Studies of Halictinae

(Apoidea: Halictidae),

II: Revision of

Sphecodogastra Ashmead,

Floral Specialists of Onagraceae

RONALD J. MCGINLEY

i

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY • NUMBER 610
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SMITHSONIA N CONTRIBUTION S T O ZOOLOG Y • NUMBE R 61 0

Studies of Halictinae (Apoidea: Halictidae),

II: Revision of Sphecodogastra Ashmead,

Floral Specialists of Onagraceae

Ronald J. McGinley

Smithsonian Institution Press

Washington, D.C.

2003
ABSTRAC T

McGinley, Ronald J. Studies of Halictinae (Apoidea: Halictidae), II: Revision of Sphecodogastra Ashmead, Floral Specialists of Onagraceae. Smithsonian Contributions to Zoology,

number 610, 55 pages, 151 figures, 3 tables, 2003.—The bee genus Sphecodogastra Ashmead

is revised from the study of approximately 3200 specimens. Eight species are recognized with

three described as new. All species are described and diagnosed. The systematic history of the

genus, including questions concerning its monophyly, is presented. Flight records are summarized with histogram plots, distributions are indicated by dot maps, and illustrated keys are provided for species identification. Floral association data and daily flight activity records are

summarized, and a literature review of nesting biology is presented for four species (5. antiochensis, S. lusoria, S. oenotherae, S. texana). The new species are S. antiochensis (a potentially

endangered species), S. danforthi, and S. potosi. Sphecodogastra lusoria (Cresson) is elevated

from junior synonymy under S. aberrans (Crawford), and Halictus galpinisae Cockerell is synonymized under 5. lusoria. New combinations are S. aberrans (Crawford), 5. lusoria

(Cresson), and S. oenotherae (Stevens).

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

recorded in the Institution's annual report, Annals of the Smithsonian Institution. SERIES COVER

DESIGN: The coral Montastrea cavernosa (Linnaeus).

Library of Congress Cataloging-in-Publication Data

McGinley, Ronald J.

Studies of Halictinae (Apoidea: Halictidae), II: revision of Sphecodogastra Ashmead, floral specialists of

Onagraceae / Ronald J. McGinley.

p. cm. — (Smithsonian contributions to zoology ; no. 610)

Includes bibliographical references (p.).

1. Sphecodogastra—Classification. I. Title. II. Series.

QLI S54no. 610

[QL568.H3]

590 s—dc21

[599.79'9]

2001020072

© The paper used in this publication meets the minimum requirements of the American

National Standard for Permanence of Paper for Printed Library Materials Z39.48—1984.
Contents

Page

Introduction 1

Material and Methods 1

Acknowledgments 1

Sphecodogastra Systematic History 3

Monophyly 3

Species Descriptions and Synonymies 6

Flight Activity 7

General Flight Records 7

Daily Flight Activity Records 7

Floral Associations 9

Onagraceae Floral Associations 9

Significance of Sphecodogastra in Onagraceae Reproductive Biology 10

Non-Onagraceae Floral Associations 11

Nesting Biology 12

Social Behavior 12

Nest Architecture 12

Nest Architecture and Social Behavior 13

Nest Architecture and Parasites 13

Systematics 14

Sphecodogastra Ashmead 14

Key to Sphecodogastra Females 16

Key to Sphecodogastra Males 19

1. Sphecodogastra aberrans (Crawford), new combination 23

2. Sphecodogastra antiochensis, new species 28

3. Sphecodogastra danforthi, new species 31

4. Sphecodogastra lusoria (Cresson), new combination 34

5. Sphecodogastra noctivaga (Linsley and MacSwain) 38

6. Sphecodogastra oenotherae (Stevens), new combination 40

7. Sphecodogastrapotosi, new species 45

8. Sphecodogastra texana (Cresson) 47

Appendix 1: Identification of Figures in Keys 52

Appendix 2: Onagraceae Taxonomy and Synonymy 53

Literature Cited 54

111

Studies of Halictinae (Apoidea: Halictidae),

II: Revision of Sphecodogastra Ashmead,

Floral Specialists of Onagraceae

Ronald J. McGinley

Introduction

The eight known species of Sphecodogastra Ashmead are

widely distributed throughout North America, ranging from

southern Canada to Veracruz, Mexico (Figure 1). Females of

this halictine genus collect pollen almost exclusively from

evening primroses (Onagraceae). This specialized floral relationship and associated behavioral adaptations make Sphecodogastra of interest to bee biologists and pollination ecologists. The scopae (pollen collecting hairs on the hind legs of

females) are reduced to a single, linear row of simple and apically curved hairs on the hind femora (Figures 2, 3). These relatively thick hairs apparently facilitate collection of the unusual pollen of their host plants (Thorp, 1979). The large pollen

grains of Onagraceae are enveloped in what botanists refer to

as viscin threads. These threads and associated pollen are

"hooked" by the modified scopal hairs of these bees. As a result, females can accumulate extremely large pollen loads (Figure 4).

The dependence of Sphecodogastra on Onagraceae pollen

affects their diurnal flight activity. Most species are both matinal and crepuscular, and two species (S. noctivaga, S. texana)

are strictly crepuscular and nocturnal. The foraging activities of

Sphecodogastra texana are correlated with lunar cycles, and

Ronald J. McGinley, Office of the Chief, Illinois Natural History Survey, Champaign, Illinois 61820, USA [formerly of the Department of

Entomology, National Museum of Natural History, Smithsonian Institution, Washington, DC 20560-0105, USA].

Review Chairman: John M. Burns, Department of Systematic Biology,

National Museum of Natural History, Smithsonian Institution, Washington, DC. 20560-0127, USA.

Reviewers: Robert W. Brooks, The University of Kansas, Natural History Museum and Biodiversity Research Center, Lawrence, Kansas

66045, USA; Bryan N. Danforth, Department of Entomology, Cornell

University, Ithaca, New York 14853, USA; Laurence Packer, Department of Biology, York University, North York, Ontario M3J1P3. Canada.

TABLE 1.—Checklist of Sphecodogastra and numbers of specimens examined.

Species

S. aberrans (Crawford)

S. antiochensis, new species

S. danforthi, new species

S. lusoria (Cresson)

S. noctivaga (Linsley and MacSwain)

& oenotherae (Stevens)

S. potosi, new species

S. texana (Cresson)

Total

Total

426

574

39

918

225

170

32

903

3287

Females

327

233

34

633

201

104

18

847

2397

Males

99

341

5

285

24

66

14

56

890

these bees apparently use their conspicuously enlarged ocelli

(Figure 11) to navigate by moonlight (Kerfoot, 1967c).

The classification of Sphecodogastra has been fraught with

questions concerning generic monophyly and with long-standing confusion surrounding the identity of two of the most common and widespread species, S. aberrans and S. lusoria. I hope

this review will encourage additional systematic and biological

work on this interesting group of bees.

MATERIAL AND METHODS.—This study was based on approximately 3200 specimens borrowed from 36 institutions. A summary of specimens examined of each species is presented in Table 1. Methods and terminology follow Harris (1979) and

McGinley (1986). Following common convention, the terms

"tergum" and "sternum" are abbreviated "T" and "S, " respectively, in the "Systematics" and figure legends. Terminology associated with flight activity follows that of The Torre-Bueno

Glossary of Entomology (Nichols, 1989:179,423): crepuscular =

"active or flying at dusk," and matinal = "of or in the morning, as

in reference to activity patterns of insects." Flight data were recorded from labels for the specimens examined in this study.

ACKNOWLEDGMENTS.—I am grateful to the curators listed

below who arranged the loan specimens for this study. The institution acronyms used follow those of Griffiths (1980) with

minor modifications.
SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

FIGURE 1.—Sphecodogastra distribution in North America (circled dots represent literature localities for S.

oenotherae).

AMNH American Museum of Natural History, New York (J.G

Rozen, Jr.)

ANSP Academy of Natural Sciences, Philadelphia (D. Otte, D.

Azuma)

ASU Arizona State University, Tempe (F.F. Hasbrouck)

BYU Brigham Young University, Provo, Utah (S.L. Wood)

CAS California Academy of Sciences, San Francisco (W.J.

Pulawski)

CDA California Department of Agriculture, Sacramento (M.S.

Wasbauer)

CMP Carnegie Museum, Pittsburgh, Pennsylvania (G Ekis)

CNC Canadian National Collection, Ottawa (L. Masner, M.

Sharkey)

CSUFC Colorado State University, Fort Collins (H.E. Evans)

CTMI Central Texas Melittological Institute (J.L. Neff)

CU Cornell University, Ithaca, New York (GC. Eickwort)

FSCA Florida State Collection of Arthropods, Gainesville (L.A.

Stange)

INHS Illinois Natural History Survey, Champaign (W.E. LaBerge)

KSU Kansas State University, Manhattan (H.D. Blocker)

KU University of Kansas, Lawrence (R.W. Brooks, CD.

Michener)

LACM Los Angeles County Museum of Natural History, California

(R.R. Snelling)

MCPM Milwaukee City Public Museum, Wisconsin (GR. Noonan)

MCZ Museum of Comparative Zoology, Harvard University,

Cambridge, Massachusetts (M.K. Thayer)

NCSU North Carolina State University, Raleigh (C. Parron)

NMNH National Museum of Natural History, Smithsonian Institution, Washington, D.C. (P.D. Hurd, Jr., S. Shanks)

NMW Naturhistorisches Museum, Wien (Vienna), Austria (M.

Fischer)

NYSM New York State Museum, Albany (T.L. McCabe)

OhS Ohio State University, Columbus (C.A. Triplehom)

OrS Oregon State University, Corvallis (W.P. Stephen, GM.

Stonedahl)

TOR University of Toronto (G Knerer, L. Packer)

UAE University of Alberta, Edmonton (GE. Ball, D. Shpeley)

UCB University of California, Berkeley (H.V. Daly)

UColB University of Colorado, Boulder (U.N. Lanham)

UCR University of California, Riverside (S.I. Frommer)

UGa University of Georgia, Athens (C.L. Smith)

UIM University of Idaho, Moscow (J.B. Johnson)

UMSP University of Minnesota, St. Paul (P.J. Clausen)

UND University of North Dakota, Grand Forks (P.B. Kannowski)

UNL University of Nebraska, Lincoln (B.C. Ratcliffe)

USU Utah State University, Logan (T.L. Griswold)

UWL University of Wyoming, Laramie (R.J. Lavigne)

UWM University of Wisconsin, Madison (S. Krauth)
NUMBER 610

FIGURES 2, 3.—Sphecodogastra scopae, anterior views: 2, female S. texana; 3,

closeup of S. oenotherae scopal hairs.

Many individuals contributed to the production of this monograph, and I would like to acknowledge their help. Elaine R.S.

Hodges drew the male terminalia, Vichai Malikul took the habitus and head photographs, and George Venable was responsible

for scanning these images and the SEM photos, for graphic enhancement, and for plate layout. Preliminary distribution maps

were generated by Daniel Cole using Arc/Info software and

were then enhanced for publication by George Venable in

Adobe Illustrator. Maureen Mello handled the laborious return

of borrowed specimens to 36 museums.

I am greatly indebted to my apoid colleagues Bryan Danforth

(CU) and John Neff (CTMI) who provided critical specimens

of the rarely collected new species Sphecodogastra danforthi.

Warren Wagner (NMNH) contributed invaluable literature and

advice on Onagraceae systematics and distribution. Michael

Caterino (UCB) and Jerry Powell (UCB) contributed important

information on the Antioch Dunes habitat, and Jerry Powell

provided me with unpublished collection records for S. antiochensis.

Sphecodogastra Systematic History

The monophyly and generic limits of Sphecodogastra and

the identity of two of its most commonly collected species, 5.

aberrans and S. lusoria, have been subject to question and controversy for 131 years. Sphecodogastra texana was originally

described by Cresson (1872) as a parasitic Sphecodes species,

due to its orange abdomen and reduced scopa. In the same paper, Cresson described Halictus lusorius, a bee with a similarly

reduced scopa but a dark abdomen. The monophyly of the Texana (orange species: S. noctivaga, S. texana. Figures 4, 5) and

Lusoria groups (dark species, Figures 6, 7) and the problems

associated with species identity within the Lusoria Group, will

be explored below.

MONOPHYLY.—The following summary of systematic

events relating to Sphecodogastra higher classification provides a chronological framework for the discussion that follows.

1872 Cresson described Sphecodes texana (mistaking this for

a parasitic bee).

1898 Cockerell recognized Sphecodes texana as Halictus

{Parasphecodes) texanus.

1899 Ashmead recognized Sphecodogastra. Although he incorrectly indicated "Parasphecodes" texana as the type

species, he correctly noted that P. texana is not related

to Parasphecodes.

1951 Michener recognized the monophyly of Sphecodogastra sensu lato as a subgenus of Lasioglossum, including

5. texana and the dark-abdomened species originally

described as Halictus.

1958 Linsley accepted Michener's inclusive concept of Sphecodogastra but cautioned that Hurd (in litt.) indicated

[incorrectly] that several species of Evylaeus have modified scopae similar to Sphecodogastra.

1960 Mitchell accepted Michener's inclusive concept of

Sphecodogastra.

1962 Linsley and MacSwain, based on Hurd's comments (in

litt.), resurrected the restricted concept of Sphecodogastra and recommended transfer of other species to the

subgenus Evylaeus.

1979 Hurd formalized the restricted concept of Sphecodogastra in his Hymenoptera catalog (restated by Moure and

Hurd, 1987).

1994 Michener et al. restored Michener's (1951) inclusive

concept of Sphecodogastra.

Sphecodogastra was established as a monobasic genus by

Ashmead (1899) for Sphecodes texana Cresson (1872), a relatively large halictid having an orange abdomen, conspicuously

enlarged ocelli, and highly modified scopae, apparently

adapted for collecting pollen of Onagraceae. A related species, Sphecodogastra noctivaga, was later described by Linsley

and MacSwain (1962). Other species (the Lusoria Group),

herein included in Sphecodogastra, have similarly modified

scopae, but being relatively small and darkly pigmented they

were originally grouped with the other small, dark species of

Halictus, and later, with Evylaeus. Recognizing the monophyly

of the then known species in question (//. aberrans Crawford,

H. lusorius Cresson, H. oenotherae Stevens, H. texanus
SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

\

FIGURES 4-7.—Figures 4, 5: Sphecodogastra lusoria: 4, female with full load of pollen from Calylophus

hartwegii (length, 8.4 mm); 5, male (length, 7.2 mm). Figures 6,7: Sphecodogastra noctivaga: 6, female (length,

10.2 mm); 7, male (length, 8.8 mm).

(Cresson)), Michener (1951) placed all of them in Lasioglossum, subgenus Sphecodogastra. Since that time the monophyly

of this taxon has been disputed, largely because of classificatory changes made on the basis of literature review, not specimen examination. Linsley (1958:567) originally accepted

Michener's inclusive concept of Sphecodogastra monophyly

but noted that "the group is closely related to the subgenus Evylaeus, which contains several species which gather pollen from

diumal Onagraceae and thus have the scopa similarly modified

but have normal ocelli (Hurd, in litt.)." Hurd's statement that

several Evylaeus species not included in Michener's Sphecodogastra have Sphecodogastra-hke characters was incorrect.

Nevertheless, it highly influenced Linsley and MacSwain

(1962:45):

Although Ashmead established Sphecodogastra as a monobasic genus, several

other species have been subsequently added which share with S. texana one or

more adaptive features associated with the collection of pollen from Onagraceae, including Halictus abberrans [sic] Crawford, Halictus galpinsiae Cockerell, Halictus lusorius Cresson, and Halictus (Evylaeus) oenotherae Stevens.

However, these added species are crepuscular or matinal bees, or both, not truly nocturnal, and as pointed out by Hurd (see Linsley 1958), they are closely

related to species included in Lasioglossum, subgenus Evylaeus by Michener

(1951). We prefer to see them assigned to that group, since they agree with

Evylaeus in basic characters and in general facies. They also share with species

in Evylaeus and most of the other subgenera of Lasioglossum a distinct lateral
NUMBER 610

carina on the propodeum, which is lacking in the subgenus Sphecodogastra as

here restricted.

This viewpoint was formalized by Hurd (1979) and Moure

and Hurd (1987) who restricted the concept of Sphecodogastra

to include only S. noctivaga and S. texana and transferred the

smaller, darkly pigmented species to Evylaeus.

Aside from the misconception that several additional species

of Evylaeus share Sphecodogastra-like characters, taxa should

not be grouped on the basis of "basic characters" and "general

fades" unless these similarities can be reasonably hypothesized or shown to be uniquely derived (i.e., synapomorphies).

That the species of the Lusoria Group share the distinct lateral

propodeal carina with most other Lasioglossum subgenera

would suggest that the well-developed propodeal carina may

be plesiomorphic and, therefore, of no value in estimating species relationships (to date, this character-state pattern remains

unexplored, and polarities should be considered unknown).

There is little doubt that S. noctivaga and S. texana are sister

species (details presented herein) and could logically be recognized at the generic, subgeneric, or informal species-group

level; however, it appears these two species (the Texana Group)

form a more inclusive monophyletic taxon with the Lusoria

Group, which is recognized herein as Sphecodogastra, consistent with Michener's (1951) subgenus Sphecodogastra. The

apparent synapomorphies supporting this grouping are as follows: (1) females have the femoral scopa reduced to a near linear row of stiff, apically recurved setae (Figures 2,3; more diffused in S. antiochensis, Figure 25); (2) females are oligoleges

of Onagraceae (a biological character correlated with the scopal modification); and (3) female foraging activity is matinal,

crepuscular, and/or nocturnal (a behavioral modification also

associated with phenology in Onagraceae). Ocellar size, also

associated with foraging time, may be of interest for further

study. Ocelli of S. noctivaga and S. texana are conspicuously

large (the ratio of the lateral ocellar diameter to the ocellar-compound eye distance ranges from 0.33 to 0.35 in these

species). Although not as large, ocelli in most other Sphecodogastra appear to be somewhat larger than in most Evylaeus

species. The above ratio ranges from 1.18 to 1.44 in S. aberrans, S. antiochensis, S. lusoria, and S. danforthi and ranges

from 1.78 to 2.00 in S. oenotherae and S. potosi. A preliminary

survey of Evylaeus species for this character (E. cinctipes (Provancher), E. nelumbonis (Robertson), E. pectinatus (Smith), E.

pectoralis (Smith), and E. truncatus (Robertson)) showed a ratio range of 1.90-2.25.

Male Sphecodogastra have monotonously similar genitalia

with the exception of those of S. aberrans and 5. texana. Unlike other species, S. texana has the membranous retrorse lobes

very broad and twisted ventrally (Figures 31, 32). It is notable

that these lobes in the closely related 5. noctivaga are quite different (Figure 33) and are similar to those of the Lusoria Group

(compare with S. antiochensis, Figure 78). The genitalia of S.

aberrans differ significantly from those of other species in the

Lusoria Group only in having the penis valves angulate and

concave dorsally (Figure 67). It should be emphasized that a

comprehensive review of New World Evylaeus male genitalia

remains to be documented, and so the cladistic significance of

Sphecodogastra genitalic similarities remains to be determined; nevertheless, these similarites lend support for Lusoria

plus Texana group monophyly. That the genitalia of S. noctivaga are so similar to those of the Lusoria Group is compelling

evidence for monophyly.

Male Sphecodogastra also have conspicuous, elongate sternal vestiture (Figures 37, 38), and the male sternum 6 has a

central, shallowly depressed area with an elevated longitudinal,

subapical, rounded median ridge (Figures 66, 99). These characters, although characteristic of the males of this genus,

should be carefully studied in other halictine taxa before they

can be considered significant cladistic characters.

The most compelling evidence for the monophyly of Sphecodogastra, in the inclusive sense, is in the results of Bryan

Danforth's (2002) molecular systematic studies of Halictinae.

He analyzed a combined nuclear and mitochondrial data set for

54 species of Lasioglossum sensu lato (including 31 species of

Evylaeus) and found the monophyly of S. noctivaga and S.

oenotherae to be strongly supported by a 100% bootstrap

value.

In summary, characters that would logically group any Sphecodogastra (sensu lato) species with other halictine taxa are

unknown at present. The alternative position is to hypothesize,

a priori, that the scopal modifications found in the Lusoria and

Texana groups are the result of convergence associated with

their use of Onagraceae pollen. As Thorp (1979:792) pointed

out, such convergence is known to occur: "The scopal hairs of

most oligolectic (pollen specialist) bees that collect pollen from

the Onagraceae tend to be long and simple in contrast to those

of their relatives." This is known in Andrena (Onagrandrena;

most species of Diandrena), Melissodes clarkiae LaBerge, and

Diadasia angusticeps Timberlake. However, the simple scopal

hairs of Sphecodogastra are highly unusual in being extremely

linear in their arrangement (except in 5. antiochensis) and in

having hooked apices (Figures 2, 3). Lacking evidence to the

contrary, I consider the female scopa, suggestive male characters, and preliminary molecular data sufficient to recognize the

Lusoria and Texana groups as being monophyletic (i.e., Sphecodogastra).

Sphecodogastra has been alternately recognized at the generic level (Hurd, 1979; Moure and Hurd, 1987) or as a subgenus of Lasioglossum (Michener et al., 1994; Michener, 2000).

