubescence throughout (becoming

white or nearly so in older specimens); legs chiefly

reddish brown pubescent, darkest basally and on

ventral surfaces, usually paling to fulvous on exterior surfaces and on tarsi; pubescence of first

metasomal tergum mostly consisting of long, erect,

whitish plumose hairs, thickest at sides and ventrally, darkening to nearly black over raised submarginal areas laterally; metasomal terga II-IV

without apical bands of pale pubescence, mostly

clothed with short, posteriorly directed, brownish

black pubescence, longest and thickest at sides;

tergum II tufted with white pubescence on ventrally exposed sides and with shorter and less conspicuous erect, white hairs on discal surface medially at base. Metasomal terga III and IV tufted

with long, whitish pubescence on extreme ventrally exposed sides; metasomal tergum V entirely

black pubescent, shortest on punctate surface basally, much longer and forming a transverse band

of simple hairs arising just before impunctate apical margin, projecting posteriorly and terminating

in thick tufts of plumose hairs laterally; metasomal

tergum VI with numerous, long, posteriorly directed, white plumose hairs arising just before impunctate apical margin, longest and somewhat tuftlike laterally, usually intermixed on median tergal

surface with brownish black or black hairs; metasomal tergum VII with pubescence chiefly white

and disposed laterally as posteriorly projecting

tufts of plumose hairs; metasomal sterna thickly

clothed with rather long, plumose hairs, usually

white or cinereous on sterna I—III and reddish

brown or brownish black thereafter.

Eyes strongly convergent above, separated on

vertex by less than distance between hind ocelli

and always by less than distance across hind ocelli

or that between rims of antennal sockets; clypeus

coarsely and nearly confluently punctured throughout, except almost impunctate, on somewhat

smoothed and reddened, narrow, apical margin;

supraclypeal area, except above, densely and confluently punctured, without shining interspaces;

frontal carina somewhat triangular in outline,

deeply excavated basally, nearly as long as diameter

of median ocellus; middle tibiae and basitarsi of

equal length; punctures on discs of metasomal terga I and II dense, rather fine with punctures

separated by mostly less than their diameters, that

on terga III and IV primarily, but larger and

much less densely disposed, especially medially and

with large impunctate or sparsely punctate areas

extending across terga basal to raised submarginal

areas; fifth metasomal tergum with discal punctures similar in size and disposition to those on
52

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

50

FIGURES 4^-52.—Mesoxaea vagans (Fox), male (6 mi N Santa Anita, Baja California, Mexico):

49, seventh metasomal tergum (left half ventral); 50, 51, seventh and eighth metasomal sterna

(left halves dorsal); 52, genitalia (left half ventral).

tergum II, but larger and less densely crowded

especially medially; structure of seventh metasomal

tergum (Figure 49), seventh and eighth metasomal

sterna (Figures 50, 51), and genitalia (Figure 52)

as illustrated.

FEMALE.—Length 16-19 mm. Length of forewing,

including tegula, 15-16.5 mm. Head and body

black, not paling perceptibly on ventrally exposed

surfaces; metasomal terga feebly metallic blue

green with some purplish tints in bright light, most

noticeably so on impunctate surfaces; antennae

dark brownish black, somewhat reddened at apex

of scape, pedicel, and variably below on usually

apical flagellar segments; mandibles black, red-
NUMBER 220 53

dened usually basally and sometimes apically; clypeus brownish black to black, not reddened on medial surface; wings similar in coloration, chiefly or

entirely hyaline basally, infuscated with black apically; wing veins dark reddish brown, irregularly

blackened; tegulae testaceous; legs mostly brownish

black to black, somewhat reddened on tarsi and below; tibial spurs dark reddish brown; claws deep

mahogany red, banded medially with black.

Vestiture of head chiefly moderately long and

white on face, especially that about ocelli, antennal

sockets, clypeus at sides, becoming brownish black

on upper part of face, and on vertex; hind margins

of head thickly clothed with long pubescence,

mostly white, but intermixed with and replaced by

dark brownish and black pubescence dorsally near

and on vertex; thorax thickly clothed with fulvous

pubescence dorsally, usually becoming ochraceous

or whitish posteriorly on metanotum, propodeum,

and on lower sides and ventrally; legs chiefly white

pubescent basally, darker pubescent, usually reddish brown on exterior surfaces and on tarsi; scopal

hairs of hind legs entirely pale, mostly white or

nearly so; discal surfaces of metasomal terga I-IV

devoid of pubescence except for very short, posteriorly directed, brownish black hairs situated laterally, which at extreme sides and on ventrally exposed surfaces are replaced on terga I and II by

scopae consisting of long, densely plumose, mainly

whitish hairs and ventroapically on terga III and

IV with long tufts of chiefly whitish plumose hairs;

metasomal tergum V apically tranversed by a thick

fringe of long, posteriorly directed, plumose hairs

characteristically white at sides and brownish black

on medial tergal surface; metasomal tergum VI

densely clothed on either side of pygidial plate

with moderately long, brownish black plumose

hairs; scopal hairs of metasomal sterna I—III white;

other surfaces of metasomal sterna II-V (except at

extreme sides) and VI thickly clothed with rather

short to moderately long, posteriorly directed, reddish brown to brownish black hairs; extreme sides

of metasomal sterna II-V with long tufts of white

plumose hairs projecting posteriorly from under

overlapping tergal surface.

Eyes weakly diverging below (51:56); clypeus,

except narrowly impunctate mediolateral corners,

coarsely, rugosely, and nearly confluently punctate

throughout; supraclypeal area, except on nearly

impunctate sides, about as coarsely punctate as

clypeus; frontal carina narrowly triangular in outline, deeply excavated, slightly longer than maximum diameter of median ocellus; discal surfaces

basal to marginal depressions of metasomal terga

I-V deeply and coarsely punctured, punctures well

separated medially, crowded laterally, sparsest on

tergum I medially, and closest and densest on tergum V.

SPECIMENS EXAMINED.—Mexico. Baja California: Cabo San

Lucas, 9 , IX-12-67 (A. E. and M. M. Michelbacher and

J. A. Chemsak, CIS). El Triunfo, 10 mi NW, 3 $, 9, VIII5-66, on flowers of Asclepias sp. (P. D. Hurd, CIS). La Paz,

$, X-9-55 (F. X. Williams, CAS). La Paz, 2 mi S, 2 $, 4 $,

VIII-6-66, on flowers of Wislizenia refracta var. mamillata

(P. D. Hurd, CIS); 5^,4$ , same data (J. A. Chemsak, CIS);

$, VIII-11-66 (J. T. Doyen, CIS). La Paz, 14 mi SE, 2$ ,

IX-20-67 (A. E. and M. M. Michelbacher and J. A. Chemsak,

CIS). La Paz, 7 mi SW, 4$, $, VIII-4-66, on flowers of

Wislizenia refracta var. mamillata (J. A. Chemsak, CIS);

3 $, same data (P. D. Hurd, CIS); 2 $, 9, same data (E. G.

Linsley CIS). La Paz, 21 mi W, $, VII-9-66 (J. A. Chemsak,

CIS). La Paz, 26 mi W, $$, 9 , VIII-10-66, one female at

flowers of Larrea tridentata (J. A. Chemsak, CIS); 5,29 ,

VIII-10-66 (J. A. Powell, CIS). Pescadero, $, X-8-41 (E. S.

Ross and G. E. Bohart, CAS). Rancho El Palmar, 2 9, X-22-

72 (E. M. and J. L. Fisher, LACM)T San Jose del Cabo, 9,

IX-11 to 16-67 (A. E. and M. M. Michelbacher and J. A.

Chemsak, CIS). San Jose del Cabo, 5 mi N, 9, IX-14-67

(A. E. and M. M. Michelbacher, CIS). San Jose del Cabo, 6

mi N, $, IX-15-67 (A. E. and Michelbacher and J. A.

Chemsak, CIS). San Jose Viejo, 4$, VII-17-68 (M. A. Cazier

and party, ASU). San Lucas, $, VII-22-68 (M. A. Cazier and

party, ASU). San Pedro, 4$, X-7-41, two males on Compositae (E. S. Ross and G. E. Bohart, CAS). San Pedro, 7

mi S, 2 $, VIII-10-66 (J. T. Doyen, CIS). Santa Anita, 6 mi

N, <J, IX-11-67 (A. E. and M. M. Michelbacher and J. A.

Chemsak, CIS). Santiago, 6 mi S, $, VIII-31-59, in light trap

(K. W. Radford and F. G. Werner, CAS). Santiago, 6 mi W,

S, X-12 to 17-72 (E. M. and J. L. Fisher, LACM). Todos

Santos, 2 $, X-10-41 (E. S. Ross and G. E. Bohart, CAS).

DISCUSSION.—Mesoxaea vagans is evidently endemic to the Cape Region of Baja California. It is

closely related to and very similar in appearance to

M. rufescens, which inhabits the adjacent Mexican

mainland from Nayarit to southern Arizona (Map

2). It is also similar in colorational features to M.

texana, which was considered to be the same as

this species by Fox (1894). Even though the females

of M. rufescens differ principally from those of M.

vagans only by the more densely punctured metasomal terga, the genitalia of the males, especially

the structure of the penis valves and the volsellae,

are strikingly different (Figures 48, 52). In addi-
54

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

tion, the eyes of the male in M. vagans are usually

more strongly convergent above than those of M.

rufescens.

Fox (1894:119-120) named the species from

specimens which had been collected in Baja California (El Taste, west side and at 3400 ft. San Jose

del Cabo, October) and Texas (Cypress Mills).

However, as we have noted elsewhere in this study,

the material from Texas represents another species, viz., M. texana. Since Fox (1894:119-120) did

not select a type specimen for vagans and since the

collections of the California Academy of Sciences

contain the Fox specimens of vagans from El Taste,

we select the male labeled Oxaea vagans, #286 as

the lectoholotype and the female labeled as Oxaea

vagans, no number, as the lectoallotype.

4. Mesoxaea texana (Friese), new combination

FIGURES 53-56; MAP 2

Oxaea texana Friese, 1898:85, $ [Texas: Dallas].