In this study, recognition of Sphecodogastra at the generic

level is done merely to correspond with my generic treatment

of Lasioglossum (McGinley, 1986). Decisions on the most useful categorical levels should await a better understanding of the

relative rankings involved. The monophyly of many currently

recognized halictine taxa is open to question, and a number of

these taxa (e.g., Dialictus, Evylaeus) are possibly artificial as

presently defined (Michener, 1974; McGinley, 1986).
SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

SPECIES DESCRIPTIONS AND SYNONYMIES.—The following

summary of systematic events relating to Sphecodogastra species descriptions and associated history provides a chronological framework for the discussion that follows.

1872 Cresson described Halictus lusorius and Sphecodes texana.

1897 Cockerell recorded one female of//, lusorius from New

Mexico. (This is the last use of the name lusorius in the

literature other than Crawford's 1907 note that he did

not include lusorius in his key to Halictus species.)

1903 Crawford described H. aberrans but did not compare it

with lusorius.

1903 Cockerell described H. galpinsiae but did not compare

it with lusorius, which he apparently had forgotten

about {galpinsiae is herein considered a junior synonym of Sphecodogastra lusoria).

1904 Vachal described H. gelidus (from a male specimen),

later synonymized under galpinsiae by Crawford

(1907).

1907 Cockerell maintained the distinctiveness of galpinsiae

and aberrans.

1911 Graenicher contrasted galpinsiae with aberrans, supporting their separate identities.

1920 Stevens described H. oenotherae.

1920 Stevens synonymized galpinsiae under aberrans.

1951 Michener listed galpinsiae as a junior synonym of aberrans.

1952 Timberlake (in litt., Moure and Hurd, 1987) presented

an inconclusive discussion of the identities of aberrans,

galpinsiae, and an undescribed "Antioch species.

1962 Linsley and MacSwain described Lasioglossum {Sphecodogastra) noctivaga.

1976 Bohart and Youssef differentiated galpinsiae from aberrans and noted the existence of an undescribed species from Antioch, California.

1979 Hurd listed galpinsiae as a junior synonym of aberrans

(agreeing with Stevens).

1987 Moure and Hurd listed galpinsiae as a junior synonym

of aberrans and offered additional justification.

Present paper McGinley synonymizes H. galpinsiae under

Sphecodogastra lusoria; recognizes the distinctiveness

of 5. aberrans and S. lusoria (agreeing with Bohart and

Youssef, Cockerell, Crawford, and Graenicher); and describes Sphecodogastra antiochensis, S. danforthi, and

S. potosi.

The confusion surrounding the identities of Sphecodogastra

aberrans and S. lusoria {— H. galpinsiae) was reviewed by

Moure and Hurd (1987:68):

Stevens (1920) has discussed the identity of this species [S. aberrans] and concluded that galpinsiae is a synonym of aberrans. Bohart and Youssef (1976)

consider these to be distinct, citing ecological criteria, and comment that a species closely related to galpinsiae from Antioch, California is an undescribed

species. Years ago, Timberlake (in litt.) looked into the identity of the Antioch

species and wrote the following on 10 July 1952: ["]The species of Halictus

that you sent from Antioch is, I think, H. galpinsiae Ckll. The males agree

closely with Colorado males that I collected in June, 1939. The fact that I had

determined females from Antioch previously as H. aberrans Cwfd. might indicate that the two names refer to one species, but against this is the fact that

Crawford considered the two distinct.f"] Cockerell [1906b:427] reports "Mr.

Crawford writes that H. galpinsiae and aberrans, so similar in the female, are

widely different in the male." In another place Cockerell states that aberrans is

a diurnal species. The male of aberrans which I presume is correctly placed

has slightly smaller ocelli, much less white tomentose hair on head and thorax,

abdomen less banded and the genitalia very distinctive. Timberlake further

comments that "Stevens' bee from North Dakota may or may not have been the

true aberrans, although presumably it was, as some of his material was determined by Crawford (but these specimens were diurnal ones from Helianthus)."

Until a critical systematic study is made of the nominal forms throughout the

range in North America, we have elected to consider that only one species [S.

aberrans] is represented in this area.

Much of the above confusion might have been avoided had

Cockerell recalled that he was aware of Cresson's (1872) description of Halictus lusorius. Cockerell (1897:167) identified

one female from New Mexico as Halictus lusorius and wrote,

"it is possible that the specimen represents a species distinct

from lusorius, but Cresson's description so nearly fits it that the

identity may be assumed until the contrary is proven." As soon

as 1901, Cockerell referred to specimens of this same species

as Halictus amicus, "var. a," and later in 1903 he described additional specimens as Halictus galpinsiae. Cockerell contrasted

H. galpinsiae with H. amicus, which is an entirely different

Evylaeus species, but did not mention H. lusorius, a species he

had previously recognized. Other than the problem of synonymy, the identity of//, galpinsiae and H. aberrans should have

been resolved in 1906 with the above quotation by Cockerell in

which he reported Crawford's note on the similarity of the females but extreme difference of males; Crawford also noted the

distinctive genitalia of//, aberrans. In his key to North American Halictus species, however, Crawford (1907) unfortunately

chose to differentiate the males on the basis of pubescence

characters (correct, but subtle without illustrations) with no

mention of genitalic characters, and he separated the females

by differences in first metasomal tergal and mesoscutal punctation (virtually useless without definition or illustration, both of

which were lacking in Crawford's treatment). Cockerell

(1907b) also differentiated H. aberrans and H. galpinsiae on

the basis of color differences of the wing stigma—another difficult character to use with unidentified material. By this time,

the synonymy question concerning H. lusorius and H. galpinsiae could have been settled, but Crawford, who mentioned

that he knew of Cresson's H. lusorius, excluded it from his key

(probably for lack of specimens).

As noted by Moure and Hurd (1987), Stevens (1920:37) was

the first worker to critically address the identity problem relating to H. aberrans and H. galpinsiae. He mentioned "2 females

and 9 males taken at Dickinson [North Dakota], by Mr. C.H.

Waldron [that] were determined by Mr. Crawford as H. aberrans." Two of these males are in the collection of the National

Museum of Natural History (Smithsonian Institution) and are

indeed aberrans, as determined by Crawford. Unfortunately,
NUMBER 610

800 -,

r T i r i ~~i r i i i i

Jan Feb Mar Apr May Jun Jul Aug Sep Oct Nov Dec

FIGURE 8.—Sphecodogastra flight records, by month of collection.

Stevens went on to say that these specimens "agree well with

Cockerell's description of galpinsiae [and that] a paratype female of galpinsiae sent by Prof. Cockerell agrees with the

North Dakota specimens." Although Stevens was mistaken on

both counts, his opinion apparently had a major influence on

Moure and Hurd's (1987) decision to recognize the synonymy

of aberrans and galpinsiae. Timberlake's cautious remarks (in

litt., Moure and Hurd, 1987) reflected the fact that the females

of S. antiochensis are similar to those of aberrans whereas the

males of S. antiochensis resemble those of galpinsiae. As recently as 1976, Bohart and Youssef emphasized the distinctiveness of aberrans and galpinsiae and, contrary to the above

quote by Moure and Hurd (1987), supported this on the basis of

morphological characters: "It [galpinsiae] differs from E. aberrans Crawford (the only other species in the Intermountain Region of the USA) in being smaller (average 8 mm in length)

and having more numerous minute appressed pale hairs on the

thorax" (Bohart and Youssef, 1976:185).

I herein synonymize H. galpinsiae under Sphecodogastra lusoria, describe as a new species the Antioch Sphecodogastra

long known to Bohart and Timberlake (S. antiochensis), and

recognize the distinctiveness of S. aberrans and S. lusoria

(agreeing with Bohart and Youssef, Cockerell, Crawford, and

Graenicher). Diagnoses for all species are provided in the species treatments below.

Flight Activity

GENERAL FLIGHT RECORDS.—Sphecodogastra specimens

examined in this study were collected from March through December (Figure 8). Most females (82%) were collected during

May through July and most males (79%) were collected

slightly later, in June and July. Given the lack of standardization in collecting methods and the broad geographic ranges

summarized, little should be concluded from these data. Variations in yearly patterns as well as subtleties of generational patterns cannot be determined from these data; for example, Bohart and Youssef (1976) reported S. lusoria to usually have in

most years two full generations and a partial third at Cornish,

Utah. The only valid comments might be that Sphecodogastra

species have been collected primarily during the late spring and

summer months, males emerge later than females, and that unlike other species, 5. antiochensis and S. oenotherae have been

primarily collected in late spring (Figures 82, 128). However,

in the northern area of its distribution in Ontario, Canada,

Knerer and MacKay (1969) observed S. oenotherae to be most

active in June and July. Data for individual species are presented in the species accounts that follow.

DAILY FLIGHT ACTIVITY RECORDS.—Most species of the

Lusoria Group (S. aberrans, S. antiochensis, S. lusoria, S.

oenotherae) have long been known to be both crepuscular and

matinal in flight activity (Stevens, 1920; Linsley et al., 1963,

1973; Turner, 1966; Knerer and MacKay, 1969; Bohart and

Youssef, 1976). I am not aware of time data for the two other

rarely collected species of this group, S. danforthi and 5. potosi, but one might assume they have activity patterns similar

to the above species, which would, in conjunction with their

known distributions, explain their relative lack of representation in museum collections.

The two species of the Lusoria Group with the most time

data associated with known specimens, S. aberrans (236

records) and S. lusoria (230 records), have flight times equally
SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

Time of Day

FIGURE 9.—Flight-time records, by hour of day, for Sphecodogastra aberrans (n = 236) and S. lusoria (n = 230)

(data from specimen labels examined in this study).

distributed between the early morning and the late afternoon to

early evening hours (Figure 9). For S. lusoria at Cornish, Utah,

Bohart and Youssef (1976) reported the earliest recorded inception of foraging during its morning activity period was 0558

hours and the latest was 0810 hours. Flights started after dawn

but usually before the sun directly illuminated the nest sites.

Morning flights usually ended between 0800 and 0840 hours in

late August. Bohart and Youssef concluded that initiation and

termination of morning foraging were primarily dependent on

temperature and increasing light levels. Foraging activity in the

evening period was apparently controlled by decreasing light

levels and the opening and availability of new Oenothera flowers. Evening foraging began around 2000 hours, which was

shortly after sunset or about 40 minutes before dusk (this

schedule was maintained throughout the season). Flights ended

around 2100 hours, at which time illumination was extremely

low. No bees were observed to fly in the dark, even during full

moon periods, and bees were never collected at lights.

In this study I examined only nine specimens of S.

oenotherae associated with time data, and all were collected

between 1930 and 2030 hours. Knerer and MacKay (1969) observed a population of S. oenotherae in Toronto, Canada, to be

initially both matinal and crepuscular; however, when pollen

resources became unavailable in the evening, the bees at this

site learned to restrict their activity to morning hours. During

their morning foraging periods, females were observed to leave

nests as early as 0600 hours. Nest plugging usually started

around 0800 hours and all nests were closed by 0900 hours.

Turner (1966) observed a similar switch in foraging activity

periods for S. antiochensis. Early in the season this species collected pollen and nectar from Oenothera both in the early

morning and early evening. Later in the season, apparently associated with pollen competition from Agapostemon texanus

Cresson (Halictidae), S. antiochensis foraged for pollen primarily in the morning. Evening activity became less intense,

and females then foraged primarily for nectar. Although pollen

competition from A. texanus may be involved, Turner emphasized that, as the season progressed, new flowers of the Oenothera host plant opened later in the evening, possibly when the

illumination levels were insufficient for flight activity by S. antiochensis.

Turner reported that in the morning, S. antiochensis opened

their nests 45-60 minutes before they initiated foraging. During his study the mean time of flight initiation was approximately two minutes before sunrise; however, some individuals

left their nests for the first time as late as 48 minutes after sunrise. Evening flight began around 47 minutes before sunset and

usually ended about 30 minutes after sunset. This could extend

as much as 79 minutes after sunset late in the season.

Unlike the Lusoria Group, examined specimens of the two

species of the Texana Group (S. noctivaga, S. texand) were crepuscular and nocturnal, being active exclusively in the evening

hours (Figure 10), as reported by Kerfoot (1967c). Of the 187

specimens of S. texana associated with time data, 175 were

simply labeled "at light." Only four specimens of S. noctivaga

had time data, but all were collected between 1901 and 2100

hours or "at light."
NUMBER 610

(0

<D

f

CO

&

180 n

160 -j

140 -j

120 -j

100 -_

80 -j

60 -_

40 -f

20 -

0

f

5

CD

O

CO

Time of Day

FIGURE 10.—Flight-time records, by hour of day, for Sphecodogastra noctivaga (n = 4) and S. texana (n = 187)

(data from specimen labels examined in this study).

Kerfoot (1967c) reported that nests were usually unplugged

just before sunset (about 1930 hours during midsummer in

Kingman County, Kansas). When the moon was visible by

2130 hours, foraging then continued long into the night until

moonset or until pollen resources were depleted. Kerfoot

termed this the "moonlight period" of the month. The latest reported flight record for S. texana was 0030 hours. If the moon

was not visible after sunset, foraging activity continued until

approximately 2100 hours. During this lunar period, females

were poised at their nest entrances, and if the moon remained

down, they closed their nest by approximately 2130 hours. If

the moon rose after this time, the nests remained closed and

foraging was not resumed. Kerfoot termed this the "crepuscular

period" for S. texana.

Floral Associations

Female Sphecodogastra collect pollen almost exclusively

from Onagraceae (Table 2) but are known to visit flowers of

other plant families for nectar; they are rarely known to collect

non-onagraceous pollen, which apparently occurs only under

duress when evening primrose resources are unavailable (Bohart and Youssef, 1976). Males, however, commonly visit several plant families for nectar.

Onagraceae pollen is apparently not required for Sphecodogastra larval development. Bohart and Youssef (1976) reported

rearing larvae on alfalfa (Medicago sativa L.) pollen. Development time was somewhat longer, but eggs were successfully

reared to adults. They speculated that the advantage of Onagraceae specialization to Sphecodogastra was pollen harvesting efficiency (e.g., S. lusoria often requires less than one

minute to gather a full pollen load; see Table 2) and the precise

timing of their foraging activity with the unusual opening periods of primrose flowers, which reduces competition from other

bees. An additional benefit may be avoidance of nest parasites

by having nests closed during the day, when most potential

cleptoparasites are active. The only report of a possible cleptoparasitic bee association with Sphecodogastra is Bohart and

YousseFs (1976) observation of an overwintering adult Sphecodes female in a nest of Sphecodogastra lusoria (see "Nest

Architecture and Parasites," below).

ONAGRACEAE FLORAL ASSOCIATIONS.—Table 3 lists the

Onagraceae floral records for the 826 female Sphecodogastra

specimens associated with floral data that were examined in

this study as well as a few reliable literature records. These

TABLE 2.—Numbers of Sphecodogastra aberrans and S. lusoria collected from

Oenothera pallida, Maryhill Ferry, 3 miles east of Briggs, Sherman County,

Oregon, 3 June 1963 (P.H. Raven, collector). Only two of the nine S. aberrans

had pollen; all S. lusoria had pollen except the female collected at 0845 hours.

Hours

0530-0559

0600-0629

0630-0659

0700-0729

0730-0759

0800-0829

0830-0859

S. aberrans

0

0

0

0

9

0

0

S. lusoria

14

26

10

3

1

0

1
10 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

TABLE 3.—Records of female Sphecodogastra associated with Onagraceae flora as well as associations indicated

in the literature by other researchers: A = Gregory (1964); B = Stevens (1920); C = Knerer and MacKay (1969);

D = Kerfoot (1967a); M&H = Moure and Hurd (1987; this catalog data should be considered with reservation).

Associated flora 5. aberrans S. antiochensis S. danforthi S. lusoria S. noctivaga S. oenotherae S. potosi S. texana

Calylopohus hartwegii

Camissonia claviformis

Camissonia lanacetifolia

Clarkia pulchella

Gaura coccinea

Gaura suffulta

Gaura sp.

Oenothera albicaulis

Oenothera biennis

Oenothera caespitosa

Oenothera deltoides

Oenothera drummondii

Oenothera elata

Oenothera engelmannii

Oenothera fruticosa

Oenothera grandis

Oenothera laciniata

Oenothera lati/olia

Oenothera macrocarpa

Oenothera nuttallii

Oenothera pallida

Oenothera pilosella

Oenothera rhombipetala

Oenothera villosa

Oenothera sp.

22

135

70

10, A

20

56

16

2

1

32

40

21

18

2

99

62

12

M&H

17

A

9

M&H

2

D

M&H

58

data indicate that Sphecodogastra species are broad oligoleges

of Onagraceae, with individual species visiting a variety of

Oenothera species and other onagraceous genera.

Among the Lusoria Group species, Sphecodogastra lusoria

had the greatest number of floral records (379) in this study and

was recorded from eight species of Oenothera and two species

of Gaura as well as Calylophus hartwegii and Camissonia

claviformis. The preponderance of floral records for S. aberrans associated with Camissonia tanacetifolia (49%) and

Oenothera elata (39%) is interesting but will have to await further study before this pattern can be considered meaningful.

The rarely collected 5. danforthi and 5. potosi are only known

to be associated with Calylophus hartwegii and Gaura coccinea, respectively, but it would be surprising to not find these

species associated with Oenothera given additional study.

Sphecodogastra antiochensis, known only from Antioch,

California, is associated almost exclusively with the similarly

restricted Oenothera deltoides howellii.

Within the crepuscular/nocturnal Texana Group (5. noctivaga, S. texana), 92 of the 94 recorded floral associations were

with Oenothera species.

The more popular Onagraceae species among Sphecodogastra species appear to be the widespread and common Calylophus hartwegii, Gaura coccinea, Oenothera caespitosa, and

the somewhat less common Oenothera macrocarpa. All four

species are known to be associated with three or more Sphecodogastra species. However, given the vagaries associated with

limited, nonsystematic collecting, not much should be concluded from these preliminary data.

SIGNIFICANCE OF Sphecodogastra IN ONAGRACEAE REPRODUCTIVE BIOLOGY.—Sphecodogastra is a highly specialized

oligolege of Onagraceae and has evolved morphological and

behavioral characters to efficiently harvest the resources of its

host plants. However, at least from the literature, these bees often appear to be at best ineffectual pollinators and at worst,

from the plant's perspective, skilled pollen thieves capable of

"stealing" large amounts of Onagraceae pollen (see Figure 4),

with little or no benefit to the plant's reproductive biology. A

similar relationship has been reported by Barrows et al. (1976)

for the andrenid bee Perdita texana, an oligolege of prickly

pear cactus (Opuntia phaeacantha). The cactus flowers are

large and so configured that the relatively small bees can collect large amounts of pollen without normally contacting the

plant's stigmas.

Evidence for this nonmutualistic type of relationship between Sphecodogastra and its host plants initially came from

Gregory's (1963, 1964) excellent studies of Oenothera reproductive biology. He concluded that the most effective pollinators of the plant species he studied were hawkmoths (Sphingidae). "At two colonies [of Calylophus hartwegii], [at]

Monahans and Ector County, over 50% of the flowers had been

pollinated, most lightly, before any hawkmoths were seen. At

Monahans four species of bees were collecting pollen before

sunset and were responsible for most of the pollen transfer"
NUMBER 610 11

(Gregory, 1964:394). He suspected that most of this was

"single flower self-pollination" in a self-incompatible species

and that most cross-pollination was effected by hawkmoths.

For Oenothera drummondii, he concluded that there was certainly some pollination by Celerio [= Hyles, Sphingidae] at

Aransas Pass [San Patricio County, Texas] but Lasioglossum

(Sphecodogastra) texanum (Cresson) took so much pollen during the hour or more before dark that pollination by Celerio

was greatly reduced. In this case the addition of a regular and

abundant visitor species seriously decreased the amount of pollination rather than supplementing it.

The most compelling support for thinking of Sphecodogastra

species as pollen thieves is from Wagner et al. (1985:8), who

reported that S. aberrans and S. lusoria [as galpinisae] were

both observed at a locality in Larimer County, Colorado, to

"collect pollen from Oenothera caespitosa subsp. macroglottis,

primarily in the early mornings." They reviewed Stockhouse's

(1973) study in which, on six successive evenings, he bagged

half the flowers of a population of this Oenothera species to

exclude hawkmoths at night and then removed the bags in the

morning so the two Sphecodogastra species could visit the

flowers. The exposed flowers were visited by hawkmoths in

the evenings. Later, marked capsules were collected and analyzed. Approximately 70% of the flowers visited by hawkmoths produced capsules, whereas only one out of the 150

flowers visited by bees produced capsules. This is strong evidence that Sphecodogastra, at that one locality and time,

played no significant role in the pollination of Oenothera.

Countering the above conclusions is a three-year study of S.

texana by Kerfoot (1967c). He concluded that whereas Oenothera rhombipetala is probably adapted for pollination by

moths, Sphecodogastra texana—although clumsy pollen collectors—were apparently responsible for cross-fertilization of

plants of this species at his study sites. This was because he

saw only one moth during his study.

In summarizing their six-year study of Oenothera pallida

and 5. lusoria, Bohart and Youssef (1976) also concluded that

the hawkmoth Hyles lineata (Fabricius) was probably the most

important pollinator. However, unlike S. texana, S. lusoria females were not clumsy at Oenothera flowers but were very efficient pollen foragers that only rarely made contact with plant

stigmas. However, similar to Kerfoot's (1967a) findings, they

observed no hawkmoths at their study site at Cornish, Utah, until 1973, the last year of their study.