Protoxaea texana.—Cockerell, 1910:275, 9, $ [Texas: Lee

County, at flowers of Polygonum sp.]; 1933:286 [Texas:

Hufsmith, Harris County, at flowers of Croton sp.];

1934a: 8 [Texas: southeast Bexar County, females at flowers

of Monarda punctata in sand dunes, males at flowers of

Aloysia ligustrina]; 1934b: 153, $ $ [Texas: Houston and

San Antonio County].—Crawford, 1915:578, $ [Texas:

Victoria].—Rayment 1935:610.—Linsley, 1951:1086, $ $

[Texas].—Linsley and Michener 1962:385.

Oxaea vagans.—Authors [not Fox 1894:119].—Fox 1894:120

[Texas: Cypress Mills; Baja California record applies to

Mesoxaea vagans, q. v.].—Friese 1898:80, 85, $.—Cameron

1901:316 [Mexico].

LOCATION OF TYPE.—Unknown.

GEOGRAPHIC RANGE (Map 2).—Southern United

States (Louisiana and Texas) and adjacent northeastern Mexico (Nuevo Leon).

MALE.—Length 17-24 mm. Length of forewing,

including tegula, 15-20 mm. Head and body black,

metasomal sterna usually somewhat reddened;

metasomal terga feebly metallic blue green with

purplish tints, most evident on impunctate apical

depressions; antennae brownish black above,

brightly reddened below; mandibles deep mahogany, reddened basally and apically; clypeus dark

brownish black, reddened apically; wings nearly

similar in coloration, forewings hyaline basally,

apical half or more heavily infuscated with brownish black and violaceous in strong light, hind wings

hyaline basally in area of closed cells, beyond these

lightly infuscated with brownish black and somewhat violaceous in bright light; wing veins chiefly

reddish brown, stained irregularly with black; tegulae testaceous, darkening basally to reddish brown;

legs dark reddish brown, tarsi and ventral surfaces

redder; tibial spurs deep reddish yellow; claws mahogany red, irregularly banded, or marked with

black.

Vestiture of head chiefly cinereous or whitish on

face below antennal sockets, and on cheeks below;

pubescence of face above antennal sockets and hind

margins of head, except cheeks below, dark reddish

brown to nearly blackish brown; thorax thickly

clothed with mostly fulvo-ochraceous to fulvous

pubescence (becoming whitish in older specimens)

darkening on sides and below (variably) to dark

reddish brown or brownish black; legs entirely dark

reddish brown pubescent, not paling on exterior

surfaces; pubescence of first metasomal tergum

mostly consisting of long, erect whitish or ochraceous plumose hairs, thickest at sides and ventrally,

darkening to black over raised submarginal areas

laterally; metasomal terga II-IV without apical

bands of pale pubescence, mostly clothed with

short, posteriorly directed, reddish brown or brownish black pubescence, longest and thickest at sides;

tergum II usually tufted with at least some white

pubescence on ventrally exposed sides and with

shorter and less conspicuous white hairs on discal

surface medially at base; terga III and IV without

long tufts of white pubescence on ventrally exposed

sides, rarely and at most with only a few white hairs

on one or more of these terga; metasomal tergum

V entirely black pubescent, shortest on punctate

surface basally, much longer and forming a transverse band of simple hairs arising just before impunctate apical margin, projecting posteriorly and

terminating in thick tufts of plumose hairs laterally; metasomal tergum VI with numerous long,

posteriorly directed, white plumose hairs, usually

intermixed on median tergal surface with brownish black or black hairs; metasomal tergum VII

with pubescence chiefly white and disposed laterally

as posteriorly projecting tufts of plumose hairs;

metasomal sterna thickly clothed with rather long

plumose hairs, usually white or cinereous on sterna

I—II and sometimes also on apical margins of

sterna III and IV, otherwise chiefly or entirely reddish brown pubescent.
NUMBER 220 55

Eyes moderately convergent above, usually separated on vertex by at least minimum distance between hind ocelli and always by less than distance

across hind ocelli or that between rims of antennal

sockets; clypeus coarsely and confluently punctured

at sides and above, becoming less strongly punctate

on slightly depressed anterior half medially and

almost impunctate on smoothed and reddened apical margin; supraclypeal area punctured about as

strongly as adjacent clypeal surface, except above

where almost impunctate, frontal carina narrowly

triangular in outline, deeply excavated basally,

nearly as long as diameter of median ocellus; middle tibiae slightly longer than basitarsi (75:70);

punctation on discs of metasomal terga I and II

dense, rather fine with punctures separated by

mostly less than their diameters, that on terga III

and IV similar, but larger and much less densely

disposed, especially medially and with larger impunctate or sparsely punctate areas extending

across terga on and basal to raised submarginal

areas; fifth metasomal tergum with discal punctures

similar in size and disposition to those on tergum

II, but larger and less densely crowded especially

medially; structure of seventh metasomal tergum

(Figure 53), seventh and eighth metasomal sterna

(Figures 54, 55,), and genitalia (Figure 56) as

illustrated.

FEMALE.—Length 20-24 mm. Length of forewing

including tegula, 16-18 mm. Head and body black,

not paling perceptibly on ventrally exposed surfaces; metasomal terga feebly metallic blue green

with some purplish tints in bright light and most

noticeable on impunctate surfaces; antennae dark

reddish brown to brownish black, reddened below

and at apex of scape, pedicel, and basal flagellar

segments; mandibles chiefly black, reddened basally

and apically; clypeus black, sometimes reddened

medially; wings somewhat similar in coloration,

chiefly or entirely hyaline basally, infuscated with

black apically, especially heavily so on forewings,

violaceous in bright light; wing veins dark reddish

brown, tinged irregularly with black; tegulae testaceous externally, dark yellowish brown basally; legs

dark reddish brown, tarsi and ventral surfaces redder; claws deep mahogany red, mottled with black.

Vestiture of head chiefly cinereous or whitish on

face below antennal sockets and on cheeks below;

pubescence of face around antennal sockets and

above including vertex and hind margins of head,

except lower genal areas, dark reddish brown to

nearly blackish brown; thorax thickly clothed with

mostly fulvo-ochraceous to fulvous pubescence

(whitish in older specimens), darkening on sides

and below (variably) to dark reddish brown or

brownish black; pubescence on thorax behind usually paler, ochraceous to nearly white; legs entirely

dark reddish brown pubescent, not paling on exterior surfaces; scopal hairs on hind femora entirely pale, mostly white or nearly so, those on hind

coxae, trochanters, and tibiae largely or entirely

dark fuscous; discal surfaces of metasomal terga

I-IV devoid of pubescence except for very short,

posteriorly directed, brownish black hairs situated

laterally, which at extreme sides and on ventrally

exposed surfaces are replaced on terga I and II by

scopae consisting of long, densely plumose, mainly

whitish hairs and ventroapically on terga III and

IV with moderately long tufts of reddish brown

plumose hairs; metasomal tergum V apically traversed by a thick fringe of long, posteriorly directed, plumose hairs characteristically white at

sides and brownish black on medial tergal surface;

metasomal tergum VI densely* clothed on either

side of pygidial plate with moderately long, brownish black plumose hairs; scopal hairs of metasomal

sterna I—III white or chiefly so; other surfaces of

metasomal sterna II-V (except extreme sides) and

VI thickly clothed with rather short to moderately

long, posteriorly directed, reddish brown to brownish black hairs; extreme sides of metasomal sterna

II-V with long tufts of white hairs projecting posteriorly from under overlapping tergal surfaces.

Eyes weakly diverging below (58:60); clypeus,

except narrowly impunctate mediolateral corners,

coarsely and rather rugosely punctate, although

punctures medioapically separated and with some

shiny interspaces; supraclypeal area for the most

part with punctures well separated and largely absent on sides and above; frontal carina narrowly

triangular in outline, deeply excavated, slightly

longer than maximum diameter of median ocellus;

discal surfaces basal to marginal depressions of

metasomal terga I-V, deeply and coarsely punctured, punctures well separated medially, crowded

laterally, sparsest on tergum I medially, and closest

and densest on tergum V.

SPECIMENS EXAMINED.—United States. Louisiana: Lafayette

Parish: Southwestern Lafayette, $, XI-27-39 (C. Guidroz,

USNM).
56

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

56

54

FIGURES 55-56.—Mesoxaea texana (Friese), male (Falfurrias, Brooks County, Texas): 53, seventh

metasomal tergum (left half ventral); 54, 55, seventh and eighth metasomal sterna (left halves

dorsal); 56, genitalia (left half ventral).

Texas. Bastrop County: $, VIII-31-51, on flowers of

perennial Monarda sp. (A. H. Alex, TAM). Bexar County:

9, VII-20-31 (H. B. Parks, TAM); 4$ , X-12-31 (H. B.

Parks, TAM); 9, same data (H. B. Parks, U. Colo.); $,

IX-6-36 (TAM); 2$, 4$ , no date (TAM). Brazos County:

9, VII-18-51, on flowers of Gossypium sp. (A. H. Alex,

TAM); 9, VII-31-51, on flowers of Sesamum indicum (A. H.

Alex, TAM); 29 , VII-1-51, on flowers of Sesamum indicum

(A. H. Alex, TAM); 9, VIII-3-51, on flowers of Sesamum

indicum (A. H. Alex, PHT); $, VIII-6-54, on flowers of

Eupatoriutn serotina (A. H. Alex, TAM); $, VII-22-55, on

flowers of Vernonia texana (A. H. Alex, TAM). College Station, 9, VIII-31-39 (USNM). Millican, $$, VI-17-56 (H. E.

Evans and E. G. Matthews, CU). Brooks County: Falfurrias,

9 , VH-7-65 (J. E. Gillaspy, GEB); &, VI-23-66 (J. E.

Gillaspy, GEB); 2<$, VI-5-67, on flowers of Monarda sp.

(J. E. Gillaspy, GEB). Burleson Co.: 3 9, IX-24-56, on flowers

of Solanum rostratum (A. H. Alex, TAM). Harris Co.: $,

VIII-13-28 (L. D. Beamer, KU). Dickinson, 2$, VI-1929

(F. M. Hull, TAM). Rose Hill, £, 9, VIII-20-32 (R. Birk-
NUMBER 220 57

mann, CAS, PHT). Jackson County, $, VIII-9-28 (J. G.