During his two-year study of S. antiochensis. Turner (1966)

observed no hawkmoths at flowers or at electric lights and concluded the moths were either rare or absent. As Turner pointed

out, this raises interesting questions concerning the pollination

of the self-incompatible host plant, Oenothera deltoides howellii, which is confined to the Antioch dune area. Turner observed a shift in foraging activity of 5. antiochensis females

during the year. Early in the season, when there was no competition from Agapostemon texanus (and apparently none from

hawkmoths), pollen was plentiful and females harvested Oenothera pollen with alacrity (Turner, 1966), visiting only one

flower during each foraging trip (of no use for cross-pollination). Later in the season, with floral resource competition from

A. texanus, foraging Sphecodogastra females appeared rather

frantic and needed to visit seven or more flowers to obtain a

full pollen load. During this period, Turner frequently observed

females with partial pollen loads land on several different stigmas and was of the opinion that they easily could have effected

pollination. He concluded that the assumption that only hawkmoths pollinated hawkmoth flowers was not necessarily true.

As documented by Gregory (1963, 1964), Stockhouse

(1973), and Wagner et al. (1985), hawkmoths (especially Hyles

lineata), when present, are undoubtedly the most effective pollinators of Oenothera and related genera. However, as reviewed above, three long-term studies of Onagraceae pollination ecology (Turner, 1966; Kerfoot, 1967a; Bohart and

Youssef, 1976) reported situations where hawkmoths were

found to be rare or apparently absent. Turner addressed this issue of erratic sphingid faunal representation and mentioned

their migratory capabilities, citing Williams (1957:168), who

(without providing references) wrote strongly concerning

Hyles lineata migration: "The insects breed during the winter

somewhere in Central or South America, almost certainly in

the semiarid climate of the western coast. Thence they move

north in the spring into the southern states where, as in Europe

[the subspecies H. lineata livornica (Esper)], they become a

minor pest of vines." Williams cited Grant (1937) as demonstrating statistically significant evidence supporting correlations for heavy migration years between the New and Old

World subspecies. In her paper, Grant documented the apparent

erratic representation of//, lineata in North America but made

no mention of long distance overwintering migration; however,

she indicated that H. lineata might migrate gregariously after

the larvae completely destroyed their host plants (many hosts

but especially vineyard vines). Whether Hyles migration involves travel to distant overwintering sites or more local movements prompted by resource depletion is apparently unresolved

(D. Ferguson, pers. comm., 1993). Whatever the cause, local

representation of these moths is erratic and undoubtedly has

implications for Onagraceae pollination. These observations

suggest that additional studies similar to those of Stockhouse

(1973) should be pursued for other Sphecodogastra-Onagraceae relationships before it can be concluded that these bees

never play a significant role in the cross-pollination of their

host plants, especially in situations where hawkmoth populations are at low densities or are absent.

NON-ONAGRACEAE FLORAL ASSOCIATIONS.—Associations

with plant genera from 19 non-Onagraceae families are listed

below, as recorded from Sphecodogastra specimens examined

in this study. The mostly commonly visited family is Asteraceae (43% of female records; 65% of male records). Bohart

and Youssef (1976) reported S. lusoria collecting pollen of

Lygodesmia grandiflora (Asteraceae): one female had a mixed

load of pollen (90% Oenothera and 10% Lygodesmia). They
12

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

also noted P.F. Torchio's observation of a female attempting to

collect pollen from Cleome serrulata (Capparidaceae).

Sphecodogastra aberrans: Allionia (3d), Arnica (1?),

Chrysothamnus (49), Coreopsis (1 d), Cucurbita(Id1), Geranium (1 9), Grindelia (2d), Gutierrezia (1 d), Helianthus

(Id1), Heterotheca (Id),Iris (19), Melilotus (1 d), Taraxacum (1 9), Verbesina(19), Viguiera(Id).

Sphecodogastra antiochensis: Layia (19).

Sphecodogastra lusoria: Arenaria (1 ?), Argemone (1 9 ),

Asclepias (Id), Barbarea (29), Brassica (\d), Callirhoe

(19), Chrysothamnus (3d), Circium (2d), Cleome (19,

9<f), Coreopsis (19), Cucurbita (109), Descurainia (19),

Engelmannia (2d), Eriogonum (Id), Galpinsia (19), Geramufif (19), Grindelia (19), Gutierrezia (19), Helianthus

(3 9, \2d), Iris (Id), Lactuca (19), Iipi/ius (2<J), Mr/oco-

/Arix (2 9), Malvastrum (19), Medicago (19), Melilotus

(3 9,2d), Monarda (3d), Poliomintha (3d), Potentilla

(Ad), Prunus (39), Pyrrhopappus (29, Id), Salsola (19),

Senecio (19), jfr/fou (2 9), Solidago (5 9 ), Sphaeralcea

(19), Tamarix (3 9), Taraxacum (19), Thelesperma (19),

Xanthisma (5 d1).

Sphecodogastra noctivaga: Chrysothamnus (1 9).

Sphecodogastra oenotherae: Carduus (3 d ) , Cichorium

(2c?), Coreopsis (\9,2d), Eupatorium (Id), Salix (\d),

Taraxacum (Id), Tragopogon (29,22d).

Sphecodogastra potosi: Asclepias (Id1).

Sphecodogastra texana: Allionia (109), itofens (19), M?«fze//a(19).

Nesting Biology

The nesting biologies of four Sphecodogastra species have

been studied: 5. antiochensis (Turner, 1966; two-year study,

Antioch, Contra Costa County, California; details of nest architecture were not reported); 5. lusoria (as galpinsiae, Bohart and

Youssef, 1976; six-year study, Cornish, Cache County, Utah);

S. oenotherae (Knerer and MacKay, 1969; one-year study, Toronto, Canada); and 5. texana (Kerfoot, 1967a; three-year study,

Kingman County, Kansas). Bohart (in Bohart and Youssef,

1976) indicated he had unpublished data for S. aberrans and

included some information on this species in his paper on S. lusoria. The following review is based on the papers cited above.

SOCIAL BEHAVIOR.—Sphecodogastra species are primarily

solitary bees but show several tendencies towards limited social behavior. Females of all species studied return to their natal nests for overwintering. For S. oenotherae this can involve

as few as one but usually three to four females and sometimes

as many as 10 individuals. Inseminated females of 5. lusoria

have been reported to number from one to nine in overwintered

nests, resulting in an early-season transitional period during

which more than one active female can be found in a single

nest. Of 33 nests of S. texana examined by Kerfoot (1967a), 12

contained more than one female; however, among all the females in these 12 nests, only one female appeared to be fertilized, with the other females showing no mandibular wear or

ovarial development. As for other species, only one female reuses the natal nest for brood rearing, and eventually surplus females leave to establish their own nests. However, one nest of

S. texana was found to contain two fertilized and two unfertilized females.

Bohart and Youssef (1976) reported a similar situation for S.

lusoria in the early season of 1970 in which a few nests contained up to three females, but later in the same year, when provisioning had begun, all nests appeared to be occupied by a

single female. Under exceptional conditions S. lusoria appeared to be truly communal. There was no Oenothera bloom

at the Cornish, Utah, site in 1972. The bees that year emerged

but finding no host flowers returned to their shared natal nests

with undeveloped ovaries and resumed their dormancy. These

bees emerged in 1973 as two-year-old sisters. Up to 30 percent

of the nests examined that year were communal, with more

than one female sharing a common opening burrow and apparently maintaining their own burrow beneath this (the number of

females in a nest always corresponded with the number of subburrows). Bohart and Youssef (1976) speculated that this communal behavior was possibly the result of increased tolerance

between these unusually old sisters who had shared their natal

nests.

Turner (1966) provided strong indications that S. antiochensis may be communal. He observed "guard" bees at nest entrances as well as several long-established nests that contained

many females. The nature of this social behavior needs further

investigation and clarification.

NEST ARCHITECTURE.—Most Sphecodogastra species are

known to nest in stabilized sand in open xeric areas (S. antiochensis, S. lusoria, S. texana). One observation was made of S.

lusoria nesting in vertical banks (Bohart and Youssef, 1976).

This occurred only among bees of the 1968 second generation

brood of S. lusoria at Cornish, Utah. However, the third (overwintering) generation of this population reverted to nesting in

flat ground. Bohart indicated that unlike S. lusoria, S. aberrans

"is generally found in more mesic and highland surroundings,

on brush-covered slopes or open woodlands" (Bohart and

Youssef, 1976:186). Sphecodogastra oenotherae was reported

to nest on gently sloping ground in patches of dirt between

well-cared-for lawns and gardens (Stevens, 1920).

All species studied showed slight to conspicuous tendencies

toward nesting gregariously and forming distinct aggregations.

Kerfoot (1967a) observed S. texana to be gregarious at times,

with 100 or more nests found within a small area, and at other

times having nests widely scattered. Bohart and Youssef (1976)

reported for S. lusoria that 15 to 20 nests per square meter were

observed in years of high population density, but only 10 nests

in a few square meters were seen during low-density years

even though similar, empty areas surrounded these aggregations. Sphecodogastra oenotherae formed aggregations with a

maximum density of about 25 nests per square meter, with a
NUMBER 610 13

minimum internidal distance of 2 cm. Turner (1966) indicated

that S. antiochensis aggregations never surpassed seven burrows per square meter.

Nests were always, at least initially, associated with conspicuous tumuli (3.0-5.5 cm in diameter for S. texana; usually 5.0

cm in diameter for S. lusoria). Main nest burrows are usually

vertical and straight—remarkably so as reported by Kerfoot

(1967a) for 5. texana. He reported that nest depths ranged from

20.0 cm (mean for S. oenotherae during provisioning period;

hibernacular cells averaged 30.0 cm deep) to 55.0-75.0 cm (5.

lusoria) and 45.5-145.0 cm (S. texana; mean = 102.0 cm). This

range possibly represents increased depths in drier soils (L.

Packer, pers. comm., 1994).

The main burrow of all Sphecodogastra nests studied was

usually plugged during nonforaging periods. For S. antiochensis, Turner (1966) said these plugs consisted of 5 or 6 distinct

and compact sand pellets. Oddly, nests of this species were often plugged before all females returned. In most cases these females made unsuccessful attempts at entry but eventually flew

away to await reopening of the nests during the next foraging

period. Kerfoot (1967a) observed that S. texana closed their

nests with sand plugs that mutillids could not penetrate (even

after concerted effort), but that bees easily passed through. He

speculated that when the bees were ready to exit the nests they

moistened the plugs (presumably with nectar).

Brood cells of Sphecodogastra are horizontal (S. lusoria) or

slope slightly downward (S. oenotherae, S. texana). The cells

are unusual for halictines in that they are only partially lined

with a waxy secretion. This is reported as being over the distal

three-fourths of the cell for S. oenotherae; the distal two-thirds

for 5. lusoria (with the lining being thickest under a depression

for the pollen ball); and only partially covering just the bottom

of the cell for S. texana. Cells were sessile (S. oenotherae),

connected to the main burrow by extremely short lateral tunnels (1-3 mm, 5. lusoria), or connected by distinct laterals (average length = 2.6 cm, S. texana; see "Nest Architecture and

Social Behavior," below). The number of cells found per nest

were 12-20 for S. oenotherae (6-20 cm below the surface) and

varied for S. lusoria, with the greatest number being 31 (20-43

cm deep). The number of cells per nest appeared to be unusually low for S. texana, but numbers increased throughout the

season (cell depth ranged from 20 to 110 cm below the surface). Kerfoot (1967a) reported that 16 nests excavated in June

averaged 1.1 old cells and 1.0 occupied cells; 8 nests in July

averaged 3.6 old and 1.9 occupied cells; 5 nests in August averaged 11.2 old and 3.2 occupied cells. This apparent low number of brood cells may be associated with Kerfoot's (1967c)

observation that S. texana actively provisions cells primarily

during its "moonlight period" of the month and does little provisioning and egg laying during the "crepuscular period" of the

month (see "Daily Flight Activity Records").

NEST ARCHITECTURE AND SOCIAL BEHAVIOR.—The nest architecture of Sphecodogastra species studied includes characteristics typical of solitary halictines as well as some characteristics correlated with primitively eusocial species. The brood

cells of most solitary halictines are not clustered and are connected to the main burrow by elongate lateral tunnels. However, Packer (1991) pointed out that there are some solitary halictines with brood-cell clusters and cavities that may represent

a reversal to solitary behavior. Some primitively eusocial species (e.g., Evylaeus malachurus) construct clustered cells that

have the surrounding soil excavated, are left open during the

early stages of larval development, and are sessile, i.e., not connected to the main burrow by lateral tunnels (Packer, 1991).

The brood cells of S. texana were connected to the main burrow by distinct lateral burrows. Cells were loosely "clumped"

at different depths, but Kerfoot (1967c) thought this might be

correlated with the different foraging activity periods observed

for this species (i.e., the crepuscular versus the moonlight periods; see "Daily Flight Activity Records").

Sphecodogastra oenotherae at Toronto, Canada, is apparently univoltine, and Knerer and MacKay (1969:293) reported

that the species "lacks even the most rudimentary social behavior." Nevertheless, the nests of this species have some eusocial

characteristics: sessile, clustered cells with cell closures consisting of a porous layer of loose soil particles. Knerer and

MacKay also observed that formation of the pollen ball was not

immediately followed by oviposition and that this delay may

exceed 24 hours. They concluded that these characteristics may

represent a preadaptation towards social behavior that might

have been arrested because of the species' specialized association with Oenothera. It would be interesting to study the nesting biology of S. oenotherae in southern areas of its range

where the possibility of multivoltinism is likely.

Bohart and Youssef (1976) observed that S. lusoria females

shape more than one pollen ball in advance of egg laying and

leave the cells (which are closely positioned but not clustered)

and short lateral burrows (1-3 mm in length) initially unplugged—characteristics of social halictines. These researchers

wondered whether this could be a step leading to social behavior or if it represented a habit retained from a more social ancestor. A large part of this question may be answered after the

phylogenetic relationships of Sphecodogastra to related halictines are resolved. Much biological work, however, remains to

be pursued—in particular, clarification of the "social" behavior

of S. antiochensis, study of 5. aberrans, and observation of 5.

oenotherae in the southern latitudes of its range. Nothing is

currently known about the nesting biologies of the newly described S. danforthi and S. potosi.

NEST ARCHITECTURE AND PARASITES.—The matinal, crepuscular, and nocturnal periods of foraging activities of Sphecodogastra combined with nest plugging during nonforaging

periods most likely contribute to the paucity of nest parasites

reported for this genus. The cleptoparasitic bees of the genus

Sphecodes, commonly associated with halictine nests, have

only tenuously been associated with Sphecodogastra. In their

six-year study of S. lusoria, Bohart and Youssef (1976) reported

only one observation of Sphecodes in a host nest. This was a
14 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

single, adult female found in a hibernacular (nonbrood) cell.

Presumably, this female made her way into the hibemaculum

of this one nest and managed to overwinter there. No other

cleptoparasitic bee associations with Sphecodogastra have

been reported.

Kerfoot (1967a) occasionally found mutillids (Myrmillodes

grandiceps Blake) in nests of 5. texana that had been left open,

but none of these were in brood cells. He speculated that these

wasps simply might have been trying to avoid the hot sand of

the ground surface. Turner (1966) observed the nest sites of S.

antiochensis also to be associated with mutillids (Photopsis

spp., Pseudomethoca anthracina (Fox)). Mutillids inspecting

nest openings of this species were always deterred from entering by brief encounters with "guard" bees. One female of P. anthracina was artificially introduced into a guarded S. antiochensis nest and was repeatedly repelled by the bee at the nest

entrance. Of 14 nests excavated at Antioch, Turner found no

evidence of parasites or predators or evidence of cell entry.

Knerer and MacKay (1969) reported finding one dying 5.

oenotherae female at the bottom of her nest with a third instar

larva of the conopid fly (Thecophora) completely filling her

abdomen. Bohart and Youssef (1976) found adult phorid flies

in the main burrows of two nests of S. lusoria, and larval

phorids in a few cells, but did not believe these flies were responsible for the absence of bee larvae in the infested cells.

Kerfoot (1967a) dissected 62 adult S. texana (57 females; five

males) and found none to have internal parasites.

Miscellaneous associates reported in Sphecodogastra nests

include nematodes in fecal material (5. lusoria) and mites in

cells and on pollen of S. lusoria (anoetids [Histiogaster], pyemotids [Parapygmephorus], and scutacarids). Kerfoot (1967a)

reported an unidentified fungus from the cells of S. texana.

General predator reports are confined to Bohart and Youssef

(1976) finding three nest cells of the bee-storing sphecid wasp,

Philanthus gibbosus (Fabricius), with several intact S. lusoria

of both sexes.

Systematics

Sphecodogastra Ashmead

Sphecodogastra Ashmead, 1899. [Type species: Parasphecodes texana (Cresson). Monotypic and original designation. (=Sphecodes texana Cresson,

1872).]

DIAGNOSIS.—Female Sphecodogastra can be distinguished

from other Halictidae by their distinctively reduced scopae

composed of simple, apically hooked, and linearly arranged setae on the ventral surface of the hind femora (Figures 2, 3, 26).

These setae are surrounded by conspicuously nonsetose areas

on the anterior and posterior femoral surfaces (this pattern is

somewhat diffused in 5. antiochensis, Figure 25). The orange

abdomens and extremely enlarged ocelli of males and females

of S. noctivaga and 5. texana make both species highly distinctive (Figures 11, 29). Males of the Lusoria Group (dark species) are more problematic to diagnose. Most helpful is the pattern of long sternal vestiture of most species (Figure 38;

vestiture highly distinctive in S. aberrans. Figure 37). Also

characteristic is the central depression of sternum 6, which has

a low, longitudinal median elevation (Figures 66, 99).

DESCRIPTION.—The format and character numbering in the

following generic description follow those of McGinley

(1986). Characters thought to be nonvariant, or primarily so,

for Sphecodogastra are presented in small capital letters (if minor variation does occur, this is described in individual species

descriptions).

FEMALES: (1) Length 6.6-11.2 mm (mean = 8.5, n = 40);

(2) wing length 1.9-3.3 mm (mean = 2.6, n = 40); (3) abdominal width 2.0-3.3 mm.

Structure: (4) Head short (Figure 129) to moderately elongate (Figure 52); length/width ratio 0.80-1.40 (mean = 0.95, n

= 40). (5) Gena, at midpoint, exceeded by width of compound

eye to greatly exceeded by eye; (6) GENA ROUNDED POSTERIORLY, NOT PRODUCED OR ANGULATE. (7) SUPRACLYPEAL AREA

EVENLY ROUNDED, (8) WEAKLY PROTUBERANT. (9) Clypeus

projecting approximately 0.40-0.92 times its length below

lower margin of eyes; (10) CLYPEAL SURFACE BROADLY

ROUNDED; (11) clypeal surface usually without median longitudinal sulcation (present only in S. noctivaga). (12) FRONTAL

CARINA WEAKLY DEVELOPED, INCONSPICUOUS. (13) Distance

between lateral ocellus and eye slightly greater than distance

between lateral ocellus and hind margin of vertex (ocular-ocellar distance ranging from approximately 0.5 to 2.0 times lateral

ocellar diameter); (14) distance between lateral ocelli slightly

to greatly exceeding ocular-ocellar distance; (15) LATERAL

OCELLI JOINED ABOVE BY AT MOST A WEAK, INCONSPICUOUS

IMPRESSED LINE. (16) Inner margins of compound eyes converging below or nearly parallel. (17) HYPOSTOMAL CARINA

WELL DEVELOPED AND UNIFORM; (18) ANTERIOR ANGLE OF

HYPOSTOMAL CARINA NARROWLY ROUNDED; (20) ANTERIOR

CARINA NEARLY PERPENDICULAR TO LONGITUDINAL CARINA.

(21) Scape reaching top of vertex or slightly beyond (S. noctivaga, S. texana); (22) PEDICEL SLIGHTLY LONGER THAN WIDE,

slightly shorter than flagellomere 1 (S. noctivaga, S. texana), or

lengths subequal; (23) FLAGELLOMERE 1 SUBEQUAL IN LENGTH

TO FLAGELLOMERE 2; flagellum with distinctive sensillar patterns in S. antiochensis and S. lusoria (Figures 42,76). (24) LABRUM WITH BASAL AREA AND DISTAL PROCESS; unlike most

species, basal area in Texana Group much broader than long

(Figures 108, 142); (25) BASAL ELEVATION WELL DEVELOPED;

(26) BASAL LATERAL DEPRESSIONS ABSENT; (27) DISTAL KEEL

NARROW AS SEEN IN FRONTAL VIEW; (28) DISTAL LATERAL

PROJECTIONS ABSENT (present in many Lasioglossum (McGinley, 1986, fig. 7)); (29) FIMBRIAL SETAE ACUTELY POINTED.

(30) Mandible moderate in length to conspicuously elongate;

(31) MANDIBLE USUALLY WITH DEFINED SUBAPICAL TOOTH

(broadly rounded, not defined in S. danforthi).