Shaw, KU). Kenedy County: Los Olmos Creek, $, $, VI7-48 (M. A. Cazier, AMNH). Mittilin, 4$, $, VII-6-52

(H. E. Cott, CIS, PHT). Kleberg County: $, IX-21-73

(J. E. Gillaspy, TAI). Lee County: $, $, VIII-10-07, on

flowers of Polygonum sp. (MCZ, PHT); $, $, VIII-26-07

(MCZ, PHT). Robertson Co.: Robertson, $, VII-10-55, on

flowers of Verbena halei (A. H. Alex, TAM). Uvalde Co.:

Concan, $, VII-6-36 (M. B. Jackson, KU). Victoria County:

Victoria, $, IX-8-11 (J. D. Mitchell, USNM; $, VIII-20-18

(J. D. Mitchell, USNM); $, VI-21-56 (H. E. Evans and

E. G. Matthews, CU). Washington County: Brenham, 11

mi NE, 6,$, VI-17-56 (H. E. Evans and E. G. Matthews, CU).

Mexico. Nuevo Leon: Vallecillo, 5 3 , VI-2 to 5-51 (P. D.

Hurd, CIS).

DISCUSSION.—Superficially this species closely resembles Mesoxaea vagans and M. rufescens in

colorational features, but is structurally most

closely related to M. nigerrima. The amount of

pale vestiture on the head and thorax is subject

to some variation in both the males and the females. In some specimens there is much pale pubescence on the face above the antennae and on

the sides and venter of the thorax, while in others

this pubescence is largely or entirely dark in

coloration.

Mesoxaea texana has been observed nesting in

very large numbers in Hufsmith, Harris County,

Texas, by the Reverend Birkmann (1932), who

published a newspaper article about his observations in The Giddings News. These observations

were subsequently summarized in part by Cockerell

(1933), who identified the species, and by Rayment

(1935), who mentioned these observations. Since

the original information contained in the Reverend

Birkmann's article has never been reproduced and

has entailed a special effort through the Library of

Congress to obtain the article, it is presented verbatim below so that it will be conveniently available for use by future investigators:

I will write now of a novel experience in natural history

which I had some few days ago right here in Harris County.

Ten days ago we had a storm, a hurricane they called it

in the papers. We however suffered no damage. Hufsmith

is in the northern part of the county, and we were only

skirted by the storm. We were fortunate in getting rain

after a pretty long drouth. The rain brought new life to

plants and animals, even the insects in the ground got busy

and began to come out and breathe some fresh air.

I am especially referring to a great swarm of wild bees

that for several days had a holiday in the air and, for a

change, made their burrows in the sand.

Now, wild bees are not honey bees that are occasionally

found in the woods, but by this term are meant all species

of insects that are related to the honeybees, for instance,

the bumble bees and the carpenter bees and many others.

In Lee County there are several hundred different species

of wild bees. The bumble bees are the only ones of the

"wild bees" that live in colonies, all the rest live in pairs,

each pair has its own burrow or nest. Some burrow in wood,

or in stems or weeds, others dig into the ground, where they

lay their eggs and have their brood, some once a year, others

twice.

Now I am coming to my subject more closely. Some days

ago my son, with whom I am now staying, tells me that he

has seen a lot of bees or wasps in the air and they are making a great ado and hubbub flying in all directions, and not

only over a small area, but for quite a djptance the noise is

heard and the ground is full of newly dug holes. Next

morning when the day grows warm we go out and I found

all this to be true, and not the half told. I could not see the

insects on account of my poor eyesight, but my son tells

me all I want to know. He tells me the bees are flying, some

of them only a half foot from the ground, others fly higher,

so as to get over the weeds, still others are pretty high up,

all of them flying fast and like so many little furies, making

all the noise passible [sic] to them. You could hear them at

some distance, the sound was similar to that heard in the

telegraph wires at times, but much louder, and to listen to

them for awhile would make me a little nervous. The question was: What is all that about? What are these hundreds

of thousands of bees after? And the ground was as stated,

full of holes. Fresh earth was lying beside these holes, you

could see the bees had just opened them. My grandson Roger

probed some of them with a switch, and found them going

down several feet, others not quite that far, the diameter

of the holes being about half an inch. The size of the female

bee is about that of a queen bee in the hives. The male is

much smaller, insignificant in appearance. It seems to take

no part in the digging and procuring a home for his family.

Roger caught about a dozen of these bees for me and I

have sent specimens for determination to Prof. Cockerell of

Boulder, Col., who is a great scholar and student of the

bees, knows all of the thousands of bees in the world, and

has collected and described many of them in scientific books

and periodicals. I suppose that the species of which I am

writing is one new to science, at least, I have never before

observed it in Lee County. And if it should prove to be a

known species, I think that their habit of building so many

homes near one another, and of swarming in such great

numbers, and with so much noise and fury has not been

observed before. I have never before seen anything like such

a great area occupied by the bees, and never before heard

such sounds produced by thousands and thousands of these

insects.

I stepped off the ground, it was about 700 feet in length,

and 230 feet in width, I counted (or rather Roger did) 29

holes in ten feet square (100 sq. ft.) and 46 holes in another ten feet square. According to this reckoning there were

about 80,000 holes and counting one pair of bees for each
58 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

hole, there would have been one hundred thousand individual bees. All of the same kind.

This spectacle lasted for about four or five days, according

to my observation. After that time, there were only a few

speciments [sic] to be noticed. They are in their burrows

now and the females will come out occasionally to gather

food for their offspring.

What food? I don't know, I don't think that anyone knows.

Bees don't collect animal food like the wasps do, but I suppose their subsistence is on sweets which they get from plants

and flowers.

G. Birkmann

(This is a very interesting article, and the editor is glad

Rev. Birkmann is yet able to find delight in nature, as he

did for so many years at Fedor.)

Friese (1898:85) cites two males with his original

description of this species and we have not been

able to locate their whereabouts.

5. Mesoxaea nigerrima (Friese), new combination

FIGURES 57-60; MAP 3

Oxaea nigerrima Friese, 1912:199, $ [Mexico: Vera Cruz].

Protoxaea nigerrima.—Linsley 1951:1086.—Linsley and Michener, 1962:386, fig. [Mexico: 14 miles south of Cuernavaca].

LOCATION OF TYPE.—Zoologisches Museum,

Humboldt-Universitat, Berlin.

GEOGRAPHIC RANGE (Map 3).—Southwestern to

southern Mexico (Chiapas, Colima, Guerrero,

Jalisco, Michoacan, Morelos, Oaxaca and Vera

Cruz).

MALE.—Length 16-22 mm. Length of forewing,

including tegula, 15-17 mm. Head and body chiefly

or entirely black; metasomal terga feebly metallic

blue green, most evident on impunctate apical marginal depressions; antennae dark reddish brown to

brownish black above, deep reddish brown basally,

becoming deep mahogany red or black apically;

clypeus black, reddened along apical margin; wings

dissimilar in coloration, forewings nearly uniformly

and heavily infuscated throughout with black,

feebly violaceous in bright light, hind wings much

paler, hyaline basally and lightly infuscated with

black apically and posteriorly; wing veins dark reddish brown; tegulae dark reddish brown, black

basally; legs brownish black to black on coxae to

femora, tibiae and tarsi reddish brown; tibial spurs

pale reddish brown; claws deep mahogany red, irregularly marked with black.

Vestiture of head chiefly ochraceous to nearly

white on face below antennal sockets and on lower

hind margins of head (variable); pubescence on

face above antennal sockets chiefly brownish black

to black, sometimes pubescence just above antennal sockets intermixed with some fulvous hairs;

vertex and hind margins of head dorsally (variable)

chiefly or entirely brownish black; thorax thickly

clothed with fulvo-ochraceous to fulvous pubescence, usually paling to ochraceous, somewhat posteriorly and usually darkening partly on lateral

and ventral surfaces (variably) to brownish black;

legs with chiefly brownish black to fulvous pubescence basally, paling to bright fulvous on tarsi;

pubescence of first metasomal tergum consisting of

long, erect whitish plumose hairs, thickest at sides

and on ventrally exposed surfaces; metasomal terga

II-IV, without apical bands of pale pubescence,

mostly clothed with short, posteriorly directed,

brownish black pubescence, (except basal discal

surface of tergum II with some erect white hairs),

longest and thickest at sides; metasomal tergum V

mostly clothed with brownish black pubescence,

turning to white at extreme sides, shortest on punctate surface basally, much longer and forming a

transverse band of long simple hairs arising just

before impunctate apical margin, projecting posteriorly and terminating in small tufts laterally;

metasomal tergum VI with numerous long, posteriorly directed, mostly or entirely white plumose

hairs arising just before apical margin, longest and

somewhat tuftlike laterally and usually intermixed

medially with some brownish black or black hairs;

metasomal tergum VII with pubescence chiefly or

entirely white (sometimes intermixed or replaced

medially with brownish black or black hairs) and

disposed laterally as posteriorly projecting tufts of

plumose hairs; metasomal sterna thickly clothed

with rather long plumose hairs, usually white on

sterna I—III and brownish black on succeeding

sterna, which in some specimens may display variable amounts of pale pubescence apically and

laterally.

Eyes moderately convergent above, separated on

vertex by nearly or more than minimum distance

between hind ocelli, but by less than distance between rims of antennal sockets; clypeus with a

rather broad, reddened, impunctate, apical margin

medially, otherwise densely and confluently punctured on sides, median surface somewhat depressed

and with punctures larger and well separated,
NUMBER 220 59

interspaces smooth and shiny; supraclypeal area

densely, finely, and confluently punctured; frontal

carina triangular in outline, very short, much

shorter than diameter of median ocellus, deeply

excavated especially basally; middle tibiae slightly

longer than basitarsi (70:65); punctation on discs

of metasomal terga I and II dense, rather fine, with

punctures mostly separated by less than their diameters, those on terga III and IV similar but less

densely punctate, especially medially and laterally

over and basad to raised submarginal areas; fifth

metasomal tergum with punctures similar in size

and disposition to those on tergum II, but less

densely crowded especially medially; structure of

seventh metasomal tergum (Figure 57), seventh

and eighth metasomal sterna (Figures 58, 59), and

genitalia (Figure 60) as illustrated.