(32) PRONOTAL LATERAL ANGLE BROADLY OBTUSE; (33)

PRONOTAL LATERAL RIDGE BROADLY INTERRUPTED BY OBLIQUE LATERAL SULCUS; (34) LOWER PORTION OF LATERAL
NUMBER 610 15

RIDGE BROADLY ROUNDED. (35 ) MESOSCUTAL ANTERIOR EDGE

ROUNDED, NOT BILOBED, AND (36 ) MODERATELY ELEVATED

FROM PRONOTUM AT CENTER; (37 ) MEDIAN MESOSCUTAL LINE

USUALLY NOT IMPRESSED (impressed only in S. antiochensis);

(38 ) PARAPSIDAL LINES ELONGATE, APPROXIMATELY 0.40-0.50

TIMES LENGTH OF MESOSCUTUM, WITH WEAKLY IMPRESSED

LINE EXTENDING TO POSTERIOR EDGE OF MESOSCUTUM (e.g.,

Figures 98, 145). (39) MEDIAN SCUTELLAR IMPRESSION VIRTUALLY ABSENT (best developed in S. aberrans). (40) Dorsal surface of propodeum 0.59-0.78 times the length of scutellum and

approximately 1.13-1.50 times length of metanotum, (41)

ONLY FAINTLY DEPRESSED CENTRALLY, (42 ) POSTERIOR MARGIN ROUNDED; (43 ) PROPODEAL TRIANGLE INCONSPICUOUS,

WITHOUT ELEVATED, MEDIAN V-SHAPED AREA OR DISTINCT

LATERAL MARGINS; (44) lateral propodeal carinae on posterior

vertical face well developed and complete to weakly developed

and extending no more than one-third basal length of posterior

surface. (45) Inner hind tibial spur usually with 4 or 5 moderately elongate teeth (e.g., Figures 58, 60); teeth somewhat

longer and more numerous (up to 7) in 5. antiochensis (Figure

59); teeth shorter and less numerous (3 or 4) in Texana Group

(Figures 64, 65).

(46) Lateral edge of metasomal T2 rounded to nearly

straight.

Sculpture: (47) FACE SHINY, (48 ) DENSELY AND CONTIGUOUSLY PUNCTATE BELOW OCELLI; AREA BETWEEN OCELLI AND

ANTENNAE UNIFORMLY PUNCTATE OR WITH PUNCTURES SEPARATED BY 1-2 TIMES THEIR WIDTHS IMMEDIATELY ABOVE AND

LATERAD OF ANTENNAE (S. noctivaga, S. texana). (49) PUNCTATION OF VERTEX NEAR EYE MUCH MORE SPARSE AND FINE

THAN THAT ON LOWER FACE, PUNCTURES SEPARATED BY 1-3

TIMES THEIR WIDTH; (50) VERTEX BEHIND OCELLI OBSCURELY

PUNCTATE, WITHOUT TRANSVERSE STRJATIONS. (51) Supraclypeal area polished or granulate, (52) densely to sparsely punctate. (53) Clypeus entirely polished or tessellate basally; (54)

punctation sparse, most punctures separated by 1-5 times their

widths. (55) Hypostoma striolate, becoming smooth and polished on anterior half in some species, PUNCTATION OBSCURE,

VIRTUALLY ABSENT.

(56) Mesoscutum shiny and polished with surface tessellation between punctures confined to anterior one-third or mesoscutum moderately dull with surface entirely tessellate; (57)

punctation moderately dense, most punctures separated by 1-2

times their widths. (58) SCUTELLAR PUNCTATION SIMILAR TO

THAT OF MESOSCUTUM, BUT USUALLY FINER. (59) METANOTUM

GRANULATE TO RUGULOSE, PUNCTATION OBSCURE OR ABSENT.

(60) PRE-EPISTERNUM USUALLY RUGULOSE (very fine in 5. noctivaga); (61) HYPOEPIMERAL AREA AND MESEPISTERNUM USUALLY RUGULOSE (finely s o in 5. lusoria, extremely fine in S.

noctivaga; mesepisternum weakly strigulate ventrally in S. texana), WITHOUT DISTINCT PUNCTURES; (62 ) UPPER PORTION OF

METEPISTERNUM STRIGULATE, BECOMING GRANULATE BELOW.

(63) Dorsal surface of propodeum strongly rugo-striate (Figure

15), rugulose (Figure 16) to ruguloso-striolate (Figure 19);

(64) surface smooth or alveolated. (65) Tl usually shiny and

polished (somewhat dull tessellate in S. noctivaga); (66) punctation fine to extremely fine, moderately dense (punctures separated by their widths) to very sparse (punctures separated by

2-3 times their widths).

Coloration: (67) HEAD AND THORAX DARK BROWN, abdomen mostly orange (S. noctivaga, S. texana) or dark brown

(other species); T1-T4 OR T2-T4 WITH APICAL HYALINE BANDS

THAT COVER ABOUT ONE-FOURTH OF TERGAL SURFACE (presence of hyaline band on Tl variable among species; bands relatively inconspicuous in species with orange abdomens). (68)

CLYPEUS WITHOUT MACULATION; LABRUM DARK BROWN. (69 )

Flagellum dark brown to amber. (70) Tegula yellow-translucent

to brown. (71) Wing membrane hyaline to pale yellowish

brown; veins and stigma pale yellowish amber to brown. (72)

Legs usually darkly pigmented (mostly amber in Texana

Group).

Vestiture: (73) PUBESCENCE OF HEAD BETWEEN VERTEX

AND ANTENNAE PLUMOSE (hairs conspicuously short in Texana

Group); (74) white to pale yellowish brown. (75) Pubescence

of thorax white to pale yellowish brown; (76) mesoscutal hairs

elongate and sparse to short and adpressed; pleuron with or

without short, suberect to adpressed hairs. (77) HIND TIBIAL

HAIRS WHITE TO YELLOWISH WHITE; HAIRS CONCOLOROUS

(HAIRS OF DORSAL SURFACE NOT DARKER THAN THOSE ON

OTHER AREAS). (78) ANTERIOR HAIRS ON Tl , (79) BASAL HAIR

BANDS ON T2-T4 WHITE. (80 ) ACARINARIUM ON ANTERIOR

SURFACE OF Tl ABSENT. (81) Basal hair bands on T2-T4

present (at most, covering basal one-fourth of terga) to absent;

apical hair bands sometimes present on T3 and T4.

MALES: As described for females except as follows: (1)

length 6.4-9.7 mm (mean = 7.9, n = 40); (2) wing length

1.4-2.8 mm (mean = 2.1, n = 40); (3) abdominal width 1.4-2.4

mm.

Structure: (4) Head length /width ratio 1.00-1.14 (mean =

1.1, n = 40). (10) CLYPEAL SURFACE BROADLY ROUNDED (this

differs from the flattened clypeal surface characteristic of the

males of most New World Lasioglossum, sensu stricto). (16)

INNER MARGINS OF COMPOUND EYES CONVERGING BELOW. (21)

SCAPE MUCH SHORTER THAN IN FEMALES, AT MOST JUST

REACHING LEVEL OF MEDIAN OCELLUS (Texana Group); (22)

PEDICEL SLIGHTLY WIDER THAN LONG, CLEARLY SHORTER

THAN FLAGELLOMERE 1; (23) flagellomere 2 approximately

1.5-2.0 times length of flagellomere 1. (24) LABRUM WITHOUT

DISTAL PROCESS (apical edge of basal process slightly projecting only in S. aberrans (Figure 55)); (25) BASAL ELEVATION

ABSENT; (26 ) BASAL LATERAL DEPRESSIONS ABSENT; (30 )

MANDIBLE SHORT, JUST REACHING OPPOSING CLYPEAL ANGLE,

(31) WITHOUT SUBAPICAL TOOTH.

(37) Median mesoscutal line impressed or not impressed.

(44 ) LATERAL PROPODEAL CARINAE PRESENT ON VERTICAL

FACE, NOT REACHING DORSAL SURFACE OF PROPODEUM. (45 )

INNER HIND TIBIAL SPUR SERRATE ON BOTH EDGES.
16 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

Sculpture: (51) SUPRACLYPEAL AREA TESSELLATE, (52)

DENSELY PUNCTATE (punctures separated by less than the

width of their diameter). (53) CLYPEUS SOMEWHAT DULL, NOT

HIGHLY POLISHED.

(57 ) MESOSCUTAL PUNCTATION DENSE, PUNCTURES SEPARATED BY LESS THAN THEIR DIAMETERS. (66) PUNCTATION ON

Tl BETTER, STRONGER THAN IN FEMALES (punctures only moderately fine, separated by 1-2 times their diameters).

Coloration: (68) CLYPEUS WITH APICAL YELLOW MACULATION; LABRUM YELLOW (yellow pigmentation relatively limited in S. potosi). (70) Tegula yellow-translucent to brown. (72)

TARSI YELLOW; TIBIAE YELLOW AT BOTH ENDS, VARYING

AMOUNTS OF DARK PIGMENTATION MEDIALLY.

Vestiture: (73) UNLIKE FEMALES, SHORT, ADPRESSED HAIRS

PRESENT ON FACE (from base of clypeus to area immediately

dorsad of antennae or extending to vertex, as in S. lusoria, S.

antiochensis). (76) Mesoscutal hairs elongate and sparse to

short and adpressed; pleuron with or without short, suberect to

adpressed hairs. (81) APICAL HAIR BANDS NOT DEVELOPED ON

T3 OR T4. (82, 83) VESTITURE ON S4 AND S5 ELONGATE, CONSPICUOUS, hairs confined to apical margin (S. aberrans; Figure

37) or distributed over entire sternal surface (Figure 38).

Terminalia: (84) S7 WITH ELONGATE, NARROW, APICALLY

ROUNDED MEDIAN PROCESS; lateral arms slender (Figure 91) to

moderately developed (Figure 81); (85) S8 usually moderately

developed (Figure 81), short and somewhat reduced in S. danforthi (Figure 91) and S. potosi (Figure 138); apex of median

process usually rounded (Figure 81), truncate (Texana Group,

Figures 115, 150); (86) GONOBASE WELL DEVELOPED; (87)

GONOSTYLUS SIMILARLY SHAPED AMONG SPECIES (somewhat

slender in S. aberrans, Figure 69); WITHOUT CONSPICUOUS

HAIR TUFTS; (88) RETRORSE MEMBRANOUS LOBE PRESENT,

WELL DEVELOPED, (89) lobes usually narrow and parallel-sided

(e.g., Figure 78, broader basally (5. aberrans, S. lusoria, Figures 67, 100), to very broad and twisted ventrally (S. texana,

Figures 146, 149)); (90) VOLSELLA SIMILAR IN SHAPE AMONG

SPECIES AS ILLUSTRATED HEREIN.

Key to Sphecodogastra Females

(See Appendix 1 for figure identifications)

Abdomen orange; ocelli conspicuously enlarged, distance between compound eye

and lateral ocellus less than lateral ocellar diameter [Figure 11] 2

Abdomen darkly pigmented, brown to dark brown; ocelli moderately large but not

conspicuously so, distance between compound eye and lateral ocellus 1.5-2.0

times lateral ocellar diameter [Figure 12] 3

2( 1). Mandibles conspicuously elongate, clearly extending beyond opposing clypeal angles [Figure 13]; face broad ventrally, inner margins of compound eyes nearly

parallel [Figure 106] 5. S. noctivaga (Linsley and MacSwain)

Mandibles not elongate, at most just reaching opposing clypeal angles [Figure 14];

face relatively narrowed ventrally, inner margins of compound eyes converging

below [Figure 140] 8. S. texana (Cresson)

3(1). Dorsal propodeal surface strongly rugo-striate [Figure 15]

6. S. oenotherae (Stevens)

Dorsal propodeal surface rugulose [Figure 16] to ruguloso-striolate [Figure 19]

4
NUMBER 610 17

4(3). Mandibles elongate, extending beyond opposing clypeal angles [Figures 17, 18];

pleuron with short adpressed pubescence beneath elongate pleural hairs 5

Mandibles not elongate, at most just reaching opposing clypeal angles [Figure 14];

pleuron with or without short adpressed pubescence beneath elongate pleural

hairs 6

5(4). Mandibles straight-edged laterally, extending conspicuously beyond clypeus [Figure 17]; mandibular subapical tooth poorly defined, broadly rounded [Figure

86]; propodeal dorsal surface ruguloso-striolate [Figure 19]; currently known

only from western Texas [Figure 83] 3. S. danforthi, new species

Mandibles broadly curved laterally, not extending conspicuously beyond clypeus

[Figure 18]; subapical tooth well defined [Figure 18]; propodeal dorsal surface

rugulose [Figure 20]; known from Mexico and southern New Mexico [Figure

83] 7. S. potosi, new species

6(4). Pleuron with short, adpressed pubescence beneath elongate pleural hairs [Figure

21]; supraclypeal area polished, punctation sparse, many punctures separated

by 3-4 times their diameters [Figure 23]; wing membranes hyaline

4. S. lusoria (Cresson)

Pleuron without short, adpressed pubescence [Figure 22]; supraclypeal area dull,

distinctly to obscurely granulate, punctation dense, most punctures separated by

twice their diameters or less [Figure 24]; wing membranes pale yellowish

brown 7
18 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY
NUMBER 610 19

7(6). Scopal hair fringe on hind femur not sharply differentiated from surrounding hairs

[Figure 25]; clypeus with basal area polished, subapical area with widely separated, small punctures; dorsal propodeal surface coarsely ruguloso-striolate

[Figure 27]; known only from Antioch (Contra Costa County), California [Figure 92] 2. 5. antiochensis, new species

Scopal hair fringe on hind femur sharply differentiated from surrounding hairs

[Figure 26]; clypeus with basal area tessellate, subapical area with contiguous,

large punctures; dorsal propodeal surface finely rugulose [Figure 28]; widespread in western North America, known in California only from Plumas and

Riverside counties [Figure 51] 1.5. aberrans (Crawford)

2(1).

Key to Sphecodogastra Males

(See Appendix 1 for figure identifications)

Abdomen orange; ocelli conspicuously enlarged, distance between compound eye

and lateral ocellus less than lateral ocellar diameter [Figure 29] 2

Abdomen darkly pigmented, brown to dark brown; ocelli moderately large, distance between compound eye and lateral ocellus slightly greater than lateral

ocellar diameter [Figure 30] 3

Retrorse lobes of genitalia very broad and twisted ventrally [Figures 31,32] ... .

8. S. texana (Cresson)

Retrorse lobes of genitalia narrow, flat, not twisted ventrally [Figure 33]

5.5 . noctivaga (Linsley and MacSwain)

[I know of no reliable external characters to differentiate the males of these two

species; see "Diagnosis" for S. noctivaga.]
20 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

3(1). Dorsal propodeal surface strongly rugo-striate [Figure 34]; primarily from eastern

North America [Figure 117] 6. S. oenotherae (Stevens)

Dorsal propodeal surface finely rugulose [Figure 35] to at most ruguloso-striolate

[Figure 36]; primarily from central and western North America [Figures 51, 83,

92] 4
NUMBER 610 21

4(3). Fringe hairs on posterior margins of S4 and S5 conspicuously developed, clearly

differentiated from shorter vestiture on S3 [Figure 37]; antennal flagellum

without distinctive sensillar pattern; central depression of S6 extremely well

developed [Figures 37, 66]; penis valves sharply angulate [Figure 39], dorsal

and apical surfaces concave and sharply carinate laterally [Figure 39]

1. S. aberrans (Crawford)

Fringe hairs on posterior margins of S4 and S5 similar to posterior hair fringe on

S3 [Figure 38]; antennal flagellum with [Figure 42] or without distinctive sensillar pattern; central depression of S6 only moderately developed [Figure 38];

penis valves rounded [Figure 40], dorsal and apical surfaces rounded [Figure

40] 5

5(4). Antennal flagellum without distinctive sensillar pattern; vertex and ocellar area

without short, adpressed pubescence [Figure 41]; antennae only moderately

elongate, second flagellomere approximately 1.3-1.5 times the length of flagellomere 1 [Figure 43]; posterior propodeal surface lacking short adpressed hairs

[Figure 45] 6

Antennal flagellum with distinctive sensillar pattern [Figures 42, 76]; vertex and

ocellar area with short, adpressed pubescence [Figure 30]; antennae elongate,

second flagellomere nearly twice the length of flagellomere 1 [Figure 44]; posterior propodeal surface with short adpressed hairs [Figure 46] 7
22

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

6(5). Mandible with extensive area of yellow pigmentation; apical half of retrorse lobes

of genitalia curved laterad [Figure 47] 3. 5. danforthi, new species

Mandible darkly pigmented, without extensive area of yellow; apical half of retrorse

lobes straight, not curved laterad [Figure 48] 7. 5. potosi, new species

7(5). Tl punctation dense at center of disc, most punctures separated by 2-3 times their

width [Figure 49; difficult to see on SEM image, much more obvious under

light microscope]; known only from Antioch (Contra Costa County), California

[Figure 92] 2. S. antiochensis, new species

Tl punctation less dense at center of disc, relatively sparse, most punctures separated by 3-4 times their width [Figure 50; see above comment]; widespread in

western North America from Canada to Mexico, known in California only from

the south in Inyo, Riverside, and San Bernardino counties [Figure 92]

4. S. lusoria (Cresson)
NUMBER 610 23

1. Sphecodogastra aberrans (Crawford), new combination

FIGURES 51-58,66-71

Halictus aberrans Crawford, 1903:336 [females].—Cockercll, 1906b:427

[floral records].—1907a:242 [key].—1907b:119 [contrasted with H. galpinsiae].—Crawford, 1907:186, 188 [key; compared with H. galpinsiae].—

Graenicher, 1911:224 [compared with H. galpinsiae].—Stevens, 1951:61

[association with Gaura coccinea and other plants].

Halictus (Evylaeus) aberrans.—Stevens, 1920:36 [taxonomy; compared with

H. galpinsiae; locality and floral records].

Lasioglossum (Sphecodogastra) aberrans.—Michener, 1951:1111 [catalog].

Lasioglossum (Evylaeus) abberans.—Linsley and MacSwain, 1962:45 [lapsus

calami].

Evylaeus aberrans.—Moldenke and Neff, 1974:60 [floral records].—Bohart

and Youssef, 1976:186 [compared with E. galpinsiae].—Hurd, 1979:1958

[catalog].—Hurd et al., 1980:66 [association with Helianthus].—Moure and

Hurd, 1987:68 [catalog].

Lasioglossum aberrans.—Poole, 1996:431 [checklist].

TYPE MATERIAL.—Crawford based his description of Halictus aberrans on a syntype series of three females with the following data: "Sioux Co., Nebr., June 3, on Symphoricarpos;

Crawford, Nebr., July 28, on Cleome; Manitou, Colorado."

None of these specimens could be located, and they are presumed to be lost. Because of the long-standing confusion between this species and Sphecodogastra lusoria (see "Species

Descriptions and Synonymies" in systematic history section) I

am designating a neotype that was also collected in Sioux

County, Nebraska. The specimen is deposited in the entomology collection of the University of Nebraska and is labeled

"Sioux Co[unty] Neb.[raska] May/L. Bruner Collector/NEOTYPE Halictus aberrans Crawford designated by] R.J. McGinley" [red label]. It is missing the right antenna, nine distal

flagellomeres of the left antenna, and two distal tarsomeres of

the right middle leg, and its abdomen is glued on the left side to

the second specimen label. Despite this damage, the specimen

is otherwise in excellent condition and shows all diagnostic

features needed to differentiate it from S. lusoria and other

known Sphecodogastra species.

DISTRIBUTION (Figure 51).—Sphecodogastra aberrans has

the most northern known distribution of the genus, having

been collected from southern Alberta, Manitoba, and

Saskatchewan. Sphecodogastra lusoria is the only other species known from central Canada, at Cardston and Medicine

Hat, Alberta (Figure 92). Both species appear to be widely
24 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

FIGURE 51.—Distribution of Sphecodogastra aberrans.

sympatric throughout the Great Basin and the intermountain

region of the central and western United States, with S. aberrans extending south only to southern California (Riverside

County), central Arizona, and northern New Mexico. Sphecodogastra lusoria in addition includes a more southeastern

distribution, east to Nebraska, Kansas, Oklahoma, and Texas

and south through southern Arizona, New Mexico, and central

Mexico (Hidalgo state). Bohart and Youssef (1976:186) noted

that "in Colorado, collection records indicate that the two species may fly together," which is known from other collections.

They also indicated that S. lusoria "nests in open, sandy areas

with typically xeric surroundings," whereas Evylaeus aberrans

"is generally found in more mesic and highland surroundings,

on brush-covered slopes or open woodlands."

The distribution records for 5. aberrans listed by Moure and

Hurd (1987) are understandably misleading because they include records for S. lusoria; however, they listed a record from

Coahuila, Mexico, presumably associated with S. lusoria. I

have not seen specimens of this species from this state, but its

occurrence there is not unexpected. They also listed a record

for one or the other of these species from Wisconsin, which

would seem to be a dubious but possible extension for either

species—most likely for S. aberrans.

DIAGNOSIS.—The absence of short, adpressed pleural hairs

differentiates the females of S. aberrans, S. antiochensis, and S.

oenotherae from all other known Sphecodogastra females. The

coarsely rugo-striate dorsal propodeal surface of 5. oenotherae

easily separates it from the other two species. The punctation

on the supraclypeal area of S. aberrans is only moderately

dense, with most punctures separated by twice their diameters

(Figure 24); this area is densely punctate in S. antiochensis,

with punctures separated by less than the width of their diameters. The femoral scopa of S. aberrans is sharply delimited as

in other species of the genus (e.g., Figure 26), whereas these

hairs in S. antiochensis are relatively diffused in pattern (i.e.,

not forming a distinctly sharp row of apically curved hairs; Figure 25). Consideration of distribution is probably the easiest

way to separate these two species (Figures 51, 92) because 5.

antiochensis is found only in the area of Antioch, California,

where S. aberrans is not known to occur.

Males of S. aberrans can be distinguished from those of

other Sphecodogastra by their distinctive sternal vestiture (Figure 37); the fringe hairs on the posterior margins of sterna 4

and 5 are conspicuously developed and differentiated from vestiture on other sternal margins. Other distinctive characters are

presented in couplet four of "Key to Specodogastra Males."

DESCRIPTION.—FEMALE: (1) Length 7.8-9.4 mm (mean =

8.4, n = 5); (2) wing length 2.4-2.7 mm (mean = 2.6, n = 5);

(3) abdominal width 2.3-2.7 mm (mean = 2.6, n = 5).

Structure: (4) Head moderately elongate (Figure 52);

length/width ratio 1.00-1.03 (mean = 1.01, n = 5). (5) Gena, at

midpoint, subequal to or slightly exceeding compound eye in

width. (9) Clypeus projecting approximately 0.69 times its

length below lower margin of eyes; (11) clypeal surface without median longitudinal sulcation. (13) Ocular-ocellar distance

greater than distance between lateral ocellus and hind margin

of vertex (ocular-ocellar space approximately 2.0 times lateral

ocellar diameter); (14) distance between lateral ocelli slightly

exceeding ocular-ocellar distance. (16) Inner margins of compound eyes converging below. (21) Scape reaching top of vertex; (22) pedicel subequal in length to flagellomere 1. (30)

Mandible moderate in length, extending to opposing clypeal

angle.

(40) Dorsal surface of propodeum about 0.60 times the

length of scutellum and approximately 1.25 times the length of

metanotum; (44) lateral propodeal carinae well developed, extending to dorsal propodeal surface. (45) Inner hind tibial spur

with 4 or 5 moderately elongate teeth (Figure 58).

(46) Lateral edge of metasomal T2 broadly rounded anteriorly, becoming straight to faintly sinuate posteriorly.

Sculpture: (51) Supraclypeal area granulate, (52) punctation moderately dense, most punctures separated by 1-2 times

the width of their diameters (Figure 24). (53) Clypeus tessellate

basally; (54) subapical area with contiguous, large punctures.

(55) Hypostoma striolate.

(56) Mesoscutum somewhat dull, surface tessellate over anterior half or more, (57) punctation as in Figure 57, punctures

separated by 1-3 times their diameters. (63) Dorsal surface of

propodeum rugulose over basal three-fourths (Figure 56), (64)

surface alveolated. (65) Tl shiny and polished, (66) punctation

fine, moderately dense, most punctures separated by their diameters.
NUMBER 610 25

FIGURES 52-57.—Sphecodogastra aberrans: 52, female head; 53, male head; 54, female labrum; 55, male

labrum; 56, female propodeum; 57, female mesoscutum.

Coloration: (67) Abdomen dark brown. (69) Flagellum

brown. (70) Tegula light brown. (71) Wing membrane pale yellowish brown; veins and stigma amber. (72) Legs dark brown.

Vestiture: (74) Head hairs pale yellowish brown. (75) Pubescence on thorax pale yellowish brown; (76) mesoscutal

hairs elongate, approximately 1.5 times median ocellar diameter; mesoscutum and pleuron without short suberect or adpressed hairs. (81) Basal hair bands on T2-T4 present, covering basal one-fourth of tergal surface; moderately developed

apical hair bands present on T3 and T4.

MALE: As described for female except as follows: (1)

Length 7.7-9.3 mm (mean = 8.7, n = 5); (2) wing length

2.0-2.2 mm (mean = 2.2, n = 5); (3) abdominal width 1.8-2.0

mm (mean = 1.9, n = 5). (4) Head (Figure 53) length/width ratio 1.10-1.12 (mean = 1.11, n = 5). (23) Flagellomere 2 approximately 1.8 times length of flagellomere 1. (37) Median mesoscutal line impressed. (70) Tegula yellow-translucent. (73)

Short, adpressed hairs on face extending dorsad to just below

median ocellus. (76) Mesoscutum with moderately elongate

hairs (short, adpressed hairs absent); pleuron without short,
26

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

FIGURES 58-65.—Sphecodogastra female inner tibial spurs: 58, S. aberrans; 59, S. antiochensis; 60, S.

danforthi; 61,5. lusoria; 62,5. oenotherae; 63, S. potosi; 64,5. noctivaga; 65, S. texana.

adpressed hairs. (82, 83) Unlike other Sphecodogastra, elongate vestiture on S4 and S5 confined to apical margin (Figure

37; vestiture distributed over entire sternal surface in other species, e.g., Figure 38); (83a) central depression of S6 extremely

well developed (Figure 66).

Terminalia (Figures 67-70): (84) S7 lateral arms slender

(Figure 70); (85) S8 moderately developed (Figure 70); apex

of median process slightly concave; (89) retrorse membranous

lobe narrow, becoming slightly broader basally. Unlike other

Sphecodogastra, penis valve sharply angulate in lateral view,

FIGURE 66.—Sphecodogastra aberrans, male S6.

dorsal surface concave, and sharply carinate laterally (Figure

67).

FLIGHT RECORDS (Figure 71).—Females of 5. aberrans were

collected from May through October, with most records (98%)

from June through August. Males appear to be active somewhat later, with an apparent peak during July.

REMARKS.—See "Species Descriptions and Synonymies" in

systematic history section.

SPECIMENS EXAMINED.—426 (327 females, 99 males).

CANADA, ALBERTA: Kipp; Lethbridge; Medicine Hat;

Medicine Hat, 25 mi NW; Morrin; Nobleford. MANITOBA:

Treesbank. SASKATCHEWAN: Elrose; Saskatoon; Val Marie.

UNITED STATES. ARIZONA: Coconino Co.: Flagstaff; Oak

Creek (W fork, 16 mi SW Flagstaff); Williams; Yavapai Co.:

Prescott; county /coordinates unknown: Oayson. CALIFORNIA: Plumas Co.: Vinton, 5.5 mi S; Riverside Co.: Indio (Pine

Flats Camp). COLORADO: Archuleta Co.: Piedra; Boulder

Co.: Altona; Altona, 3 mi SW; Boulder; Copeland Park; Jim

Creek (near Boulder); Lyons, 9 mi NW; Ward; Clark Co.: Lake

George; Clear Creek Co.: Echo Lake (Mt. Evans); Denver Co.:

Denver; Elbert Co.: Elbert; El Paso Co.: Colorado Springs;

Manitou [male specimen; not part of type series]; Gunnison

Co.: Mt. Princeton Hot Springs; Sapinero, 4 mi E; Huerfano

Co.: Cuchara; Jefferson Co.: Crossons; Golden; Larimer Co.:

Cameron Pass; Hewlett Gulch (near Poudre Park); Montrose

Co.: Buckeye Reservation; Cimarron, 1.2 mi W; Park Co.:

Wilkerson Pass; Routt Co.: Oak Creek (20 mi S Steamboat
NUMBER 610

27

FIGURES 67-70.—Sphecodogastra aberrans, male terminalia: 67, genital capsule, ventral view; 68, genital capsule, dorsal view; 69, right gonostylus, posterior view; 70, S7 and S8, ventral view.

140-

120-

(0

8

E

I

CO

•sber

100

80

60

40-

20-

0 r i i i i i i i T r

Jan Feb Mar Apr May Jun Jul Aug Sep Oct Nov Dec

FIGURE 71.—Sphecodogastra aberrans flight records, by month of collection.
28 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

Springs); Teller Co.: Florissant; county /coordinates unknown:

Mishawauka. IDAHO: Elmore Co.: Mountain Home; Twin

Falls Co.: Twin Falls, 18 mi S. MONTANA: Dawson Co.:

Glendive; Gallatin Co.: Bozeman; Missoula Co.: Missoula.

NEBRASKA: Sioux Co. NEW MEXICO: Sandoval Co.: Jemez

Springs. NORTH DAKOTA: Barnes Co.: Valley City; Bottineau

Co.: Bottineau; Bowman Co.: Gascoyne; Cass Co.: Fargo;

Dickey Co.: Oakes; Golden Valley Co.: Beach; Sentinel Butte;

Kidder Co.: Tappen; La Moure Co.: Kulm; LaMoure; McLean

Co.: Garrison; Washburn; Pierce Co.: Rugby; Ransom Co.:

Lisbon; Slope Co.: Marmarth; Stark Co.: Dickinson; Stutsman

Co.: Jamestown; Kensal; Ward Co.: Lone Tree. OREGON:

Grant Co.: Long Creek, 3.6 mi S; Homey Co.: Narrows, 20 mi

S; Sherman Co.: Maryhill Ferry (3 mi E Briggs). UTAH:

Cache Co.: Logan (Green Canyon); Daggett Co.: Palisade

Park Camp; Sheep Creek Canyon; Sanpete Co.: Fairview; Tooele Co.: Tooele; Utah Co.: Provo; Washington Co.: Lower

Deep Creek; Virgin; Pinto; Upper Deep Creek; Zion National

Park. WASHINGTON: Adams Co.: Ritzville, 4 mi N. WYOMING: Albany Co.: Laramie; Laramie, 2 mi S; Laramie, 8 mi

SSE; Converse Co.: Glenrock; Fremont Co.: Lander, 8.5 mi

SW (Sinks Canyon); Riverton; Sheridan Co.: Sheridan.

2. Sphecodogastra antiochensis, new species

FIGURES 59,72-82

As Lasioglossum {Sphecodogastra) aberrans (Crawford).—Turner, 1966

[foraging activity, host plant biology, mating biology, predation, parasitism,

nest site].

TYPE MATERIAL.—The holotype female, from the collection

of the Bee Biology and Systematics Laboratory, Utah State

University, has been deposited in the National Museum of Natural History, Smithsonian Institution. It is in excellent condition and is labeled "Antioch Calif.[ornia] IV[April]-4-[19]39

[handwritten]/GE Bohart Collector/HOLOTYPE Sphecodogastra antiochensis R.J. McGinley" [red label]. Paratypes are listed

in "Specimens Examined," below.

ETYMOLOGY.—The specific name was a label manuscript

name used by George E. Bohart, who along with P.H. Timberlake was among the first workers to recognize this species as

being distinct. As the name implies, this species is known

only from the vicinity of Antioch, Contra Costa County, California.

DISTRIBUTION (Figure 92).—Sphecodogastra antiochensis

is a geographically isolated species of this genus. It is known

to occur only in the sand dune habitat of Antioch, Contra Costa

County, California. Its host plant is a similarly isolated subspecies of primrose, Oenothera deltoides howellii (Klein, 1970).

The nearest occurrences of another Sphecodogastra species I

am aware of are two separate collections of S. lusoria from

Delhi and Livingston, Merced County, California, approximately 70 miles southeast of Antioch (Figure 92).

DIAGNOSIS.—Females: see "Diagnosis" for S. aberrans.

Males: see "Diagnosis" for 5". lusoria.

DESCRIPTION.—FEMALE: (1) Length 8.2-9.5 mm (mean =

8.8, n = 5); (2) wing length 2.6-3.0 mm (mean = 2.8, n = 5); (3)

abdominal width 2.6-2.9 mm (mean = 2.7, n = 5).

Structure: (4) Head moderately elongate (Figure 72);

length/width ratio 0.95-1.00 (mean = 0.98, n = 5). (5) Gena, at

midpoint, slightly exceeding width of compound eye. (9)

Clypeus projecting approximately 0.63 times its length below

lower margin of eyes; (11) clypeal surface without median longitudinal sulcation. (13) Ocular-ocellar distance exceeding

distance between lateral ocellus and hind margin of vertex (ocular-ocellar space approximately 1.5 times lateral ocellar diameter); (14) distance between lateral ocelli slightly exceeding

ocular-ocellar distance. (16) Inner margins of compound eyes

converging below. (21) Scape reaching top of vertex; (22)

pedicel subequal in length to flagellomere 1. (30) Mandible

moderate in length, reaching opposing clypeal angle.

(40) Dorsal surface of propodeum about 0.59 times the

length of scutellum and approximately 1.13 times the length of

metanotum; (44) lateral propodeal carinae well developed, extending to dorsal propodeal surface. (45) Inner hind tibial spur

with 5-7 teeth, more numerous and somewhat longer than

those of other Sphecodogastra species (Figure 59).

(46) Lateral edge of metasomal T2 rounded anteriorly, becoming straight to faintly sinuate posteriorly.

Sculpture: (51) Supraclypeal area mostly polished, tessellate only near lateral margins, (52) very densely punctate, most

punctures separated by less then the width of their diameters.

(53) Clypeus polished; (54) punctation extremely sparse, apical punctures only slightly larger than basal ones, most separated by at least three times their diameters. (55) Hypostoma

striolate throughout.

(56) Mesoscutum somewhat dull, tessellate on anterior half,

(57) punctation as in Figure 77, most punctures separated by

1-2 times their diameters. (63) Dorsal surface of propodeum

ruguloso-striolate, nearly to posterior edge (Figure 27), (64)

surface alveolated. (65) Tl shiny and polished, (66) punctation

fine, moderately sparse, punctures separated by 1-2 times their

diameters.

Coloration: (67) Abdomen dark brown. (69) Flagellum

dark brown. (70) Tegula brown to light brown. (71) Wing

membrane pale yellowish brown; veins and stigma amber. (72)

Legs dark brown.

Vestiture: (74) Hairs on head pale yellowish brown. (75)

Pubescence on thorax pale yellowish white; (76) mesoscutal

hairs elongate, length approximately 1.5 times median ocellar

diameter; mesoscutum and pleuron without short, suberect or

adpressed hairs. (81) Basal hair bands on T2-T4 present, covering basal one-fourth of tergal surface; moderately developed

apical hair bands on T3 and T4.

MALE: As described for female except as follows: (1)

Length 8.0-9.3 mm (mean = 8.5, n = 5); (2) wing length

2.2-2.6 mm (mean = 2.4, n = 5); (3) abdominal width 1.7-2.2

mm (mean = 2.0, n = 5). (4) Head (Figure 73) length/width ratio 1.08-1.11 (mean = 1.09, n = 5). (23) Flagellomere 2 ap-
NUMBER 610 29

FIGURES 72-77.—Sphecodogastra antiochensis: 72, female head; 73, male head; 74, female labrum; 75, male

labrum; 76, male flagellum; 77, female mesoscutum.

proximately 2.0 times length of flagellomere 1. (37) Median

mesoscutal line impressed. (70) Tegula yellow-translucent to

brown. (73) Short, adpressed hairs on face extending dorsad to

ocellar area, nearly enclosing median ocellus. (76) Mesoscutum with moderately elongate hairs and extensive layer of

short, adpressed hairs; pleuron with short, adpressed hairs.

Terminalia (Figures 78-81): (84) S7 lateral arms moderately well developed; (85) S8 moderately developed; apex of

median process rounded; (89) retrorse membranous lobe narrow.

FLIGHT RECORDS (Figure 82).—Females of 5. antiochensis

examined in this study were collected from March through August, with most records (67%) collected during May. Male collections ranged from May through October, with an overwhelming number (96%) taken in June.

Turner (1966) reported the earliest records of female flight

activity as 15 March and 5 April. During his 1964-1965 study,

females were first active usually in late April. The flight period

of this species normally extends through August; however, in
30

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

FIGURES 78-81.—Sphecodogastra antiochensis, male terminalia: 78, genital capsule, ventral view; 79, genital

capsule, dorsal view; 80, right gonostylus, posterior view; 81, S7 and S8, ventral view.

350

300

M 250

3

S 200

150

2

z

100

50

Jan Feb Mar Apr May Jun Jul Aug Sep Oct Nov Dec

FIGURE 82.—Sphecodogastra antiochensis flight records, by month of collection.
NUMBER 610 31

1965, unusual rainfall prolonged the season, with females and

males being observed as late as 19 and 22 September, respectively. Turner indicated that Sphecodogastra antiochensis apparently produced two generations each year.

REMARKS.—Sphecodogastra antiochensis is known exclusively from the bluffs of Antioch, Contra Costa County, California, as is its host plant, Oenothera deltoides howellii (Klein,

1970). This area was recognized in 1980 as the Antioch Dunes

National Wildlife Refuge, largely for the protection of the endangered Lange's metalmark butterfly (Apodemia mormo

langei) and its host plant, Eriogonum nudum (Caterino, 1997).

Caterino indicated that under protection and proper dune management the situation for the butterfly and its host plant looked

relatively good.

Jerry Powell, who has long been involved with conservation

efforts on behalf of the Antioch Dunes (e.g., Powell, 1978),

added (Powell, pers. comm., 1996) the following:

The last time I checked on Sphecodogastra at Antioch was June 1982, and it

was present then (22 individuals, 0630-0830). Although I have visited a few

times in early A.M. in recent years to look for Apodemia larvae, I didn't look for

Sphecodogastra. Since that time there has been a concerted effort to plant

Oenothera, even including trucking in and contouring whole new sandhills,

with considerable success. The plant is much more abundant now than in the

lowest ebb, when weediness was choking out seedling establishment. The

plants set seed, so I assume the bee population has increased. I don't know that

it will be of any advantage to propose the population as endangered because the

property is already in Federal hands and because they target the Oenothera as

an endangered species in their recovery plans.

I agree with Powell's opinion that it is probably unnecessary

to propose endangered species status for S. antiochensis, but I

hope this unique Oenothera—Sphecodogastra relationship will

be monitored in the future.

SPECIMENS EXAMINED.—574 (233 females, 341 males); 573

paratypes are designated and so labeled. They are deposited in

the following collections: KU, NMNH, UCB, UCR, UIM,

UNL, and USU.

3. Sphecodogastra danforthi, new species

FIGURES 60,83-91

TYPE MATERIAL.—The female holotype of S. danforthi is

deposited in the NMNH collection. It is in excellent condition

and is labeled "K07461/TEXAS: Winkler Co. Rd. 404, 8 mi E

Hwy 18 10-IX-1989 J.L. Neff/on flowers of Calylophus

Hartwegii/HOLOTYPE Sphecodogastra danforthi des. R.J.

McGinley" [red label]. Paratypes (33 females, 5 males) are

designated and listed in "Specimens Examined," below.

ETYMOLOGY.—I was originally aware of this undescribed

species from one female in very poor condition collected in

1959, 9.5 miles south of Monahans, Texas, but I was reluctant

to describe it as new. Subsequently, Bryan N. Danforth, for

whom this species is named, sent me a freshly collected sample

of 11 females and two males from 10 km northeast of Monahans, which confirmed my initial opinion that the one previously known specimen represented a new species. Later, addiFlGURE 83.—Distribution of Sphecodogastra danforthi (squares) and S. potosi

(bullets).

tional specimens of this rarely collected species were sent to

me by J.L. Neff, an important contribution that I would like to

acknowledge.

DISTRIBUTION (Figure 83).—Sphecodogastra danforthi is

presently only known from Ward and Winkler counties, Texas.

It is definitely sympatric with S. lusoria, S. noctivaga, and S.

texana. From the UCB collection I have examined specimens

of all four species collected by D.P. Gregory on 11 May 1959,

9.5 miles south of Monahans, from the flowers of Calylophus

hartwegii (all specimens had Onagraceae pollen in their scopae).

DIAGNOSIS.—The dark abdomen and conspicuously elongate and projecting mandibles, which lack defined subapical

teeth (Figure 86), easily separate S. danforthi females from

other known congenerics. Females of S. potosi are similar in

having elongate mandibles, but these are broadly rounded laterally, weakly projecting, and have well-defined subapical teeth

(Figure 18). The males of 5. danforthi and 5. potosi are difficult

to differentiate, a situation parallel to S. noctivaga and S. texana where the females are easily separated on the basis of mandibular morphology but the respective males are nearly identical. The darkly pigmented abdomens of male S. danforthi and

S. potosi in combination with the finely rugulose dorsal propodeal surface and "normal" antennal flagella (lacking distinctive sensillar patterns) will differentiate them from other congenerics. The head of male S. potosi (Figure 130) appears to be

broader than that of 5. danforthi (Figure 85), but this is not obviously reflected in head length/width ratios (1.04-1.08 for the

former and 1.10-1.14 for the latter).
32

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

FIGURES 84-87.—Sphecodogastra danforthi: 84, female head; 85, male head; 86, female mandibles; 87, female

propodeum.

DESCRIPTION.—FEMALE: (1) Length 6.7-7.7 mm (mean =

7.2, n = 5); (2) wing length 2.0-2.2 mm (mean = 2.1, n = 5); (3)

abdominal width 2.0-2.3 mm (mean = 2.2, n = 5).

Structure: (4) Head short, rounded (Figure 84); length/

width ratio 0.91-0.95 (mean = 0.95, n = 5). (5) Gena, at midpoint, exceeding compound eye in width. (9) Clypeus projecting approximately 0.77 times its length below lower margin of

eyes; (11) clypeal surface without median longitudinal sulcation. (13) Ocular-ocellar distance slightly shorter than distance

between lateral ocellus and hind margin of vertex (ocularocellar space approximately 1.5 times lateral ocellar diameter);

(14) distance between lateral ocelli exceeding ocular-ocellar

distance. (16) Inner margins of compound eyes only slightly

converging below, nearly parallel. (21) Scape reaching top of

vertex; (22) pedicel nearly subequal in length to flagellomere

1. (30) Mandible conspicuously elongate, strongly projecting

beyond clypeus (Figure 86); (31) unlike other Sphecodogastra,

subapical tooth broadly rounded, not defined.