FEMALE.—Length 18-23 mm. Length of forewing,

including tegulae, 16-17.5 mm. Head and body

black, somewhat paler on metasomal sterna; metasomal terga feebly metallic blue green with some

purplish tints evident in bright light and most

visible on impunctate apical margins; antennae

dark reddish brown to brownish black, somewhat

reddened at apex of scape and pedicel and to a

variable extent on flagellar segments, especially

below; mandibles chiefly deep mahogany red to

black, especially basally and sometimes apically;

clypeus black, narrowly reddened apically; wings

dissimilar in coloration, forewings weakly violaceous, nearly uniformly infuscated throughout

with brownish black, but slightly less so basally,

hind wings hyaline or nearly so, slightly violaceous

in bright light; wing veins chiefly brownish black,

sometimes somewhat reddened; tegulae dark reddish brown; legs mostly blackish brown dorsally,

usually reddened below on femora, tibiae, and tarsi;

tibial spurs deep reddish brown; claws reddish basally, blackened apically.

Vestiture of head chiefly moderately long and

white on face about antennal sockets and clypeus

at sides; face above antennal sockets rather thickly

clothed with long pubescence, mostly white pubescence on cheeks at sides, becoming intermixed with

and replaced by dark brownish and black pubescence dorsally; thorax thickly clothed with dark

brownish black pubescence; legs, except whitish

scopal hairs on hind femora, clothed with rather

long brownish to blackish pubescence; discal surfaces of metasomal terga devoid of pubescence except for very short, posteriorly directed, brownish

black hairs situated laterally, which at extreme

sides and on ventrally exposed surfaces are replaced on terga I and II by scopae consisting of

long, densely plumose, mostly whitish hairs and

ventroapically on terga III and IV with tufts of

brownish black plumose hairs usually intermixed

with some white hairs; metasomal tergum V apically traversed by a thick fringe of long, posteriorly

directed, plumose hairs characteristically white at

sides and brownish black on medial tergal surface;

metasomal tergum VI densely clothed on either

side of pygidial plate with moderately long, brownish black, plumose hairs; scopal hairs of metasomal

sternum I medially white, laterally brownish black,

those on sternum II and median tuft on sternum

III largely or entirely white; other surfaces of

metasomal sterna II-V (except extreme sides) and

VI thickly clothed with rather short to moderately

long, posteriorly directed, reddish brown to brownish black hairs; extreme sides of metasomal sterna

II-V with long tufts of white plumose hairs projecting posteriorly from under overlapping tergal

surfaces.

Eyes weakly diverging below (51:56); clypeus

densely and rather rugosely punctured throughout,

punctures on median surface larger than those on

sides, and with rugae polished and shining; supraclypeal area densely and nearly confluently punctured except at sides and above; frontal carina

narrowly triangular in outline, short, longer than

maximum diameter of median ocellus, deeply excavated and polished; discal surfaces before marginal

depressions of metasomal terga I-V deeply punctured virtually throughout, punctures well separated medially, crowded laterally, sparsest on tergum I medially and closest and densest on tergum

V.

SPECIMENS EXAMINED.—Mexico. Chiapas: El Sumidero, $,

IX-14-74 (G. E. Bohart and W. J. Hanson, GEB). Colima:

Colima, 10 mi W, $, VIII-1-54 (M. A. Cazier, W. J. Gertsch,

and Bradts, AMNH). Guerrero: Iguala, 3 mi S, $, VII-12-

70 (E. M. Fisher and P. Sullivan, LACM). Mexcala (vicinity), 1 mi S Rio Balsas on Hwy 95, 2 $, IX-3 to 4-70 (E. M.

and J. M. Fisher, LACM). Rio Balsas, 3.7 mi N on Hwy 95,

4 9 , VIII-5-63 (G. W. Byers and party, KU). Jalisco: Plan

de Barrancas, 3 mi SE, 2 $, VII-S-63 (F. D. Parker and L. A.

Stange, UCD); 2$ , VIII-25-63 (F. D. Parker and L. A.

Stange, UCD). La Quemada, 39 , VII-27-54 (M. A. Cazier,

W. J. Gertsch and Bradts, AMNH). Tequila, 3 mi NW, 4000

ft, S, VIII-19-53, on flowers of Vitex pyramidata (University
60

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

58

FIGURES 57-60.—Mesoxaea nigerrima (Friese), male (21 mi W Tehuantepec, Oaxaca, Mexico):

57, seventh metasomal tergum (left half ventral); 58, 59, seventh and eighth metasomal sterna

(left halves dorsal); 60, genitalia (left half ventral).

of Kansas Mexican Expedition, KU). Tequila, 8 km W, 4 $,

VII-18-51 (H. E. Evans, KU); 4 $ , $, same data (P. D. Hurd,

CIS). Michoacan: Apatzingan, 1200 ft, $, VIII-14-41 (H.

Hoogstraal, AMNH); $, VIII-21-41 (H. Hoogstraal, AMNH).

Apatzingan, 11 mi E, $, VIII-20-54 (E. G. Linsley, J. W.

MacSwain, R. F. Smith, CIS). Morelos: Alpuyeca, $, VII5-51 (H. E. Evans, KU). Cuautla, 10 mi W, 3$, $, VII22-62 (H. E. Milliron, CNC). Cuernavaca, 14 mi S, 3250 ft,

$, VIII-3-54 (C. D. Michener, Linsley, and Michener, 1962:

386). Tequesquitengo, 3 mi NW, 1$, $, VIII-16-62, on

flowers of Kallstroemia sp. (R. B. Roberts and N. Marston,

KU). Yautepec, 3$ , VII-13-63 (F. D. Parker and L. A.

Stange, UCD). Yautepec, 4 mi SW, 3800 ft, $, VII-2-61, on

flowers of Cassia sp. (C. D. Michener, KU). Oaxaca: El

Camaron, 2400 ft, 16$, VII-7-53 (University of Kansas

Mexican Expedition, KU). Chivela, 12 mi S, 2 $, $, VIII18-59 (A. A. Menke and L. A. Stange, UCD). S. Geronimo,

$, VIII-5-23 (E. G. Smyth, USNM). Matias Romero, 23 mi

S, 5$, VIII-14-63 (F. D. Parker and L. A. Stange, UCD).

Tehuantepec, 2 $, IX-16-74 (G. E. Bohart and W. J. Han-
NUMBER 220 61

son, GEB). Tehuantepec, 21 mi W, VII-21-52 (E. E. Gilbert

and C. D. MacNeill, CIS). Tehuantepec, 35 mi W, $, IX16-74 (G. E. Bohart and W. J. Hanson, GEB). Vera Cruz:

Cotaxtla Experiment Station, $, VII-17-62, one hour before

to one-half hour after sunrise, plant 516 (D. H. Janzen,

CIS). Vera Cruz, $, no date (F. Baker, LACM).

DISCUSSION.—This species, which occurs in the

southernmost area occupied by the genus Mesoxaea (Map 3), shares many characters with M.

arizonica and M. texana. As presently known these

species are allopatric in distribution and it appears

they were derived from a common ancestry. In

colorational features M. nigerrima closely resembles

M. arizonica from which it may be readily distinguished by the accompanying key. Although the

geographic ranges of M. nigerrima and M. arizonica are nearly contiguous in southwestern Mexico

there is a large geographic hiatus separating the

ranges of M. texana and M. nigerrima in the gulf

coast region of eastern Mexico (Maps 2,3). Most

likely this comparatively wide separation of their

known ranges is due to a lack of collecting.

6. Mesoxaea arizonica (Cockerell),

new combination and new status

FIGURES 61-64; MAP 3

Protoxaea nigerrima arizonica Cockerell, 1936:6, $ [Arizona:

Tumacacori Mts.].—Linsley, 1951:1086, $ [Arizona].

LOCATION OF TYPE.—American Museum of Natural History, New York.

GEOGRAPHIC RANGE (Map 3).—Southwestern

United States (Arizona) and adjacent northwestern

Mexico (Nayarit, Sinaloa, and Sonora).

MALE.—Length 16-20 mm. Length of forewing,

including tegula, 14-16 mm. Head and body chiefly

or entirely black; metasomal terga faintly metallic

blue green, most evident on impunctate, apical,

marginal depressions; antennae dark reddish

brown to brownish black above, reddened below

and scape usually somewhat reddened also; mandibles dark reddish brown basally, becoming deep

mahogany red or black apically; clypeus black,

reddened along apical margin; wings dissimilar in

coloration, forewings, except pale basal cells (R

and 1st Cu), uniformly and heavily infuscated

throughout with black, feebly violaceous in bright

light, hind wings entirely hyaline or nearly so;

wing veins dark reddish brown; tegulae reddish

brown to nearly testaceous; legs brownish black to

black on coxae to femora, tibial and tarsi dark

reddish brown; claws deep mahogany red, irregularly blackened.

Vestiture of head chiefly pale ochraceous to

white on face below antennal sockets and on hind

margins of head; pubescence on face above antennal sockets ochraceous to fulvous (variable) turning

to brownish black to black on top of face and

vertex; pubescence on hind margins of head adjacent to vertex chiefly or entirely white, sometimes intermixed with a few brownish black or

black hairs; thorax thickly clothed with ochraceous to fulvous pubescence (variable), usually

paling to white or pale ochraceous posteriorly on

propodeum and sometimes darkening partly on

lateral and ventral surfaces (variably) to brownish

black; legs with chiefly brownish black pubescence

basally, paling to fulvous or ochraceous on tarsi,

sometimes also on hind femora; pubescence of

first metasomal tergum consisting of long, erect

whitish plumose hairs, thickest at sides and on

ventrally exposed surfaces; metasomal terga II-IV,

without apical bands of pale pubescence, mostly

clothed with short, posteriorly directed, brownish

black pubescence (except basal discal surface of

tergum II with some erect white hairs), longest

and thickest at sides; metasomal tergum V mostly

clothed with brownish black pubescence, turning to

white at extreme sides, shortest on punctate surfaces basally, much longer and forming a transverse band of long simple hairs arising just before

impunctate apical margin, projecting posteriorly

and terminating in small tufts laterally; metasomal

tergum VI with numerous long, posteriorly directed, mostly or entirely white plumose hairs

arising just before apical margin, longest and somewhat tuftlike laterally and usually intermixed medially with some brownish black or black hairs;

metasomal tergum VII with pubescence chiefly or

entirely white (sometimes intermixed or replaced

medially with brownish black or black hairs) and

disposed laterally as posteriorly projecting tufts of

plumose hairs, usually white on at least sterna

I—III and brownish black on succeeding sterna,

which in some specimens may display variable

amounts of pale pubescence apically and laterally.