(40) Dorsal surface of propodeum about 0.71 times the

length of scutellum and approximately 1.36 times the length of

metanotum; (44) lateral propodeal carinae well developed, extending to dorsal propodeal surface. (45) Inner hind tibial spur

usually with four moderately elongate teeth (Figure 60).

(46) Lateral edge of metasomal T2 straight.

Sculpture: (51) Supraclypeal area finely granulate, (52)

punctation moderately dense, most punctures separated by two

times the width of their diameters. (53) Clypeus weakly tessellate basally, mostly polished and shiny; (54) subapical area

with a few large, poorly defined punctures. (55) Hypostoma

smooth, nonstriolate.

(56) Mesoscutum somewhat dull, surface tessellate over anterior half or more; (57) mesoscutal punctures separated by

their diameters or slightly less. (63) Dorsal surface of propodeum ruguloso-striolate throughout (Figure 87), (64) surface

shiny, nonalveolate. (65) Tl shiny and polished, (66) punctation extremely fine and sparse, most punctures separated by

2-5 times their diameters.

Coloration: (67) Abdomen dark brown. (69) Flagellum

brown. (70) Tegula light brown. (71) Wing membrane pale yellowish brown; veins and stigma amber. (72) Legs dark brown.

Vestiture: (74) Head hairs yellowish white-brown. (75) Pubescence on thorax pale yellowish brown; (76) mesoscutal

hairs moderately short, only slightly exceeding diameter of me-
NUMBER 610 33

FIGURES 88-91.—Sphecodogastra danforthi, male terminalia: 88, genital capsule, ventral view; 89, genital capsule, dorsal view; 90, right gonostylus, posterior view; 91, S7 and S8, ventral view.

dian ocellus; mesoscutum and pleuron without short suberect

or adpressed hairs. (81) Basal hair bands on T2-T4 absent; apical hair bands on T3 and T4 absent.

MALE: As described for female except as follows: (1)

length 6.4-7.7 mm (mean = 7.0, n = 5); (2) wing length 1.4-1.8

mm (mean = 1.8, n = 5); (3) abdominal width 1.4-1.8 mm

(mean = 1.7, n = 5). (4) Head (Figure 85) length/width ratio

1.10-1.14 (mean = 1.12, n = 5). (23) Flagellomere 2 approximately 1.3 times length of flagellomere 1. (37) Median mesoscutal line not impressed. (70) Tegula yellow-translucent. (73)

Short, adpressed hairs on face extending dorsad to ocellar area,

not on vertex. (76) Mesoscutal hairs mostly very short, adpressed (longer hairs relatively inconspicuous); pleuron with

very short, adpressed to suberect hairs.

Terminalia (Figures 88-91): (84) S7 lateral arms very slender; (85) S8 short, somewhat reduced; apex of median process

rounded; (89) retrorse membranous lobe narrow, parallel sided,

curved laterad.

FLIGHT RECORDS.—Only three collections of S. danforthi

have been made. Females were taken in May, August, and September; males were taken in August and September.

REMARKS.—The elongate, blade-like mandibles of S. danforthi females (Figure 86) are similar to those of S. noctivaga

(Figure 13). The mandibles of the latter species differ by having small but defined subapical teeth. Elongate mandibles are

also found in S. potosi (Figure 18) and S. oenotherae, but in

these species they are broadly rounded, not straight and bladelike on their outer edges.

It is possible these elongate mandibles are important in harvesting onagraceous pollen. Knerer and MacKay (1969) reported that S. oenotherae females worked pollen free from the

anthers by using their mandibles. Sphecodogastra antiochensis, which have "normal" mandibles, apparently use them to

clamp themselves to Oenothera anthers. Once so anchored the

bees use both their front and middle legs to remove pollen and

transfer it to the scopae on the hind legs (Turner, 1966). Additional observations of pollen harvesting among Sphecodogastra species would be interesting, especially a comparison between the closely related S. noctivaga with elongate mandibles

(Figure 13) and S. texana with "normal" mandibles (Figure

14).

SPECIMENS EXAMINED.—39 (34 females, 5 males).

UNITED STATES, TEXAS: Ward Co.: Monahans, 9.5 mi S.,

11 May 1959, Oenothera hartwegii, D.P. Gregory (1 female;

UCB); Monahans, 10 km (6.2 mi) NE, 29 Aug 1986, B.N. Danforth, A.M. Simons (11 females; 2 males; K.U); Winkler Co.:
34 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

Road 404 (8 mi E Highway 18), 10 Sep 1989, Calylophus

hartwegii, J.L. Neff (22 females; 3 males; gift to NMNH; one

male and four female paratypes from this series have been deposited in the CTMI collection).

4. Sphecodogastra lusoria (Cresson), new combination

FIGURES 61,92-104

Halictus lusorius Cresson, 1872:252 [two females].—Dalla Torre, 1896:69

[catalog].—Cockerell, 1897:166 [one female, west fork of Gila River, New

Mexico, July 16].—Crawford, 1907:186 [noted that lusorius was excluded

from key].

Halictus amicus. van a, Cockerell, 1901:126 [female].

Halictus galpinsiae Cockerell, 1903:342 [female].—1906a:294 [recorded from

Pecos, Las Vegas, New Mexico].—1907b: 119 [locality, floral records; contrasted with H. aberrans].—1919:288 [recorded from Rocky Mountain National Park, just above timberiine].—Crawford, 1907:186-188 [key].—

Graenicher, 1911:224 [compared with H. aberrans]. [New synonymy.]

Halictus gelidus Vachal, 1904:481 [male].—Crawford, 1907:188 [synonymy].

Lasioglossum (Sphecodogastra) lusorium.—Michener, 1951:1111 [catalog].

Lasioglossum (Evylaeus) galpinsiae.—Linsley and MacSwain, 1962:45 [taxonomy].—Linsley et al., 1963:19, 42, 43 [pollen records from Gaura and

Oenothera].—Gregory, 1964:394 [floral records].—Linsley et al., 1973:38,

39 [pollen records from Oenothera].

Lasioglossum (Evylaeus) lusorium.—Linsley and MacSwain, 1962:45 [taxonomy].

Evylaeus galpinsiae.—Bohart and Youssef, 1976:185-234 [nesting biology,

life history, taxonomy].

Evylaeus lusorius.—Hurd, 1979:1960 [catalog].—Moure and Hurd, 1987:75

[catalog].

Lasioglossum lusorius.—Poole, 1996:578 [checklist].

TYPE MATERIAL.—The female lectotype of Halictus lusorius, herein designated, is deposited in the Academy of Natural

Sciences of Philadelphia. The specimen is in good condition

but is missing the three distal tarsomeres of the left hind leg,

and the hairs over most of the body are slightly soiled and matted. Label data are as follows: "Tex.[Texas]/TYPE No. 2115

[red label]/Halictus lusorius 667 Cr [handwritten on folded

label]/ANSP [yellow label]/LECTOTYPE Halictus lusorius

Cresson des. [designated by] McGinley [red label]."

The female holotype of Halictus galpinsiae, in the collection

of the University of Colorado, Boulder, is in excellent condition and is labeled "Pecos. N.M.[New Mexico], June 22.

(Ckll.)/Collecting pollen from Galpinsia fendleri [= Calylophus hartwegii fendleri], 7.30 P.M./Halictus galpinsiae Ckll.

TYPE" [handwritten with red-inked borders].

Vachal's male holotype of Halictus gelidus is in the collection of the Naturhistorisches Museum, Wien (Vienna, Austria).

The specimen is in good condition but is missing the five distal

flagellomeres of the right antenna; the terminalia have been extruded, which apparently resulted in soiling of the pleurae, portion of the left scutum, and left gena. Label data are as follows:

"Morrison Colorado 1879. I./gelidus Vachal det.[determined]

J. Vachal/H. gelidus Vach.[Vachal]/LASlOGLOSSUM o Evylaeus

gelidum (Vach. 1906) Holotypus, Unikum det. A.W. Ebmer

1980" [red-inked right border].

DISTRIBUTION (Figure 92).—See "Distribution" for S. aberrans.

FIGURE 92.—Distribution of Sphecodogastra lusoria (bullets) and S. antiochensis (circled dot).

DIAGNOSIS.—The short, adpressed pubescence on the

pleura, beneath longer hairs will distinguish the females of

Sphecodogastra lusoria and S. potosi from other Sphecodogastra having dark abdomens. The head of female S. lusoria is

only moderately broad (Figure 93) and has short mandibles that

at most extend as far as the opposing clypeal angle (e.g., Figure

14). Sphecodogastra potosi has a conspicuously broad head

(Figure 18) and moderately elongate mandibles that slightly

exceed the opposing clypeal angles.

Males of both S. lusoria and S. antiochensis have distinctive

antennal sensillar patterns: flagellomeres are primarily covered

by plate sensillae, with peg sensillae found only on the anterior

surface and further limited to areas immediately adjacent to

flagellomere margins (Figures 42,76). This pattern can be seen

through the light microscope, where the areas with peg sensillae appear lighter or darker than the surrounding surfaces. In

addition, these are the only males having short, adpressed hairs

extending from the clypeal base to the vertex (in other species,

the short, adpressed hairs do not reach the vertex, extending

only slightly above the antennae). The only detected morphological differences between the males of these two species are
NUMBER 610 35

FIGURES 93-98.—Sphecodogastra lusoria: 93, female head; 94, male head; 95, female labrum; 96, male labrum;

97, female propodeum; 98, female mesoscutum.

that S. antiochensis are larger (body length mean = 8.5 mm versus 7.2 mm for S. lusoria) and have slightly denser punctation

on tergum 1 than do the males of S. lusoria (Figures 49, 50).

The easiest way to differentiate these species is to consider distribution data; the two are not known to be sympatric—S. antiochensis is found only in the Antioch area of California (Figure

92) where S. lusoria is not known to occur.

DESCRIPTION.—FEMALE: (1) Length 7.5-9.1 mm (mean =

8.4, n = 5); (2) wing length 2.4-2.7 mm (mean = 2.6, n = 5); (3)

abdominal width 2.4-2.7 mm (mean = 2.5, n = 5).

Structure: (4) Head moderately rounded (Figure 93);

length/width ratio 0.99-1.40 (mean = 1.07, n - 5). (5) Gena, at

midpoint, slightly exceeded by width of compound eye. (9)

Clypeus projecting approximately 0.70 times its length below

lower margin of eyes; (11) clypeal surface without median longitudinal sulcation. (13) Ocular-ocellar distance slightly

greater than distance between lateral ocellus and hind margin

of vertex (ocular-ocellar space approximately 1.5 times lateral

ocellar diameter); (14) distance between lateral ocelli slightly

exceeding ocular-ocellar distance. (16) Inner margins of compound eyes converging below. (21) Scape reaching top of vertex; (22) pedicel subequal in length to flagellomere 1. (30)

Mandible moderate in length, extending to opposing clypeal

angle.
36 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

(40) Dorsal surface of propodeum about 0.78 times the

length of scutellum and approximately 1.5 times the length of

metanotum; (44) lateral propodeal carinae well developed over

three-fourths of propodeum, often reaching dorsal propodeal

surface (weakly developed in some specimens). (45) Inner hind

tibial spur with 4 or 5 moderately elongate teeth (Figure 61).

(46) Lateral edge of metasomal T2 broadly rounded anteriorly, becoming straight to faintly sinuate posteriorly.

Sculpture: (51) Supraclypeal area polished, (52) sparsely

punctate, many punctures separated by 3-4 times their diameters (Figure 23). (53) Clypeus mostly polished (sometimes with

obscure tessellation basally), (54) with large, widely spaced

punctures over apical half. (55) Hypostoma striolate posteriorly, becoming smooth and shiny on anterior half.

(56) Mesoscutum mostly shiny and polished; tessellation, if

present, confined to anterior one-fourth; (57) punctation as in

Figure 98, most punctures separated by 1-2 times their diameters. (63) Dorsal surface of propodeum rugulose (Figure 97),

(64) surface alveolated. (65) Tl shiny and polished; (66) punctation extremely fine, very sparse; punctures separated by 2-3

times their diameters.

Coloration: (67) Abdomen dark brown. (69) Flagellum

brown to amber ventrally. (70) Tegula light brown to yellowtranslucent. (71) Wing membrane hyaline; veins and stigma

pale yellowish amber. (72) Legs dark brown.

Vestiture: (74) Hairs on head white. (75) Pubescence on

thorax white; (76) mesoscutal hairs moderately short, subequal

in length to median ocellar diameter; mesoscutum and especially pleuron with short, adpressed hairs. (81) Basal hair bands

on T2-T4 present, covering basal one-fourth of tergal surface;

moderately developed apical hair bands on T3 and T4.

MALE: As described for female except as follows: (1)

length 6.7-8.0 mm (mean = 7.2, n = 5); (2) wing length 1.6-2.0

mm (mean = 1.8, n = 5); (3) abdominal width 1.3-1.7 mm

(mean = 1.5, n = 5). (4) Head (Figure 94) length/width ratio

1.07-1.10 (mean = 1.09, n = 5). (23) Flagellomere 2 approximately 2.0 times length of flagellomere 1. (37) Median mesoscutal line not impressed. (70) Tegula yellow-translucent. (73)

Short, adpressed hairs on face extending dorsad to vertex, enclosing median ocellus. (76) Mesoscutal hairs primarily short,

adpressed (longer hairs relatively inconspicuous); pleuron with

conspicuous short, suberect to adpressed hairs. (76a) Central

depression of S6 only moderately developed (Figure 99).

Terminalia (Figures 100-103): (84) S7 lateral arms moderately well developed; (85) S8 moderately developed; apex of

median process rounded; (89) retrorse membranous lobe narrow, becoming broader basally.

FLIGHT RECORDS (Figure 104).—Females of S. lusoria have

been collected from March through October, with most records

(87%) from May through July. Males appear to be active somewhat later than the females, with an apparent peak in June.

At Cornish, Utah, Bohart and Youssef (1976) reported flight

activity for this species to range from late May until SeptemFIGURE 99.—Sphecodogastra lusoria, male S6.

ber. In most years two full generations and a partial third were

produced. Bohart and Youssef indicated that the number of

generations produced was primarily dependent on the availability of Oenothera pollen.

REMARKS.—See "Species Descriptions and Synonymies" in

"Sphecodogastra Systematic History."

SPECIMENS EXAMINED.—918 (633 females, 285 males).

CANADA, ALBERTA: Cardston, 3 mi N; Medicine Hat.

MEXICO, CHIHUAHUA: Ahumada. DURANGO: Durango.

GUANAJUATO: Silao. HIDALGO: Tlaxcoapan, 5.6 mi E.

SAN LUIS POTOSI: San Luis Potosi, 16 mi E. ZACATECAS:

Fresnillo, 5 mi S; Fresnillo, 5-11 mi N; Juan Aldama.

UNITED STATES: ARIZONA: Cochise Co.: Douglas, 1 mi E;

Portal, 2 mi NE; Portal, 5 mi W (Southwestern Research Station); Coconino Co.: Cameron; Flagstaff, 10 mi N (San Francisco Mountains); Flagstaff, 20 mi N; Vermillion Cliffs; Maricopa Co.: Mesa, 1 mi N; Pima Co.: Continental; Yavapai Co.:

Granite Dells, 4 mi N. CALIFORNIA: lnyo Co.: Bartlett; Big

Pine, 3 mi N; Wyman Canyon (White Mountains); Merced Co.:

Delhi, 2.5 mi S; Livingston, 2 mi SW; Riverside Co.: Hopkins

Well, 2 mi W; Riverside; San Bernardino Co.: Apple Valley;

Kelso Dunes. COLORADO: Adams Co.: Berkeley; Utah Junction; Boulder Co.: Beaver Reservoir; Boulder; Longs Peak;

Nederland; Niwot Ridge (near Ward); Pinecliffe; Ward, 2 mi E;

Clark Co.: Lake George; Clear Creek Co.: Bear Creek Canyon;

Clear Creek; Echo Lake (Mt. Evans); Guanella Pass; Custer

Co.: Westcliff, 7.5 mi W; Denver Co.: Denver; Franktown, 5

mi SSE; Larkspur; El Paso Co.: Colorado Springs; Foster

Ranch; Gunnison Co.: Mt. Princeton Hot Springs, 1 mi E; Salida, 1.5 mi N (Highway 291); Gilpin Co.: Lump Gulch, near

Gilpin; Huerfano Co.: Walsenburg, 3 mi W; Jefferson Co.:

Coal Creek Canyon (7 mi S Crescent Village); Conifer; Larimer Co.: Buckhorn Creek; Estes Park; Ft. Collins; Glen Haven; Hewlett Gulch (near Poudre Park); White Rock; Moffat

Co.: Maybell; Montrose Co.: Cimarron, 1.2 mi W; Otero Co.:

La Junta; Park Co.: Pennsylvania Mountain; Wilkerson Pass;
NUMBER 610

37

FIGURES 100-103.—Sphecodogastra lusoria, male terminalia: 100, genital capsule, ventral view; 101, genital

capsule, dorsal view; 102, right gonostylus, posterior view; 103, S7 and S8, ventral view.

250

i i r

Jan Feb Mar Apr May Jun Jul Aug Sep Oct Nov Dec

FIGURE 104.—Sphecodogastra lusoria flight records, by month of collection.
38 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

Sedgwick Co.: Julesburg; Teller Co.: Florissant; Weld Co.:

Nunn (Pawnee Grassland Pasture); Roggen; coww/y/coordinates unknown: Pingee Park; Platts Gunon. IDAHO: Franklin

Co.: Preston, 5 mi W; St. Anthony; St. Anthony, 5.5 mi WNW;

Owyhee Co.: Oreana. KANSAS: Barber Co.: Medicine

Lodge; Medicine Lodge, 4 mi WNW; Ellis Co.: Hays; Finney

Co.: Garden City; Kalvesta, 15 mi W; Greeley Co.: Tribune;

Hamilton Co.: Syracuse; Logan Co.: Monument; Marshall Co.:

Blue Rapids; Pratt Co.: Sawyer, 7 mi N, 3 mi E; Reno Co.: Medora (sand dunes); Scott Co.: Scott City; Sheridan Co.: Quinter,

9 mi N; Sumner Co.: Caldwell, 9 mi N; Wallace Co.: Wallace;

Wichita Co.: Leoti. NEBRASKA: Chase Co.: Wauneta, 2 mi E;

Morrill Co.: Bridgeport; Perkins Co.: Grant, 3 mi N; Saline

Co.: Crete; Scotts Bluff Co.: Mitchell; Scottsbluff, 12 mi N;

Sioux Co.: Agate. NEVADA: Churchill Co.: Fallon, 14 mi S

and 10 mi W; Fallon, 25 mi SE (Sand Mountain); Eureka Co.:

Eureka, 70.8 mi N; Humboldt Co.: Winnemucca; Winnemucca,

11 mi N; Lander Co.: Austin; Austin, 9.5 mi W; Mineral Co.:

Hawthorne; Washoe Co.: Reno, 10 mi S on Highway 395; Wadsworth, 10-17 mi W. NEW MEXICO: Chaves Co.: Mesa, 2 mi

NW; DeBaca Co.: Ft. Sumner, 10 mi S; Ft. Sumner, 15 mi E;

Dona Ana Co.: Las Cruces; Eddy Co.: Carlsbad, 19 mi E;

Guadalupe Co.: Dilia, 1 mi S; Santa Rosa; McKinley Co.: Gallup, 19 mi N; Otero Co.: Silver Spring Canyon (Mescalero

Apache Indian Reservation); White Sands National Monument;

Quay Co.: Logan; Logan, 6.3 mi NE; Roosevelt Co.: Clovis, 10

mi SW; San Miguel Co.: Las Vegas; Pecos; Sandoval Co.: Jemez Springs; Santa Fe Co.: Santa Fe; Torrance Co.: Clines

Corners; Union Co.: Grenville; Valencia Co.: Pueblo

Laguna. NORTH DAKOTA: Golden Valley Co.: Beach; Richland Co.: Walcott, 11 mi W; Slope Co.: Marmarth. OKLAHOMA: Ellis Co.: Gage; Grady Co.: Rush Springs, 11 mi N;

Jackson Co.: Altus; Kingfisher Co.: Dover, 4 mi N; Woodward

Co.: Woodward; Woodward, 6.4 km (4 mi) NE. OREGON:

Harney Co.: Denio, 2.5 mi NE; Umatilla Co.: Hermiston;

Umatilla. SOUTH DAKOTA: Custer Co.: Custer; Jones Co.:

Murdo, 10 mi S. TEXAS: Blanco Co.: Davis Ranch (NW part

of county); Brewster Co.: Alpine; Big Bend National Park;

Marathon, 39 mi S; Brown Co.: Bangs; Comal Co.: New

Braunfels; Culberson Co.: Van Horn, 14.5 mi S; Fisher Co.:

Roby, 9 mi E; Gillespie Co.: Cherry Spring; Harper; Hall Co.:

Estelline, 3 mi W; Howard Co.: Big Spring; Jeff Davis Co.: Ft.