Eyes moderately to strongly convergent above,

separated on vertex usually by much less than minimum distance between hind ocelli and always by
62

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

considerably less than distance between rims of

antennal sockets; dypeus with a rather broad, reddened, impunctate, apical margin medially, otherwise densely and confluently punctured on sides,

median surface somewhat depressed and with punctures much larger, well separated and with interspaces smooth and shining; supraclypeal area

densely, finely, and confluently punctured; frontal

carina narrowly triangular in outline, short, about

as long or longer than maximum diameter of median ocellus, deeply excavated virtually throughout

its length; middle tibiae and basitarsi of equal

length; punctation on discs of metasomal terga I

and II dense, rather fine with punctures separated

by mostly less than their diameters, those on terga

III and IV similar, but less densely punctate especially medially and laterally over and basal to raised

submarginal areas; fifth metasomal tergum with

punctures similar in size and disposition to those

on tergum II, but less densely crowded especially

medially; structure of seventh metasomal tergum

(Figure 61), seventh and eighth metasomal sterna

(Figures 62, 63), and genitalia (Figure 64) as

illustrated.

FEMALE.—Length 18-23 mm. Length of forewing

including tegula 15-17 mm. Head and body black,

metasomal sterna somewhat paler, dark brownish

black; metasomal terga feebly metallic blue green

with some purplish tints evident in bright light and

most visible on impunctate apical margins; antennae dark reddish brown, scape and pedicel reddened narrowly at apices and to some extent on

ventral surfaces of flagellar segments; mandibles

dark brownish black, reddened basally and at apex;

clypeus black, narrowly margined with red; wings

dissimilar in coloration, forewings weakly violaceous, nearly uniformly and heavily infuscated

with brownish black, slightly less so in basal cells

(R and 1st Cu), hind wings hyaline basally, faintly

infuscated apically and scarcely violaceous in

bright light; wing veins brownish black; legs mostly

brownish black dorsally, somewhat paler below;

tibial spurs bright reddish brown; claws chiefly

black, reddened basally.

Vestiture of head short to moderately long, and

mostly white on face about antennal sockets and

clypeus at sides; face above antennal sockets rather

thickly clothed with erect reddish brown pubescence; hind margins of head thickly clothed with

long pubescence mostly white on cheeks at sides,

usually becoming intermixed with and replaced by

brownish black pubescence dorsally (variable); thorax thickly clothed with brownish black pubescence, that on metanotum sometimes paling marginally to a fringe of fulvous hairs; legs, including

scopal hairs, clothed entirely with rather long,

brownish black pubescence; discal surfaces of metasomal terga devoid of pubescence except for a very

few, short, posteriorly directed, brownish black

hairs situated laterally, which at extreme sides and

on ventrally exposed surfaces are replaced on terga

I and II by scopae consisting of long, densely plumose reddish brown hairs (often intermixed with

some white hairs) and ventroapically on terga III

and IV with tufts of brownish black plumose hairs

sometimes intermixed with white hairs; metasomal

tergum V apically traversed by a thick fringe of

long, posteriorly directed, plumose hairs characteristically white at sides and brownish black on medial tergal surface; metasomal tergum VI densely

clothed on either side of pygidial plate with moderately long, brownish black, plumose hairs; scopal

hairs on sternum I brownish black, those on sternum II and median tuft on sternum III variably

either chiefly whitish or mostly brownish, but with

some whitish hairs; other surfaces of metasomal

sterna II-V (except extreme sides) and VI thickly

clothed with rather short to moderately long, posteriorly directed, reddish brown to brownish black

hairs; extreme sides of metasomal sterna II-V with

long tufts of white plumose hairs projecting posteriorly from under overlapping tergal srufaces.

Eyes weakly diverging below (48:54); clypeus

densely and rugosely punctured especially on sides,

punctures on median surface larger than those at

sides and much less crowded and with interspaces

shining; supraclypeal area densely and nearly confluently punctured, except at sides and above; frontal carina narrowly triangular in outline, short,

longer than maximum diameter of median ocellus,

deeply excavated basally; discal surfaces before

marginal depressions of metasomal terga I-V

deeply punctured, punctures well separated medially, crowded laterally and closest and densest on

tergum V.

SPECIMENS EXAMINED.—United States. Arizona. Santa Cruz

County: Atascosa Mountains, 5.6 mi W Pena Blanca Lake,

4800 ft, 2$, VIII-15-74, on flowers of Acacia angustissima,

1200-1215 MST (T. J. Zavortink, TJZ). Nogales, $, VIII24-39 (R. H. Crandall, LACM). Pena Blanca, 2$ , IX-3-68
NUMBER 220 63

61

62

FIGURES 61-64.—Mesoxaea arizonica (Cockerell), male (Sonoita, Santa Cruz County, Arizona): 61,

seventh metasomal tergum (left half ventral); 62, 63, seventh and eighth metasomal sterna (left

halves dorsal); 64, genitalia (left half ventral).

(W. J. Hanson, GEB). Pena Blanca Lake, 9.5 mi WNW of

Nogales, 4000 ft, 55 4 , $, VIII-18-74, on flowers of Kalhtroetnia grandiftora from 0830 to 1200 MST (T. J. Zavortink,

TJZ); $, VIII-19-74, on flowers of Tephrosia leiocarpa, 1515-

1530 MST (T. J. Zavortink, TJZ); 29 £, VIII-20-74, on

flowers of Kallstroemia grandifiora from 0830-1145 (T. J.

Zavortink, TJZ); 5$, VIII-20-74, on flowers of Tephrosia

leiocarpa, 1145-1200 (T. J. Zavortink, TJZ). Pena Blanca

Lake, 5.3 mi W, 2 $ , VII-29-72, on flowers of Acacia angustissima, between 1000-1215 MST (T. J. Zavortink, TJZ). Sonoita,

6 mi SE, $, VIII-27-68 (R. W. Thorp, UCD). Tumacacori

Mountains, 3$ , no date, on flowers of Lotus sp. (I. Wilson,

AMNH, CAS, KU). Nayarit: Acaponete, $, IX-29-66 (G. E.

& A. S. Bohart (GEB). Acaponete, 8 mi SE, $, X-19-64 (A. E.

Michelbacher, CIS). Navarrete, 2$, $, VII-28-53 (C and P.

Vaurie, David Rockefeller Mexican Expedition, 1953). Tepic,

44 mi NW Tepic, $, VIII-30-71 (W. J. Hanson, GEB).

Sinaloa: Arroyo del Saucillo, 4 mi NW of Choix, 2 $, VIII-
64

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

26-68 (T. A. Sears, R. C. Gardner and C. S. Glaser, UCD).

Choix, 5.5 mi NW, $, 9, VII-5-68 (T. A. Sears, R. C.

Gardner, and C. S. Glaser, UCD). Concordia, $, VIII-1-70

(M. S. Wasbauer, CIS). Concordia, 13 mi E, 9, VIII-5-64

(J. F. McAlpine, CNC). Concordia, 10 mi W, 4$ , X-30-65

(G. E. and A. S. Bohart, GEB). Escuinapa, 5$, IX-29-66

(G. E. and A. S. Bohart, GEB). Esquinapa, 33 km SE, 6$ ,

IX-11-74 (E. M. Fisher, LACM). Mazatlan, $, IX-15-18

(J. A. Kusche, USNM); $, X-28-65 (G. E. and A. S. Bohart,

GEB). Mazatlan, 5 mi N, 2 $, VIII-9 to 15-70 (J. A. Chemsak,

CIS); 6,$, 3$ , VII-29-73, flowers of Turnera diffusa (J. A.

Chemsak, E. G. Linsley, and A. E. and M. M. Michelbacher,

CIS); $, VII-28-64, on flowers of Buddleja wrightii (J. A.

Powell, CIS); 2 $, VII-30-73, on flowers of Buddleja wrightii

(J. A. Chemsak, E. G. and J. M. Linsley, and A. E. and M. M.

Michelbacher, CIS). Mazatlan, 35 mi N, 9, IX-14-64 (A. E.

and M. M. Michelbacher, CIS). Mazatlan, 40.2 mi N, $,

VIII-30-64 (G. R. Noonan, LACM). Mazatlan, 46 mi NW,

4 $ , IX-5-71 (W. J. Hanson, GEB). Piaxtla, $ , VIII-19-54

(R. E. Ryckmarin, D. Spencer, and C. P. Christianson,

USNM); $, IX-14-64 (A. E. and M. M. Michelbacher, CIS).

Venadillo, 9, IX-26-50 (R. F. Smith, AMNH). Villa Union,

6 mi N, 9, IX-9-70 (W. J. Hanson and T. L. Whitworth,

GEB). Villa Union, 24 mi E, $, VII-24-64 (J. A. Powell,

Cis). Sonora: San Bernardo, 9, VII-16-35 (GEB).

DISCUSSION.—The females of this species, like

those of Mesoxaea nigerrima but unlike those of

the other species of the genus, have the thorax

entirely dark pubescent. They may be easily distinguished from each other since the scopal hairs of

hind femora in M. arizonica are entirely dark while

those of M. nigerrima are mostly white. The males

of these species are strikingly similar in appearance, but the forewings of M. arizonica, though

heavily infuscated with brownish black like those of

M. tiigerrima, differ by being hyaline or nearly so

in cells R and 1st Cu. Mesoxaea arizonica is most

closely related to M. nigerrima and may have been

derived from the same ancestry which led to the

formation of that species and M. texana.

7. Mesoxaea clypeata, new species

FIGURES 5, 65-68; MAP 3

MALE.—Length 16-21 mm. Length of forewing,

including tegula 16-18 mm. Head and body chiefly

or entirely black; metasomal terga feebly metallic

blue green, most evident on impunctate apical

marginal depressions; antennae black or brownish

black above, deep reddish brown or mahogany below; mandibles dark reddish brown basally, becoming deep mahogany red or black apically; clypeus

black laterally, paling to deep mahogany red on

medial surface; wings dissimilar in coloration, forewings nearly uniformly infuscated throughout with

black, somewhat darker apically, hind wings hyaline; wing veins dark reddish brown; tegulae dark

reddish brown, irregularly mottled with yellow;

legs chiefly dark reddish brown, tarsi usually redder; tibial spurs pale reddish brown; claws deep

mahogany red, banded medially with black.

Vestiture of head chiefly ochraceous to nearly

white on hind margins of head and face below

antennal sockets; pubescence on face above antennal sockets bright fulvo-ochraceous darkening to

brownish black on upper part of face and vertex;

thorax thickly clothed with mostly ochraceous to

fulvo-ochraceous pubescence, usually paling posteriorly to nearly white and sometimes darkening

on lateral and ventral surfaces (variably) to brownish black; legs with chiefly brownish black to fulvous pubescence basally, paling to bright fulvous

on tibiae and tarsi; pubescence of first metasomal

tergum consisting of long, erect whitish plumose

hairs, thickest at sides and on ventrally exposed

surfaces; metasomal terga II-IV, without apical

bands of pale pubescence, mostly clothed with

short, posteriorly directed, brownish black pubescence, longest and thickest at sides, sometimes terga

II and IV tufted with whitish pubescence at extreme ventrally exposed sides and usually basal

discal surface of tergum II with erect whitish or

cinereous pubescence; metasomal tergum V mostly

or entirely clothed with brownish black pubescence

(sometimes becoming whitish laterally), shortest

on punctate surface basally, much longer and forming a transverse band of long simple hairs arising

just before impunctate apical margin, projecting

posteriorly, and terminating in thick tufts of plumose hairs laterally; metasomal tergum VI with

numerous long, posteriorly directed, white plumose

hairs arising just before impunctate apical margin,

longest and somewhat tuftlike laterally and occasionally intermixed medially with brownish black

or black hairs; metasomal tergum VII with pubescence chiefly white and disposed laterally as posteriorly projecting tufts of plumose hairs; metasomal

sterna thickly clothed with rather long plumose

hairs, usually mostly or entirely white on sterna

I-III and brownish black on succeeding sterna,

which in some specimens may display variable

amounts of pale pubescence apically and laterally.
NUMBER 220 65

Eyes strongly convergent above, separated on

vertex by less than transverse diameter of median

ocellus (Figure 5); clypeus, except at sides, nearly

impunctate, at most with a few, well-separated

large punctures on depressed median surface; supraclypeal area densely punctured, without shining

interspaces; frontal carina triangular in outline,

very short, much shorter than diameter of median

ocellus, deeply excavated basally; middle tibiae and

basitarsi of equal length; punctation on discs of

metasomal terga I and II dense, rather fine with

punctures separated by mostly less than their diameters, those on terga III and IV similar, but less

densely punctate especially medially and laterally

over and basal to raised submarginal areas; fifth

metasomal tergum with discal punctures similar in

size and disposition to those on tergum II, but less

densely crowded especially medially; structure of

seventh metasomal tergum (Figure 65), seventh

and eighth metasomal sterna (Figures 66, 67), and

genitalia (Figure 68) as illustrated.

FEMALE.—Length 18-20 mm. Length of forewing, including tegula, 18-21 mm. Head and body

chiefly black, paling somewhat ventrally to very

dark mahogany; metasomal terga feebly metallic

blue green with some purplish tints evident in

bright light and most evident on impunctate apical

margins; antennae dark brownish black, somewhat

reddened at apex of scape and pedicel and to a

variable extent on flagellar segments; mandibles

chiefly deep mahogany red to black, usually near

bases and apices; clypeus black laterally, paling to

deep mahogany on medial surface; wings dissimilar

in coloration, forewings rather uniformly infuscated throughout and violaceous in bright light,

hind wings mainly hyaline except for some blackish

infuscation apically and posteriorly; wing veins

dark brownish black; tegulae dark mahogany, somewhat reddened externally; legs mostly blackish

brown dorsally, usually much reddened on tarsal

segments and below; tibial spurs deep reddish

brown; claws deep mahogany red, banded medially

with black.

Vestiture of head chiefly moderately long and

white on face, especially that about ocelli, antennal

sockets and clypeus at sides; hind margins of head

thickly clothed with long pubescence, mostly white

on lower genae, becoming intermixed with and

replaced by dark brownish and black pubescence

dorsally; thorax thickly clothed with dark brownish black pubescence at least dorsally, usually paling

irregularly at sides and below to dark brown; legs

mostly clothed with rather long brownish pubescence, sometimes paling to reddish brown on tarsal

segments; scopal hairs of hind legs mostly reddish

brown, sometimes with intermixture of brownish

black or black hairs; discal surfaces of metasomal

terga I-IV devoid of pubescence except for very

short, posteriorly directed, brownish black hairs

situated laterally, which at extreme sides and on

ventrally exposed surfaces are replaced on terga I

and II by scopae consisting of long, densely plumose, chiefly cinereous to whitish hairs and ventroapically on terga III and IV with long tufts of

chiefly whitish plumose hairs; metasomal tergum V

apically traversed by a thick fringe of long, posteriorly directed, plumose hairs characteristically

white at sides and brownish black on medial tergal

surface; metasomal tergum VI densely clothed on

either side of pygidial plate with moderately long,

brownish black plumose hairs; scopal hairs of metasomal sternum I brownish black; those on sternum

II and median tuft on sternum III largely or entirely white; other surfaces ofx

 metasomal sterna

II-V (except extreme sides) and VI thickly clothed

with rather short to moderately long posteriorly

directed, reddish brown to brownish black hairs;

extreme sides of metasomal sterna II-V with long

tufts of white plumose hairs projecting posteriorly

from under overlapping tergal surfaces.

Eyes weakly diverging below (49:55), clypeus,

except at sides, nearly impunctate, at most with a

few, well-separated large punctures on smoothed

and reddened median surface; supraclypeal area

densely and nearly confluently punctured except at

sides and above; frontal carina triangular in outline, short, distinctly less than diameter of median

ocellus, deeply excavated and polished; discal surfaces before marginal depressions of metasomal

terga I-V deeply punctured, well separated medially, crowded laterally, sparsest on tergum I medially, and closest and densest on tergum V.

GEOGRAPHIC RANGE (Map 3).—Southwestern

Mexico (Nayarit).

TYPES.—Holotype male, allotype female, and 9

paratypes ( $ , 89) were collected 4 miles south of

Acaponeta, Nayarit, Mexico, on 2 September 1962,

by D. H. Janzen. Additional paratypes (7 $ , 4 9)

are from the following localities in Nayarit: Acaponete, $, 29 September 1966 (G. E. and A. S.
66

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

66

FIGURES 65-68.—Mesoxaea clypeata Hurd and Linsley, holotype male (4 mi S Acaponeta, Nayarit,

Mexico): 65, seventh metasomal tergum (left half ventral); 66, 67, seventh and eighth metasomal

sterna (left halves dorsal); 68, genitalia (left half ventral).

Bohart, GEB); 8 miles southeast of Acaponete,

9, 19 October 1964 (A. E. Michelbacher, CIS);

Navarrete, 6 $ , 28 July 1953 (C. and P. Vaurie,

1953 David Rockefeller Mexican Expedition,

AMNH); 18 miles north of Tepic, 2 9 , 16 August

1960 (P. H. Arnaud, Jr., E. S. Ross, and D. C.

Rentz, CAS) and 4.5 miles southeast of Tuxpan,

9 , 9 September 1965, on flowers of Sesbania macrocarpa (R. D. Sage, CIS). The primary types are

deposited in the collections of the California Academy of Sciences, San Francisco.

DISCUSSION.—The sparsely punctate and shining

clypeus distinguishes Mesoxaea clypeata from all

its congeners. Structurally it is most closely related
NUMBER 220 67

to M. nigerrima, which is very similar in colorational features. At present this species is known

only from a few localities in the State of Nayarit,

Mexico, and has been collected at some localities

there with M. arizonica and M. rufescens. Nothing

is known about its nesting habits or its pollen

sources.

Literature Cited

Ashmead, William H.

1899. Classification of the Bees, or the Superfamily

Apoidea. Transactions of the American Entomological Society, 26:49-100.

Bertoni, A. de Winkelried

1911. Contribuci6n a la Biologfa de las Avispas y Abejas

del Paraguay (Hymenoptera). Anales Museo Nacional de Historia Natural de Buenos Aires, 15:97-

146. [1912.]

Birkmann, G.

1932. [Habits of Protoxaea texana.] Giddings News (Giddings, Texas), 2 September 1932: page 7, columns

6 and 7.

Bischoff, H.

1934. Gedanken zu einem natiirlichen System der Bienen

(Hym. Apid.). Deutsche Entomologische Zeitschrift,

1934:324-331, 3 figures.

Blanchard, fimile

1845. Histoire des Insectes, traitant de leurs Moeurs et

leurs Metamorphoses en gendral et comprenent une

Nouvelle Classification fondee sur leurs rapports

natureles, Hyminopteres et Coleopteres. v + 398

pages. Paris: Librairie de Firmin Didot Freres.

Bowers, Karen A. W.

1975. The Pollination Ecology of Solanum rostratum

(Solanaceae). American Journal of Botany, 62:633-

638, 2 figures, 4 tables.

Cameron, P.

1901. Description of a New Genus and Five New Species

of Aculeate Hymenoptera from the Santa Fe Mountains, New Mexico. Transactions of the American

Entomological Society, 27:311-316.

Cazier, M. A., and E. G. Linsley

1963. Territorial Behavior among Males of Protoxaea

gloriosa (Fox). Canadian Entomologist, 94:547-556,

2 figures.