Davis, 10 mi W; Kerr Co.: Kerrville; Lamb Co.: Olton; La

Salle Co.: Cotulla; Encinal; Lee Co.: Fedor; Reeves Co.:

Toyahvale, 19 mi S; Shackelford Co.: Albany, 16 mi N; Terrell

Co.: Sanderson; Sanderson, 30 mi W; Ward Co.: Barstow, 3 mi

SW; Monahans, 6.7 mi W; Monahans, 9.5 mi S; Wilbarger Co.:

Vernon, 19 mi N. UTAH: Cache Co.: Cornish, 1.2 mi N; Carbon Co.: Greenriver, 15 mi S; Emery Co.: Calf Canyon (San

Rafael Swell); Gilson Butte (3.2^.0 mi N); Goblin Valley State

Preserve (2 mi N); Little Gilson Butte (2 mi E); San Rafael

Desert (Temple Mountain, 3 mi SSE); South Temple Wash (San

Rafael Reef); Wild Horse Creek (N of Goblin Valley); Garfield

Co.: Blind Trail Wash (SSE Notom); Lake Powell (4 mi NNW

Bullfrog); Ticaboo, 6 mi S; Grand Co.: Crescent Junction;

Moab; Kane Co.: Bullfrog; Millard Co.: Hatton; San Juan Co.:

Kane Springs (E of Natural Bridges National Monument);

Tooele Co.: Tooele, 3.6 mi S; Uintah Co.: Bonanza; Washington Co.: Leeds Canyon; Shivwits. WASHINGTON: Benton Co.:

Richland; Kittitas Co.: Yakima River (Morgan's Ferry).

WYOMING: Albany Co.: Laramie, 8 mi SE (Dirt Farm); Carbon

Co.: Kortes Dam; Converse Co.: Glenrock; Fremont Co.:

Shoshoni, 5 mi N; Shoshoni, 5 mi S; Niobrara Co.: Lusk;

Platte Co.: Chugwater; Glendo; Wheatland; Sweetwater Co.:

Bitter Creek, 25 mi S; Weston Co.: Upton, 4 mi N.

5. Sphecodogastra noctivaga (Linsley and MacSwain)

FIGURES 64,105-116

Lasioglossum (Sphecodogastra) noctivaga Linsley and MacSwain, 1962:46

[female].—Linsley et al., 1963:43 [locality records; floral association with

Oenothera hartwegii].—Gregory, 1964:394 [floral records].

Sphecodogastra noctivaga.—Hurd, 1979:1962 [catalog].—Moure and Hurd,

1987:85 [catalog].

Lasioglossum noctivaga.—Poole, 1996:611 [checklist].

TYPE MATERIAL.—The female holotype is the property of

the University of California, Berkeley, and is on loan deposit to

the California Academy of Sciences, San Francisco. The specimen is missing the four distal tarsomeres of the hind left leg but

otherwise is in excellent condition. It is labeled "Roosevelt

[Uintah County], Ut.fUtah], VI[June]-15-1956/J.L. Eastin

Collector/HOLOTYPE Lasioglossum (Sphecodogastra) noctivagum Linsley & MacSwain" [red label].

DISTRIBUTION (Figure 105).—Sphecodogastra noctivaga is

sympatric with the closely related and more widely distributed

S. texana (Figure 139) through portions of New Mexico, Nebraska, Oklahoma, and northwestern Texas. Unlike S. texana,

FIGURE 105.—Distribution of Sphecodogastra noctivaga.
NUMBER 610 39

5. noctivaga has been recorded from Utah. Sphecodogastra

texana is known from southeastern Montana, North Dakota,

east to Michigan, and south to Michoacan and Veracruz, Mexico, which are the most southern records known for the genus.

Moure and Hurd (1987) listed S. texana from Indiana. I have

not seen specimens from this state, but its occurrence there

would not be unexpected. Specimens of 5. noctivaga from Arizona, Kansas, and Oklahoma examined in this study apparently

represent new state records.

DIAGNOSIS.—The orange abdomen and conspicuously enlarged ocelli (Figures 11, 106) will distinguish both sexes of 5.

noctivaga and S. texana from other known species of Sphecodogastra. The elongate mandibles of female 5. noctivaga

(Figure 13) easily distinguish them from female 5. texana,

which have short ("normal") mandibles (Figure 14). The males

can be differentiated on the basis of the morphology of the genitalic retrorse lobes. These lobes are very broad and twisted

ventrally in 5. texana (Figures 31,32) but are narrow and flat in

5. noctivaga (Figure 33). Unfortunately, I currently know of no

external characters that will reliably distinguish the males of

these two species. The head of male S. noctivaga appears to be

slightly broader than that of S. texana. The head length/width

FIGURES 106-111.—Sphecodogastra noctivaga: 106, female head; 107, male head; 108, female labrum; 109,

male labrum; 110, female propodeum; HI, female mesoscutum.
40 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

ratio for S. noctivaga is 1.03-1.07 (mean = 1.05, n = 5) and for

S. texana is 1.00-1.03 (mean = 1.02, n = 5). The sternal vestiture, especially on sternum 5, appears to be more flocculent in

S. noctivaga than in S. texana. The only other detected difference appears to be in the coloration of the middle tibiae—the

central, pigmented portion being brown to dark brown in S. texana and very pale, yellowish brown in S. noctivaga. These

characters appear to be somewhat variable and are difficult to

define and use.

DESCRIPTION.—FEMALE: (1) Length 8.5-11.1 mm (mean =

10.2, n = 5); (2) wing length 2.8-3.3 mm (mean = 3.1, n = 5); (3)

abdominal width 2.6-3.3 mm (mean = 3.0, n = 5).

Structure: (4) Head broad (Figure 106); length/width ratio

0.80-0.88 (mean = 0.82, n = 5). (5) Gena, at midpoint, greatly

exceeded by width of compound eye. (9) Clypeus projecting

approximately 0.49 times its length below lower margin of

eyes; (11) clypeal surface with weakly impressed, median longitudinal depression. (13) Ocular-ocellar distance less than distance between lateral ocellus and hind margin of vertex (ocelli

enormous, ocular-ocellar space approximately 0.5 times lateral

ocellar diameter); (14) distance between lateral ocelli approximately 4.0 times ocular-ocellar distance. (16) Inner margins of

compound eyes nearly parallel. (21) Scape reaching beyond top

of vertex; (22) pedicel slightly shorter than flagellomere 1. (30)

Mandible conspicuously elongate, reaching beyond opposing

clypeal angle (Figures 13,106).

(40) Dorsal surface of propodeum about 0.70 times the

length of scutellum and approximately 1.35 times the length of

metanotum; (44) lateral propodeal carinae absent or weakly developed, extending no more than one-third the distance of posterior surface. (45) Inner hind tibial spur with 3 or 4 moderately

short teeth (Figure 64), similar to those of S. texana (Figure

65).

(46) Lateral edge of metasomal T2 nearly straight.

Sculpture: (51) Supraclypeal area tessellate near lateral

margins, (52) punctation moderately dense, most punctures

separated by their diameters. (53) Clypeus mostly polished,

tessellate only along basal and lateral margins; (54) apical

punctures only slightly larger than basal ones, very sparse, separated by 2-5 times their diameters. (55) Hypostoma striolate

throughout.

(56) Mesoscutum somewhat dull, surface tessellate throughout, (57) punctation as in Figure 111, most punctures separated

by 1-2 times their diameters. (63) Dorsal surface of propodeum

rugulose over basal half (Figure 110), (64) surface conspicuously alveolated. (65) Tl somewhat dull, surface obscurely tessellate, (66) punctation extremely fine, sparse, most punctures

separated by twice their diameters.

Coloration: (67) Abdomen mostly orange (tergum 5 usually orange-brown). (69) Flagellum and (70) tegula light orange-brown. (71) Wing membrane pale yellowish brown; veins

and stigma pale amber. (72) Legs light orange-brown.

Vestiture: (74) Hairs on head white. (75) Pubescence on

thorax white; (76) mesoscutal hairs short, many suberect or adpressed; pleuron with conspicuous amount of adpressed hairs.

(81) Basal hair bands on T2-T4 virtually absent, inconspicuous; moderately developed apical hair band present on T4.

MALE: As described for female except as follows: (1)

length 8.0-9.7 mm (mean = 8.8, n = 5); (2) wing length 2.4-2.8

mm (mean = 2.6, n = 5); (3) abdominal width 2.0-2.4 mm

(mean = 2.2, n = 5). (4) Head (Figure 107) length/width ratio

1.03-1.07 (mean = 1.05, n = 5). (23) Flagellomere 2 approximately 1.5 times length of flagellomere 1. (37) Median mesoscutal line not impressed. (70) Tegula yellow-translucent. (73)

Short, adpressed hairs on face extending dorsad to ocellar area,

weakly enclosing median ocellus. (76) Mesoscutum with conspicuously short, adpressed hairs and more elongate, less conspicuous hairs; pleuron with short, suberect to adpressed hairs.

Terminalia (Figures 112-115): (84) S7 lateral arms slender;

(85) S8 moderately developed; apex of median process truncate; (89) retrorse membranous lobe narrow, parallel sided.

FLIGHT RECORDS (Figure 116).—Females of S. noctivaga

were collected from May through October, with most records

from July. Most males were taken in July, but a few records

were collected in September and October.

SPECIMENS EXAMINED.—225 (201 females, 24 males).

UNITED STATES. ARIZONA: Coconino Co.: Cameron;

Navajo Co.: Holbrook, 17 mi NE; Winslow. COLORADO: Baca

Co.: Springfield, 31 mi SSW. KANSAS: Clark Co.; Seward

Co.: Liberal. NEW MEXICO: Bernalillo Co.: Albuquerque;

Chaves Co.; McKinley Co.: Pinedale; Roosevelt Co.: Portales;

Portales, 3.5 mi N (Oasis State Park); Santa Fe Co.: Santa Fe;

Tao Co.: Ojo Caliente; Torrance Co.: Gran Quivira.

OKLAHOMA: Ellis Co.: Shattuck; Harper Co.; Jackson Co.:

Elmer; Kiowa Co.: Lugert; Meade Co.: Cimarron River (NW

Oklahoma); Roger Mills Co.: Cheyenne; Tillman Co.: Grandfield; Woods Co.: Little Sahara State Park. TEXAS: Dickens

Co.: Dumont, 9 mi SW; Hemphill Co.: Canadian; Canadian,

4-8 mi NE; Ward Co.: Monahans, 9.5 mi S; Monahans State

Park. UTAH: Emery Co.: Big Flat Top, 2.5 mi NE (Dugout

Springs); Gilson Butte; Little Gilson Butte, 2 mi W; San Rafael

Desert (3 mi SSE Temple Mountain); Wild Horse Creek (N of

Goblin Valley); Grand Co.: Moab; Uintah Co.: Roosevelt.

6. Sphecodogastra oenotherae (Stevens), new combination

FIGURES 62,117-128

Halictus (Evylaeus) oenotherae Stevens, 1920:37 [female; compared with H.

aberrans, H. pectoralis].

Halictus ralenci Crawford, 1932:70 [female].—Mitchell, 1960:365 [synonymy].

Lasioglossum (Evylaeus) ralenci.—Michener, 1951:1110 [catalog].

Lasioglossum (Sphecodogastra) oenotherae.—Michener, 1951:1111 [catalog].

Lasioglossum (Evylaeus) oenotherae.—Linsley and MacSwain, 1962:45

[taxonomy].

Evylaeus oenotherae.—Knerer and Arwood, 1964:958 [taxonomy; noted misidentification of male of this species as E. truncatus by Mitchell (I960)].—

Knerer, 1969:142 [nesting biology].—Knerer and MacKay, 1969 [biology].—Hurd, 1979:1960 [catalog].—Moure and Hurd, 1987:78 [catalog].

Lasioglossum oenotherae.—Poole, 1996:617 [checklist].
NUMBER 610

41

FIGURES 112-115.—Sphecodogastra noctivaga, male terminalia: 112, genital capsule, ventral view; 113, genital

capsule, dorsal view; 114, right gonostylus, posterior view; 115, S7 and S8, ventral view.

120 -

100 -

3

60 -

40 -

20 -

0

Jan Feb Mar Apr May Jun Jul Aug Sep Oct Nov Dec

FIGURE 116.—Sphecodogastra noctivaga flight records, by month of collection.
42 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

TYPE MATERIAL.—The female holotype of Halictus oenotherae is in the collection of the National Museum of Natural

History, Smithsonian Institution. The specimen is labeled

"Blue Rapids, Ks.[Kansas] 8/30 P.M. Jun[e] 20 1919 O.A.

Stevens/Megapterium missouriense [handwritten]/12033

[Stevens' type number]/Type No. 23848 U.S.N.M. [red label]/

Halictus oenotherae Stevens ? type" [handwritten]. The type is

in excellent condition except for missing the last tarsomere of

the left front leg and the last three tarsomeres of the right front

leg. The female holotype of Halictus ralenci is deposited in the

American Museum of Natural History. It is labeled "Raleigh

NC [North Carolina] C S Brimley 29-V[May]-[19]24/Acc.

33827/Halictus ralenci Type Cwfd" [handwritten]. The specimen is missing the last tarsomere of the right middle leg and

the entire tarsus of the right hind leg.

DISTRIBUTION (Figure 117).—Sphecodogastra oenotherae

is the one member of this genus occurring in far eastern North

America. Only S. texana is also known to occur east of the

Mississippi River, being recorded in Wisconsin, Illinois, and

Michigan—states in which S. oenotherae apparently does not

occur. I have seen specimens of S. oenotherae from New

Brunswick and Ontario, the northern parts of its range. Moure

and Hurd (1987) also listed it from Nova Scotia. Its range extends south to Georgia. Interestingly, there are four disjunct

locality records of this species west of the Mississippi: Marshall County, Kansas (the type locality); Calcasieu County,

Louisiana; and Lee and Upshur counties, Texas. Moure and

Hurd (1987) also listed it from North Dakota. Whether this

disjunct pattern is real or was a result of inadequate collecting

FIGURE 117.—Distribution of Sphecodogastra oenotherae.

is unknown. The western records make S. oenotherae potentially sympatric, albeit narrowly, with S. texana and 5. lusoria

and also with S. aberrans if the North Dakota record is correct.

Knerer and MacKay (1969:289) correctly noted that the wide

distribution of this species in eastern North America "is seldom

reflected in representative local collections." In this current review of Sphecodogastra based on approximately 3200 specimens, I have examined only 170 S. oenotherae. An effort to

collect specimens from Oenothera in the early morning hours

and twilight period would undoubtedly extend our knowledge

of the distribution of this species. Oenothera laciniata, an

evening primrose from which S. oenotherae has been collected,

has an eastern distribution (Dietrich and Wagner, 1988) similar

to that of S. oenotherae and should be considered a primary target for potential sampling.

DIAGNOSIS.—The coarsely rugo-striate dorsal propodeal

surface (Figure 122) easily differentiates both males and females of Sphecodogastra oenotherae from all other known

Sphecodogastra.

DESCRIPTION.—FEMALE: (1) Length 6.6-8.2 mm (mean =

7.6, n = 5); (2) wing length 1.9-2.3 mm (mean = 2.1, n = 5); (3)

abdominal width 2.1-2.5 mm (mean = 2.4, n = 5).

Structure: (4) Head broad (Figure 118); length/width ratio

0.89-0.94 (mean = 0.92, n = 5). (5) Gena, at midpoint, slightly

exceeding compound eye in width. (9) Clypeus projecting approximately 0.67 times its length below lower margin of eyes;

(11) clypeal surface without median longitudinal sulcation.

(13) Ocular-ocellar distance subequal to distance between lateral ocellus and hind margin of vertex (ocular-ocellar space

approximately 2.0 times lateral ocellar diameter); (14) distance

between lateral ocelli slightly exceeding ocular-ocellar distance. (16) Inner margins of compound eyes nearly parallel.

(21) Scape reaching top of vertex; (22) pedicel subequal in

length to flagellomere 1. (30) Mandible elongate, but because

of broad head only reaching opposing clypeal angle; subapical

tooth somewhat reduced.

(40) Dorsal surface of propodeum about 0.71 times the

length of scutellum and approximately 1.25 times the length of

metanotum; (44) lateral propodeal carinae very well developed,

encircling propodeal surface. (45) Inner hind tibial spur with 4

or 5 moderately elongate teeth (Figure 62).

(46) Lateral edge of metasomal T2 broadly rounded.

Sculpture: (51) Supraclypeal area polished, (52) punctation

sparse, most punctures separated by two times their diameters.

(53) Clypeus polished; (54) punctation sparse, punctures small,

only slightly larger apically, separated by 2-3 times their diameters. (55) Hypostoma obscurely striolate throughout.

(56) Mesoscutum mostly shiny, tessellation confined to anterior one-fourth, (57) punctation as in Figure 123, most punctures separated by 1-2 times their diameters. (63) Dorsal

surface of propodeum strongly and entirely striate (Figure

122), (64) surface smooth, not alveolated. (65) Tl shiny and
NUMBER 610

43

FIGURES 118-123.—Sphecodogastra oenotherae: 118, female head; 119, male head; 120, female labrum; 121,

male labrum; 122, female propodeum; 123, female mesoscutum.

polished, (66) punctation fine, moderately sparse, punctures

separated by 1-3 times their diameters.

Coloration: (67) Abdomen dark brown. (69) Flagellum

brown. (70) Tegula light brown. (71) Wing membrane pale yellowish brown; veins and stigma light brown. (72) Legs dark

brown.

Vestiture: (74) Hairs on head white. (75) Pubescence on thorax white; (76) mesoscutal hairs moderately elongate, approximately 1.5 times median ocellar diameter; mesoscutum and pleuron without short suberect or adpressed hairs. (81) Basal hair

bands on T2-T4 weakly developed, inconspicuous; terga lacking

apical hair bands.

MALE: AS described for female except as follows: (1)

Length 7.3-8.1 mm (mean = 7.7, n = 5); (2) wing length 1.9-

2.0 mm (mean = 2.0, n = 5); (3) abdominal width 1.7-1.9 mm

(mean = 1.8, n = 5). (4) Head (Figure 119) length/width ratio

1.05-1.09 (mean = 1.07, n = 5). (23) Flagellomere 2 approximately 1.5 times length of flagellomere 1. (37) Median mesoscutal line impressed. (70) Tegula yellowish brown. (73) Short,

adpressed hairs on face extending dorsad just below median

ocellus. (76) Mesoscutal hairs moderately elongate; short, adpressed hairs inconspicuous to absent; pleuron without short,

suberect to adpressed hairs.
44 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

Terminalia (Figures 124-127): (84) S7 lateral arms slender; (85) S8 usually developed; apex of median process

rounded; (89) retrorse membranous lobe narrow, parallel sided.

FLIGHT RECORDS (Figure 128).—Females of Sphecodogastra oenotherae, like S. antiochensis, were most often collected

in late spring (unlike the other Sphecodogastra species examined in this study), with 64% of specimens taken during May.

Males were more often collected in later months, with 87%

from July and August.

At Toronto, Canada, Knerer and MacKay (1969) reported activity of this species to be much delayed, apparently to coincide

with the Oenothera bloom in that area. Females first appeared

in mid-June and some nests were not established before July.

Males were not observed at flowers until the end of July. Nest

activity continued until late August, but all nests remained

closed after September. The authors thought S. oenotherae to

be univoltine at this latitude.

SPECIMENS EXAMINED.—170 (104 females, 66 males).

CANADA, NEW BRUNSWICK: Nerepis. ONTARIO: Ottawa;

Toronto.

UNITED STATES. CONNECTICUT: Fairfield Co.: Stamford.

DISTRICT OF COLUMBIA: Washington, D.C. GEORGIA: Dade

Co.: Head River; DeKalb Co.: Stone Mountain; Rabun Co.:

Rabun Bald; White Co.: Yonah Mountain. KANSAS: Marshall

Co.: Blue Rapids. LOUISIANA: Calcasieu Co.: Vinton.

MARYLAND: Prince Georges Co.: Greenbelt. MASSACHUSETTS: Worcester Co.: Petersham. NEW HAMPSHIRE: Strafford Co.: Durham. NEW JERSEY: Bergen Co.: Alpine;

Closter; Ramsey; Essex Co.: Great Piece Meadows; Morris

Co.: Boonton. NEW YORK: Albany Co.: Rensselaerville

(Huyck Preserve); Nassau Co.: Floral Park; Rockland Co.:

Nyack; Suffolk Co.: Orient Point (Cold Spring Harbor), Setauket, Sound Beach; Sullivan Co.: White Lake; Tompkins Co.:

Ithaca (B. Danforth, pers. comm., 1996). NORTH CAROLINA:

Harnett Co.: Lillington, 10 mi S; Haywood Co.; Macon Co.:

Highlands; Wayah Bald; Wayah Gap; Transylvania Co.: Rich

Mountain; Wake Co.: Raleigh; Watanga Co.: Grandfather

Mountain; Yancey Co.: Mt. Mitchell. PENNSYLVANIA:

Cumberland Co.: Craigheads. TEXAS: Lee Co.: Lexington;

Upshur Co.: Lake McClellan. VIRGINIA: Arlington Co.:

Falls Church; Glencarlyn.

FIGURES 124-127.—Sphecodogastra oenotherae, male terminalia: 124, genital capsule, ventral view; 125, genital capsule, dorsal view; 126, right gonostylus, posterior view; 127, S7 and S8, ventral view.
NUMBER 610 45

CO

i

z 20 -

10 -

\ i i r i i i i i i i i

Jan Feb Mar Apr May Jun Jul Aug Sep Oct Nov Dec

FIGURE 128.—Sphecodogastra oenotherae flight records, by month of collection.