1974. Foraging Behavior of Some Bees and Wasps at

Kallstroemia grandiflora Flowers in Southern Arizona and New Mexico. American Museum Novitales, 2546:1-20, 6 figures, 2 tables.

Claude-Joseph, F.

1926. Recherches biologique sur les Hymenopteres du

Chili. Annales des Sciences Naturelles, Zoologie

(Paris)(10) 9:113-268.

Cockerell, T. D. A.

1898. Tables for the Determination of New Mexico Bees.

Bulletin of the University of New Mexico, 1:41-73.

[Reprinted from Bulletin of the Science Laboratories of Denison University, vol. XI.]

1904. Some bees from San Miguel County, New Mexico.

The Entomologist, 37:5-9.

1906a. Descriptions and Records of Bees, XII. Annals and

Magazine of Natural History, series 7, 18:69-75.

1906b. The Bees of New Mexico. Transactions of the

American Entomological Society, 32:289-314.

1910. Descriptions and Records of Bees, XXXII. Annals

and Magazine of Natural History, series 8, 6:272-

284.

1912. The Stanford Expedition to Brazil. Psyche, 19:

41-61.

1917a. Some Records of Bees. Journal of the New York

Entomological Society, 25:135.

1917b. Descriptions and Records of Bees, LXXV. Annals

and Magazine of Natural History, series 8, 19:

473-481.

1918. A Remarkable New Bee of the Genus Oxaea

(Hym.). Entomological News, 29:252.

1920. Bees in the Collection of the United States National Museum, 3. Proceedings of the United States

National Museum, 55:167-221.

1926. Some Bees in the Collection of the California

Academy of Sciences. Pan-Pacific Entomologist, 3:

80-90.

1933. The Excessive Abundance of Certain Bees. American Naturalist, 67:286-288.

1934a. Records of American Bees. American Museum

Novitates, 697:1-15.

1934b. Two Interesting New Bees. Canadian Entomologist,

66:152-153.

1936. Records of Western Bees. American Museum

Novitates, 831:1-6.

Cockerell, T. D. A., and Wilmatte Porter

1899. Contributions from the New Mexico Biological

Station, VII: Observations on Bees, with Descriptions of New Genera and Species. Annals and

Magazine of Natural History, series 7, 4:403-421.

Crawford, J. C.

1915. Descriptions of New Hymenoptera, No. 9. Proceedof the United States National Museum, 48:577-

586, 11 figures.

Dalla Torre, C. G. de

1896. Catalogus Hymenopterorum. Volume 10, vii + 643

pages. Leipzig.

Ducke, A.

1901. Beobachtungen iiber Blutenbesuch Erocheinungszeit etc. der bei Para vorkommenden Bienen.

Zeitschrift fur Systematische Hymenopterologie und

Dipterologie, 1:25-32, 49-65.
68 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

1907. Contribution a la connaissance de la fauna Hymenopterologique du Nord-Est du Bresil. Revue

d'Entomologie (Caen), 26:75-96.

1908. Contributions a la connaissance des Hymenopteres

des deux AmeYiques. Revue d'Entomologie (Caen),

27:28-^5.

1910. Contribution a la Connaissance de la faune Hymenopterologique du Nord-Est du Bresil, III: Hymenopteres recoltes dans 1'fitat de Ceara en 1909 et

supplements aux deux listes anterieures. Revue

d'Entomologie (Caen), 28:78-122.

1912. Die natiirlichen Bienengenera Siidamerikas. Zoologische Jahrbucher Abteilung fur Systematik, Geographie und Biologie der Tiere (Jena), 34:51-116.

Evans, Howard E., and E. Gorton Linsley

1960. Notes on a Sleeping Aggregation of Solitary Bees

and Wasps. Bulletin of the Southern California

Academy of Sciences, 59:30-37, 1 plate.

Fox, William J.

1893. Synopsis of the N. American Species of Megacilissa.

Psyche, 6:421:422.

1894. Second Report on Some Hymenoptera from Lower

California, Mexico. Proceedings of the California

Academy of Sciences, series 2, 4:92-121.

Friese, H.

1898. Monographic der Bienengattungen Megacilissa,

Caupolicana, Diphaglossa, und Oxaea. Annalen des

K. K. Naturhistorischen Hofmuseums, 13:59-86.

1899. Monographic der Bienengattungen Megacilissa,

Caupolicana, und Oxaea. Annalen des K. K. Naturhistorischen Hofmuseums, 14:239-246.

1904. Zweiter Nachtrag zu den Bienengattungen Caupolicana, Ptiloglossa und Oxaea (Hym.). Zeitschrift

fur Systematische Hymenopterologie und Dipterologie, 4:17-20.

1912. Neue und wenig bekannte Bienenarten der neotropische Region. Archiv fiir Naturgeschichte,

Abtheilung A, 6:198-226, 16 figures.

Gerstaecker, A.

1867. Bericht iiber die wissenschaftlichen Leistungen im

Gebeite der Entomologie wahrend der Jahre 1865-

66, Zweite Halfte: Hymenoptera. Archiv fur Naturgeschichte, 33:305-345.

Graf, Vinalto

1966. A posicao systematica de Oxaeinae (Hymenoptera -

Apoidea). Ciencia e Cultura, 18:137-138.

Gribodo, Giovanni

1894. Note Imenotterologische, Nota II: Nuovi generi e

nuove specie di Imenotteri Antophili ed osservazioni sopra alcune specie gia conosciuti. Bullettino

della Societa Entomologica Italiana, 25:248-287.

Halsted, Byron D.

1890. Notes on Stamens of Solanaceae. Botanical Gazette,

15:103-106.

Harris, J. Arthur, and Oscar M. Kuchs

1902. Observations on the Pollination of Solanum rostratum Dunal and Cassia chamaecrista L. University

of Kansas Science Bulletin, 1:15-41, 1 plate.

Heinrich, Bernd, and Peter H. Raven

1972. Energetics and Pollination Ecology. Science, 176:

597-602.

Heiser, C. B. Jr., and T. W. Whitaker

1948. Chromosome Number, Polyploidy, and Growth

Habit in California Weeds. American Journal of

Botany, 35:179-186.

Hurd, Paul D., Jr., and E. Gorton Linsley

1975. The Principal Larrea Bees of the Southwestern

United States (Hymenoptera: Apoidea). Smithsonian Contributions to Zoology, 193:1-74, 18 figures, 15 tables.

Illiger, Karl

1806. William Kirby's Familien der Bienenartigen Insekten mit Ziisatzen, Nachweisungen und Bemerkungen. Magazin fiir Insekten (Braunschweig,

Reichard), 5:28-175.

Janvier, H.

1933. Etude biologique de quelques Hymenopteres du

Chile. Annales des Sciences Naturales Zoologie

(Paris), series 10, 16:209-356, 63 figures.

Joergensen, C. A.

1928. The Experimental Formation of Heteroploid Plants

in the Genus Solanum. Journal of Genetics, 19:

133-211.

Jorgensen, P.

1909. Beobachtungen iiber Blumenbesuch, Biologie verbreitung usw. der Bienen von Mendoza (Hym.).

Deutsche Entomologische Zeitschrift (Teil II):211-

227.

1912a. Los Crisfdidos y los Himen6pteros Aculeatos de la

Provincia de Mendoza. Anales del Museo Nacional

de Historia Nacional de Buenos Aires, series 3,

22(15):267-338. [1911.]

1912b. Revision der Apiden der Provinz Mendoza, Republica Argentina (Hym.). Zoologische Jahrbucher

Abteilung fiir Systematik, Geographic und Biologie

der Tiere (Jena), 32:89-163, 8 figures.

Klug, J. C. F.

1807. Oxaea, eine neue Gattung aus der Ordnung der

Piezaten Magazin der Gesellschaft Naturforschender Freunde zu Berlin, 1:261-263.

Laroca, Sebastiao

1970. Contribuicao para o Conhecimento das Relac,6es

entre Abelhas e Flores: Coleta de P61en das Anteras

Tubulares de certas Melastomataceae. Revista

Floresta, 2:69-74.

Latreille, P. A.

1809. Genera Crustaceorum et Insectorum secundum

ordinem naturalen in familias disposita, iconibus

exemplisque plurimis explicata. Volume 4, pages

1-399. Paris and Strasburg: Konig.

1810. Considerations generates, sur I'ordre naturel des

Animaux composant les classes des Crustacis des

Arachnides, et des Insectes; avec un Tableau

methodique de leurs genres, disposes en families.

viii + 444 pages. Paris: Schoell.
NUMBER 220 69

1811. Volume VIII [in part] of Olivier, Histoire Naturelle: Insectes. In Encyclopedic Methodique. . . .

1825. Families naturelles du Regne Animal, exposees

succinctement et dans un ordre analytique, avec

Vindication de leurs genres. 570 pages. Paris: J. B.

Bailliere, Baudouin Freres.

Lepeletier de Saint-Fargeau, M. le Compe Amedee Louis

Michel

1841. Histoire naturelle des insects: Hyme'nopteres.

[Suites a Buffon.] Volume 2, pages 1-680. Paris:

Roret.

Linsley, E. Gorton

1951. Oxaeini. In Muesebeck, Krombein, and Townes,

editors, Hymenoptera of America North of Mexico:

Synoptic Catalog. United States Department of

Agriculture Agricultural Monograph, 2:1086.

1962. The Colletid Ptiloglossa arizonensis Timberlake, a

Matinal Pollinator of Solanum. Pan-Pacific Entomologist, 38:75-82.

Linsley, E. Gorton, and Mont A. Cazier

1963. Further Observations on Bees Which Take Pollen

from Plants of the Genus Solanum. Pan-Pacific

Entomologist, 39:1-18, 6 figures, 2 tables.

1972. Diurnal and Seasonal Behavior Patterns among

Adults of Protoxaea gloriosa (Hymenoptera, Oxaeidae). American Museum Novitates, 2509:1-25, 4

tables.

Linsley, E. Gorton, and Charles D. Michener

1962. Brief Notes on the Habits of Protoxaea (Hymenoptera: Andrenidae). Journal of the Kansas Entomological Society, 35:385-389, figures.

Macior, Lazarus Walter

1971. Co-evolution of Plants and Animals—Systematic

Insights from Plant-Insect Interactions. Taxon,

20:17-28, figures 1-9.

1974. Behavioral Aspects of Coadaptions between Flowers

and Insect Pollinators. Annals of the Missouri

Botanical Gardens, 61:760-769, 21 figures.

Meehan, Thomas

1886. On the Fertilization of Cassia marilandica. Proceedings of the Academy of Natural Sciences of

Philadelphia, pages 314-318.

Meeuse, B. J. D.

1961. The Story of Pollination, vii + 243 pages, 1 plate,

83 figures. New York: The Ronald Press.

Michener, Charles D.

1944. Comparative External Morphology, Phylogeny, and

a Classification of the Bees (Hymenoptera). Bulletin of American Museum of Natural History, 82:

151-326, 246 text figures, 13 diagrams.

1954. Bees of Panama. Bulletin of the American Museum

of Natural History, 104:1-176, 155 figures, 3 tables.

1962. An Interesting Method of Pollen Collecting by

Bees from Flowers with Tubular Anthers. Revista

de Biologia Tropical, Universidad de Costa Rica,

10:167-175.

1965. A Classification of the Bees of the Australian and

South Pacific Regions. Bulletin of the American

Museum of Natural History, 130:1-362, 789 figures,

15 plates, 4 tables.

1966. The Classification of the North American Diphaglossinae and North American Species of the Genus

Caupolicana (Hymenoptera, Colletidae). University

of Kansas Science Bulletin, 46:717-751, 48 figures.

1974. The Social Behavior of the Bees, xii + 404 pages,

illustrated. Cambridge, Massachusetts: The Belknap

Press of Harvard University Press.

Michener, C. D., and R. D. Lange

1957. Observations on the Ethology of Some Brazilian

Colletid Bees. Journal of the Kansas Entomological

Society, 30:71-80.

Mitchell, Theodore B.

1960. Bees of the Eastern United States, volume 1.

North Carolina Agricultural Experiment Station,

Technical Bulletin, 141: 538 pages, 134 figures, 16

tables.

Moure, Jesus S.

1944. Apoidea de Colec.ao do Conde Amadeu A. Barbiellini (Hym. Apoidea). Revista de Entomologia, 15:

1-18.

1946. Notas sobre as Mamangabas. Boletim Agricola, 4:

3-32, 5 text figures, 3 plates.

1947. Notas sobre Albunas Abejas de la Provincia de

Salta. Revista de la Sociedad Entomoldgica Argentina, 13:218-253.

1950. Euherbstiinae nova sub-familia de Andrenidae

(Hymenopt.-Apoidea). Dusenia, 1:303-306.

Moure, Jesus S., and B. Lucas de Oliviera

1962. Novo g£nero de Panurginae para America do Sul

(Hymenoptera-Apoidea). Boletim da Universidade

do Parana, Zoologia, 15:1-14, 3 figures.

Moure, J. S., and C. A. C. Seabra

1962. A New Species of the Genus Oxaea from Brazil

(Hymenoptera: Apidae). Journal of the New York

Entomological Society, 70:235-238.

Moure, J. S., and D. Urban

1963. Uma Novo Esp£ce de "Oxaea" de Mato Grosso,

Brasil (Hymenoptera, Apoidea). Revista Brasilero

de Biologia, 23:361-364, 3 figures..

Miiller, Fritz

1883. Two Kinds of Stamens with Different Functions in

the Same Flower. Nature, 27:364-365.

Olivier, M.

1809. Volume V of Histoire Naturelle: Insectes. In Encyclopedic Methodique. . . .

Popov, V. V.

1941. Family Oxaeidae and Process of Morphological

Reduction in Bees (Hymenoptera, Apoidea).

Comptes Rendus (Doklady) de I'Academie des Sciences de I'URSS, new series, 30:82-85, 4 figures.

1945. On the Morphological Reduction in the Male

Genitalia of Bees (Hymenoptera, Apoidea). Zoologischeskii Zhurnal (Moscow), 9:895-949, 19 figures.

1951. Parasitic Bees of the Genus Ammobates Latr. (Hymenoptera, Anthophoridae). Trudy Zoologicheskoga

Instituta SSSR, 9:895-949, 19 figures. [In Russian.]
70

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

Rayment, Tarlton

1935. A Cluster of Bees. 752 pages, 65 plates, 100 figures,

frontispiece. Sidney: Endeavour Press.

Roberts, Raddyffe B.

1971. Biology of the Crepuscular Bee Ptiloglossa guinnae

n. sp. with Notes on Associated Bees, Mites, and

Yeasts, Journal of the Kansas Entomological Society,

44:283-294.

1973. Nest Architecture and Immature Stages of the Bee

Oxaea flavescens and the Status of Oxaeidae (Hymenoptera). Journal of the Kansas Entomological

Society, 46:437^146.

Robertson, Charles

1890. Flowers and Insects, V. Botanical Gazette, 15:199-

204.

Rozen, Jerome G., Jr.

1951. A Preliminary Comparative Study of the Male Genitalia of Andrenidae (Hymenoptera, Apoidea).

Journal of the Kansas Entomological Society, 24:

142-150, figures 1-17.

1964. Phylogenetic-taxonomic Significance of Last Instar

of Protoxaea gloriosa Fox, with Descriptions of

First and Last Instars (Hymenoptera: Apoidea).

Journal of the New York Entomological Society,

72:223-230, figures 1-7.

1965. The Biology and Immature Stages of Melitturga

clavicornis (Latreille) and of Sphecodes albilabris

(Kirby) and the Recognition of the Oxaeidae at

the Family Level (Hymenoptera, Apoidea). American Museum Novitates, 2224:1-18.

1966. The Larvae of Anthophoridae (Hymenoptera,

Apoidea), Part 2: The Nomadidae. American Museum Novitates, 2244:1-38.

Sandhouse, Grace A.

1943. The Type Species of the Genera and Subgenera of

Bees. Proceedings of the United States National

Museum, 92:519-619.

Schrottky, C.

1902a. Ensaio sobre as Abelhas Solitarias do Brasil. Revista do Museo Paulista, 5:330-613, plates 12-14.

1902b. Les especes de genres Megacilissa, Caupolicana,

Oxaea, Epicharis, Centris, Meliphila et Euglossa

dans la collection du Musee National de Buenos

Ayres. Anales del Museo Nacional de Historia Naturel de Buenos Aires, 7:317-327.

1903. Neue Brasilianische Hymenopteren. Zeitschrift fur

Systematische Hymenopterologie und Dipterologie,

3:39-45.

1904. Beitrag zur Kenntnis einiger siidamerikanischer Hymenopteren. Allgemeine Zeitschrift fiir Entomologie, 9:344-349

1905. Enumeration des Hymenopteres connus jusqu'ici

de la Republique Argentine, de l'Uruguay et du

Paraguay. Anales de la Sociedad Cientifica Argentina, 15:80-91, 118-124, 176-186.

1907. Contribuci6n al conocimiento de los Himenopteros

de Paraguay. Anales Cientificos Paraguayos, 7:1-78.

1908. Blumen und Insekten in Paraguay. Zeitschrift fiir

wissenschaftliche Insektenbiologie, 4:22-26, 47-52,

73-78.

1913. La Distribuci6n Geografica del los Himen6pteros

Argentinos. Anales de la Sociedad Cientifica Argentina, 75:114-115, 180-286.

Sichel, J.

1865. £tudes hymenopteiologiques: Essai d'une monographie du genre Oxaea Klug. Annales de la Sociite

Entomologique de France, 34:331-334, plate 9,

figures 1-3.

Smith, Frederick

1854. Apidae. Part II in Catalogue of Hymenopterous

Insects in the Collection of the British Museum.

Pages 199-465, plates vii-xii. London.

Snodgrass, Robert Evans

1941. The Male Gcnitalia of Hymenoptera. Smithsonian

Miscellaneous Collections, 99:1-85, 6 figures, 33

plates.

Stebbins, G. L., Jr., and E. F. Paddock

1949. The Solanum nigrum Complex in Pacific North

America. Madrono, 10:70-81.

Stephen, W. P., G. E. Bohart, and P. F. Torchio

1969. The Biology and External Morphology of Bees

with a Synopsis of the Genera of Northwestern

America. 140 pages. Corvallis: Agricultural Experiment Station, Oregon State University.

Stow, I.

1927. A Cytological Study on Pollen Sterility in Solanum

tuberosum L. Japanese Journal of Botany, 3:217-

237.

Strand, Embrik

1910. Beitrage zur Kenntnis der Hymenopterenfauna

von Paraguay auf Grund der Sammlungen und

Beobachtungen von Prof. J. D. Anisits. Zoologische

Jahrbucher Abteilung fiir Systematik, Geographie

und Biologie der Tiere, 29:455-562.

Thorp, Robbin, and James R. Estes

1975. Intrafloral Behavior of Bees on Flowers of Cassia

fasciculata. Journal of the Kansas Entomological

Society, 48:175-184, 6 figures.

Todd, J. E.

1882. On the Flowers of Solanum rostratum and Cassia

chamaecrista. American Naturalist, 16:281-287.

Truxal, F. S.

1962. Observations on the Nests of Oxaea (Hymenoptera: Andrenidae). Journal of the Kansas Entomological Society, 35:432.

Wille, Alvaro

1963. Behavioral Adaptations of Bees for Pollen Collecting from Cassia Flowers. Revista de Biologia Tropical (Universidad de Costa Rica), 11:205-210.

Yager, Karen, and Jerome G. Rozen, Jr.

1966. Preliminary Systematic Study of the Pupae of

Andrenid Bees (Hymenoptera, Apoidea). American

Museum Novitates, 2265:1-13.
Plates 1-3

NUMBER 220 73

PLATE 1.—Protoxaea gloriosa (Fox): male (above) hovering in territorial flight; female (below)

in flower of Kallstroemia grandiflora (Torrey) Gray.
74 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

PLATC 2.—"Sleeping" aggregation of males of Protoxaea gloriosa (Fox).
NUMBER 220 75

PLATE 3.—Closeup details of "sleeping" aggregation of males.

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