7. Sphecodogastra potosi, new species

FIGURES 63,129-138

TYPE MATERIAL.—The female holotype is in excellent condition and is deposited in the National Museum of Natural History, Smithsonian Institution. It is labeled "Durango Dgo.

[Durango], MEX.[ico] VIII [August]-13-1962/Domestic

squash/A.E. & M.M. Michelbacher Collectors/200.2 [green

label]/0635-0735 [flight activity]/HOLOTYPE Sphecodogastra

potosi R.J. McGinley" [red label]. Thirty-one paratypes are

designated and listed in the "Specimens Examined" section

that follows.

ETYMOLOGY.—The specific name is a reference to the Mexican state of San Luis Potosi where the largest series of this species has been collected. This was a label manuscript name used

by George E. Bohart who apparently was the first to recognize

that these specimens represented a new species.

DISTRIBUTION (Figure 83).—Sphecodogastra potosi is

known only from Hidalgo County, New Mexico, USA, and the

Mexican states of Durango, San Luis Potosi, and Zacatecas. It

is currently known to be sympatric only with S. lusoria, both

having been collected at Fresnillo (Zacatecas state) and adjacent areas in New Mexico.

DIAGNOSIS.—The presence of short, adpressed hairs on the

pleuron (e.g., Figure 21) combined with the broad head and

elongate mandibles (Figure 18) will distinguish the females of

S. potosi from other Sphecodogastra species having dark abdomens. For further details, see "Diagnosis" for S. lusoria.

Males of 5. potosi can be identified by their darkly pigmented mandibles; mandibles of other congeneric males have

conspicuous areas of yellow pigmentation. Other helpful characteristics for recognizing the males of this species are the lack

of antennal sensillar patterns (found in the very common S.

lusoria as well as S. antiochensis. Figures 42, 76) and the relatively broad head (Figure 130).

DESCRIPTION.—FEMALE: (1) Length 7.2-8.5 mm (mean =

8.0, n = 5); (2) wing length 2.3-2.5 mm (mean = 2.4, n = 5); (3)

abdominal width 2.2-2.6 mm (mean = 2.4, n = 5).

Structure: (4) Head broad (Figure 129); length/width ratio 0.90-0.92 (mean = 0.91, n = 5). (5) Gena, at midpoint, exceeding width of compound eye. (9) Clypeus projecting approximately 0.63 times its length below lower margin of eyes;

(11) clypeal surface without median longitudinal sulcation.

(13) Ocular-ocellar distance subequal to distance between lateral ocellus and hind margin of vertex (ocular-ocellar space

approximately 2.0 times lateral ocellar diameter); (14) distance

between lateral ocelli slightly exceeding ocular-ocellar distance. (16) Inner margins of compound eyes nearly parallel.

(21) Scape reaching top of vertex; (22) pedicel subequal in

length to flagellomere 1. (30) Mandible elongate, reaching

slightly beyond opposing clypeal angle (Figure 18).

(40) Dorsal surface of propodeum about 0.78 times the

length of scutellum and approximately 1.38 times the length of

metanotum; (44) lateral propodeal carinae well developed, extending to dorsal propodeal surface. (45) Inner hind tibial spur

with 4 or 5 moderately elongate teeth (Figure 63).

(46) Lateral edge of metasomal T2 broadly rounded.

Sculpture: (51) Supraclypeal area polished, (52) punctation

only moderately dense, most punctures separated by 1-2 times

their diameters. (53) Clypeus polished; (54) apical punctures
46

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

FIGURES 129-134.—Sphecodogastra polosi: 129, female head; 130, male head; 131, female labrum; 132, male

labrum; 133, female propodeum; 134, female mesoscutum.

slightly larger than basal ones, separated by 2-4 times their diameters. (55) Hypostoma weakly striolate throughout.

(56) Mesoscutum mostly shiny and polished, tessellation

confined to the anterior half, (57) punctation as in Figure 134,

most punctures separated by 2-3 times their diameters (somewhat more sparse than other species). (63) Dorsal surface of

propodeum entirely rugulose (Figure 133), (64) surface alveolated. (65) Tl shiny and polished, (66) punctation extremely

fine, moderately sparse, punctures separated by 1-3 times their

diameters.

Coloration: (67) Abdomen dark brown. (69) Flagellum

dark brown. (70) Tegula brown. (71) Wing membrane hyaline;

veins and stigma light brown. (72) Legs brown.

Vestiture: (74) Hairs on head white. (75) Pubescence on

thorax white; (76) mesoscutal hairs moderately short, length

subequal to diameter of median ocellus; mesoscutum and especially pleuron, with short, adpressed hairs. (81) Basal hair

bands on T2-T4 present, covering basal one-fourth of tergal

surface; moderately developed apical hair bands present on T3

and T4.
NUMBER 610 47

FIGURES 135-138.—Sphecodogastra potosi, male terminalia: 135, genital capsule, ventral view; 136, genital

capsule, dorsal view; 137, right gonostylus, posterior view; 138, S7 and S8, ventral view.

MALE: As described for female except as follows: (1)

length 6.6-7.8 mm (mean = 7.2, n = 5); (2) wing length 1.8-2.1

mm (mean = 2.0, n = 5); (3) abdominal width 1.5-1.8 mm

(mean = 1.7, n = 5). (4) Head (Figure 130) length/width ratio

1.04-1.08 (mean = 1.05, n = 5). (23) Flagellomere 2 approximately 1.5 times length of flagellomere 1. (37) Median mesoscutal line only faintly impressed. (70) Tegula yellowish

brown. (73) Short, adpressed hairs on face extending dorsad

just below median ocellus. (76) Mesoscutum with both moderately elongate and short, adpressed hairs; pleuron with some

short, suberect to adpressed hairs.

Terminalia (Figures 135-138): (84) S7 lateral arms slender; (85) S8 short, somewhat reduced; apex of median process

rounded; (89) retrorse membranous lobe narrow, parallel sided.

FLIGHT RECORDS.—Only six collections of S. potosi were

examined. Females were collected from May through August;

males were collected in June and August. Most specimens of

both sexes (83%) were collected in June.

SPECIMENS EXAMINED.—32 (18 females, 14 males).

MEXICO. DURANGO: Durango, 13 Aug 1962, Cucurbita,

A.E. and M.M. Michelbacher (1 $, NMNH; holotype). SAN

LUIS POTOSI: San Luis Potosi, 16 mi E, 18-19 Jun 1966,

Gaura coccinea, D.E. Breedlove (129, lid", CU). ZACATECAS: Fresnillo, 9 mi S, 24 Jun 1956, J.W. MacSwain, D.D.

Linsdale (2d, UCB); Fresnillo, 17 mi N, 16 Jul 1954, J.W.

MacSwain, E.I. Schlinger (3 9, UCB).

UNITED STATES, NEW MEXICO: Hidalgo Co.: Rodeo, 1 mi

W, 3 Aug 1961, J.G Rozen (Id, AMNH); Rodeo, 3 mi SW, 5

May 1965, J.G Rozen (2 9, AMNH).

8. Sphecodogastra texana (Cresson)

FIGURES 65,139-151

Sphecodes texana Cresson, 1872:249 [female, male, four specimens].—

1916:109 [lectotype].

Halictus (Parasphecodes) lexanus.—Cockerell, 1898b:45 [key].

Halictus lexanus.—Cockerell, 1898a: 185 [taxonomy; floral association with

pear, Senecio].—Pearson, 1933:386, 392 [recorded from Chicago, Illinois

area].—Graenicher, 1935:302 [localities; association with Oenothera rhombipetala].—Stevens, 1951:61 [floral associations with Oenothera nuttallii, O.

strigosa].

Sphecodogastra texana.—Ashmead, 1899:93 [new genus].—Graenicher,

1911:222, 223, 233 [locality records; association with Oenothera rhombipetala].—Mitchell, 1960:365 [redescription, locality, and floral records].—

Kerfoot, 1967a [nesting biology; behavior].—1967c [lunar periodicity].—

Hurd, 1979:1962 [catalog].—Moure and Hurd, 1987:85 [catalog].
48 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

Halictus (Megalopta?) texanus.—Stevens, 1920:35 [taxonomy; floral associations; localities; flight activity].

Sphecodogastre texana.—Hicks, 1936:51 [lapsus calami; nest architecture].

Lasioglossum (Sphecodogastra) texanum.—Linsley and MacSwain, 1962:47

[key separating females of L. texanum and L. noctivaga].—Linsley et al.,

1963:43 [locality records; floral associations with Oenothera caespitosa, O.

runcinata].—Gregory, 1964:400, 417 [floral records].—Kerfoot, 1967b

[ocellar size and nocturnal behavior].

Lasioglossum texanus.—Poole, 1996 [checklist].

TYPE MATERIAL.—The female lectotype of Sphecodes texana

is deposited in the Academy of Natural Sciences of Philadelphia. The specimen is structurally in good condition, but the

hairs are soiled and matted. It is labeled "Tex. [Texas]/LectoTYPE 2134 [red label]/Sphecodes Cress., 2 specimens [handwritten and folded]/ANSP" [yellow label].

DISTRIBUTION (Figure 139).—See "Distribution" for S.

noctivaga.

DIAGNOSIS.—See "Diagnosis" for S. noctivaga.

DESCRIPTION.—FEMALE: (1) Length 8.0-11.2 mm (mean =

9.5, n = 5); (2) wing length 2.7-3.1 mm (mean = 2.9, n = 5); (3)

abdominal width 2.5—3.3 mm (mean = 3.0, n = 5).

Structure: (4) Head broad (Figure 140); length/width ratio

0.90-0.96 (mean = 0.93, n = 5). (5) Gena, at midpoint, greatly

FIGURE 139.—Distribution of Sphecodogastra texana.

exceeded by width of compound eye. (9) Clypeus projecting

approximately 0.60 times its length below lower margin of

eyes; (11) clypeal surface without median longitudinal sulcation. (13) Ocular-ocellar distance less than distance between

lateral ocellus and hind margin of vertex (ocelli very large,

ocular-ocellar space approximately 0.5 times lateral ocellar diameter); (14) distance between lateral ocelli approximately 4.0

times ocular-ocellar distance. (16) Inner margins of compound

eyes converging below. (21) Scape reaching beyond top of

vertex; (22) pedicel slightly shorter than flagellomere 1. (30)

Mandible moderate in length, reaching opposing clypeal angle

(Figure 14).

(40) Dorsal surface of propodeum about 0.66 times the

length of scutellum and approximately 1.46 times the length of

metanotum; (44) lateral propodeal carinae absent or at most extending one-fourth the distance to dorsal propodeal surface.

(45) Inner hind tibial spur with 3 or 4 moderately short teeth

(Figure 65), similar to that of 5. noctivaga (Figure 64).

(46) Lateral edge of metasomal T2 broadly rounded.

Sculpture: (51) Supraclypeal area tessellate only along extreme lateral margins, (52) punctation somewhat sparse, most

punctures separated by 1-2 times their diameters. (53) Clypeus

mostly polished, tessellate along basal and lateral margins; (54)

apical punctures only slightly larger than basal ones, very

sparse, separated by 2-5 times their diameters. (55) Hypostoma

striolate, weakly so on anterior half.

(56) Mesoscutum somewhat dull, surface tessellate throughout, (57) punctation as in Figure 145, punctures separated by

1-2 times their diameters. (63) Dorsal surface of propodeum

rugulose over basal two-thirds (Figure 144), (64) surface alveolated. (65) Tl shiny and polished, (66) punctation extremely

fine, sparse, most punctures separated by twice their diameters.

Coloration: (67) Abdomen mostly orange (T4 and T5 usually dark orange-brown; T3 often darkly pigmented, and rarely,

entire abdomen darkly pigmented). (69) Flagellum orangebrown. (70) Tegula light brown. (71) Wing membrane hyaline;

veins and stigma amber to light brown. (72) Legs orangebrown.

Vestiture: (74) Hairs on head white. (75) Pubescence on

thorax white; (76) mesoscutal hairs short, many suberect or adpressed; pleuron with conspicuous amount of adpressed hairs.

(81) Basal hair bands on T2-T4 virtually absent, inconspicuous; moderately developed apical hair band present on T4.

MALE: As described for female except as follows: (1)

length 7.1-9.2 mm (mean = 8.3, n = 5); (2) wing length 2.0-2.6

mm (mean = 2.3, n = 5); (3) abdominal width 1.8-2.4 mm

(mean = 2.1, n = 5). (4) Head (Figure 141) length/width ratio

1.00-1.03 (mean = 1.02, n = 5). (23) Flagellomere 2 approximately 1.5 times length of flagellomere 1. (37) Median mesoscutal line not impressed. (70) Tegula yellow-translucent. (73)

Short, adpressed hairs on face extending dorsad to ocellar area,

weakly enclosing median ocellus. (76) Mesoscutum with conspicuous short, adpressed hairs and more elongate, less conspicuous hairs; pleuron with short, suberect to adpressed hairs.
NUMBER 610 49

FIGURES 140-145.—Sphecodogastra texana: 140, female head; 141, male head; 142, female labrum; 143, male

labrum; 144, female propodeum; 145, female mesoscutum.

Terminalia (Figures 146-150): (84) S7 lateral arms moderately well developed; (85) S8 moderately developed; apex of

median process truncate; (89) unlike other Sphecodogastra,

retrorse membranous lobe very broad and twisted ventrally.

FLIGHT RECORDS (Figure 151).—Females of S. texana were

collected from April to December, but like most species of

Sphecodogastra, they were collected primarily from May

through July (88%). Males were collected from June through

October. Graenicher (1911) reported this species to be active

from May to September in northwestern Wisconsin, and

Kerfoot (1967a) observed nesting activity during this same

period in Kingman County, Kansas. No information is available concerning the number of generations produced each

year.

SPECIMENS EXAMINED.—903 (847 females, 56 males).

MEXICO, CHIHUAHUA: Juarez. COAHUILA: Cabos; Parras,

7 mi N. MICHOACAN: Morelia, 13 mi W. VERACRUZ: El

Naranjo, 10 mi W.

UNITED STATES. ARIZONA: Yavapai Co.: Camp Verde;

Cornville, 3 mi S. COLORADO: Boulder Co.: Boulder; White

Rocks; Clark Co.: Lake George; Denver Co.: Denver; El Paso

Co.: Colorado Springs; Foster Ranch; Kiowa Co.: Eads;
50

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

FIGURES 146-150.—Sphecodogastra texana, male terminalia: 146, genital capsule, ventral view; 147, genital

capsule, dorsal view; 148, right gonostylus, posterior view; 149, right retrorse lobe, lateral view; 150, S7 and S8,

ventral view.

Larimer Co.: Fort Collins; Morgan Co.: Fort Morgan; county I

coordinates unknown: Fuente; Walsenberg, 2 mi W. ILLINOIS:

Henderson Co.: Gladstone, 1.5 mi SW; Mason Co.: Forest

City, 4 mi SW; Morgan Co.: Meredosia; Peoria Co.: Peoria.

IOWA: Johnson Co.: Iowa City. KANSAS: Butler Co.; Clark

Co.: Kingstown, 1 mi W (State Park); Cowley Co.: Winfield;

Douglas Co.: Lawrence; University of Kansas Natural History

Reservation; Finney Co.: Garden City; Kingman Co.: Kingman; Kingman, 8 mi W; Marshall Co.: Blue Rapids; Pottawatomie Co.: Little Gobi Desert; Pratt Co.: Pratt; Reno Co.:

Hutchinson; Medora (sand dunes); Riley Co.: Manhattan;

Sedgwick Co.: Wichita; Seward Co.; Summer Co.: Wellington.

MICHIGAN: Clinton Co.: Bath; Ingham Co.: Dansville; Kalamazoo Co.: Gull Lake Biological Station; Kent Co.: Grand

Rapids; Livingston Co.: E. S. George Reserve. MISSOURI:

Buchanan Co.: Mo-Kan Bridge. NEBRASKA: Antelope Co.:

Grove Lake; Cherry Co.: Valentine, 25 mi S (F. Crowe Ranch);

Cuming Co.: West Point; Dawes Co.: Chadron; Hooker Co.:

Mullen; Mullen, 1.5 mi N; Lancaster Co.: Lincoln; Scotts Bluff

Co.: Mitchell; Sheridan Co.: Hay Springs; Sioux Co.: Agate;

Thomas Co.: Halsey; Halsey, 2.5 mi W (National Forest).

NEW MEXICO: Catron Co.: Glenwood, 5 mi N; Chaves Co.;

Dona Ana Co.: Las Cruces; Eddy Co.: Artesia; Malaga, 7 mi

W; Grant Co.: Bayard; Silver City; Lincoln Co.: Hondo; Union

Co.: Clayton Lake (12 mi NW Clayton). NORTH DAKOTA:

Cass Co.: Leonard; Dickey Co.: Oakes; Ransom Co.: Sheldon;

Richland Co.: Walcott Dunes. OKLAHOMA: Alfalfa Co.:

Cherokee; Great Salt Plains; Beaver Co.: Gate; Beckham Co.:

Sayre; Caddo Co.: Hinton; Canadian Co.: El Reno; Choctaw

Co.: Grant; Hugo; Cimarron Co.: Kenton, 7 mi E (North

Crease Creek); Jackson Co.: Elmer; Jefferson Co.: Waurika;

Kiowa Co.: Lugert; Love Co.: Oswalt. Marietta Co.: Thackerville, 1 mi S; Marshall Co.: University of Oklahoma Biological

Station; Okfuskee Co.: Okemah; Oklahoma Co.: 11 mi N Oklahoma City; Okmulgee Co.: Okmulgee; Pawnee Co.: Pawnee;

Payne Co.: Stillwater; Pontotoc Co.: Roff; Roger Mills Co.:

Cheyenne; Texas Co.: Guymon; Tillman Co.: Grandfield; Tulsa

Co.: Tulsa; Woods Co.: Little Sahara State Park; Waynoka, 3
NUMBER 610 51

300 -4

250 -

g 200 -I

150 -

100 -

50 -

CO

0 -

Z7I

77"

r r r i r

Jan Feb Mar Apr May Jun Jul Aug Sep Oct Nov Dec

FIGURE 151.—Sphecodogastra texana flight records, by month of collection.

mi W; Woodward Co.: Mooreland; Woodward. SOUTH

DAKOTA: Brookings Co.: Brookings; Brown Co.: Hecla; Buffalo Co.: Ft. Thompson; Fall River Co.: Angostura Dam; Hyde

Co.: Highmore. TEXAS: Bastrop Co.: Bastrop; Brazos Co.:

Bryan; Brewster Co.: Alpine; Cameron Co.: Brownsville; Clay

Co.: Henrietta; Coryell Co.: Eddy, 12 mi W; Ft. Hood; Eastland Co.: Romney; El Paso Co.: El Paso; Gonzales Co.:

Luling; Palmetto State Park; Hale Co.: Hale Center; Memphis,

5 mi W; Harris Co.: Houston; Harrison Co.: Karnack; Hemphill Co.: Canadian; Hildago Co.: Alamo, 7 mi SE (Santa Ana

Wildlife Refuge); Howard Co.: Big Spring, 1 mi NW; Jeff

Davis Co.: Phantom Lake (Davis Mountains); Kenedy Co.:

Padre Island National Seashore; Kerr Co.: Kerrville; Kleberg

Co.: Kingsville; Lamar Co.: Paris; Mitchell Co.: Colorado

City; Nacogdoches Co.: Nacogdoches; Randall Co.: Buffalo

Lake State Park; Palo Duro Canyon; San Patricio Co.: Port

Aransas; Taylor Co.: Abilene; Buffalo Gap; Val Verde Co.: Del

Rio; Ward Co.: Monahans, 9.5 mi S; Wichita Co.: Burkburnett

(Red River); Burkburnett, 4 mi E. WISCONSIN: Pierce Co.:

Prescott; Wood Co: Port Edwards. WYOMING: Converse Co.:

Douglas; Niobrara Co.: Lusk; Platte Co.: Ft. Laramie, 11.5 mi

SW; Wheatland; Weston Co.: Newcastle, 6 mi NW.
Appendix 1

Identification of Figures in Keys

The entries below identify the figures without legends used in the illustrated keys to

Sphecodogastra species.

Key to Sphecodogastra Females

11, S. texana, head. 12, 5. aberrans, head. 13,5. noctivaga, mandibles. 14, S. texana, mandibles. 15, 5. oenotherae, propodeum. 16, S. lusoria, propodeum. 17, S. danforthi, head.

18, S. potosi, head. 19, 5. danforthi, propodeum. 20, 5. potosi, propodeum. 21,5. lusoria,

pleuron. 22, S. aberrans, pleuron. 23, S. lusoria, supraclypeal area. 24, S. aberrans, supraclypeal area. 25, 5. antiochensis, right hind femur, anterior view. 26, 5. aberrans, right

hind femur, anterior view. 27, 5. antiochensis, propodeum. 28, S. aberrans, propodeum.

Key to Sphecodogastra Males

29, 5. texana, head. 30, S. lusoria, head. 31,5. texana, genital capsule, ventral view. 32, 5.

texana, right retrorse lobe, outer lateral view. 33, 5. noctivaga, genital capsule, ventral

view. 34. 5. oenotherae, propodeum. 35, 5. antiochensis, propodeum. 36, 5. danforthi,

propodeum. 37, 5. aberrans, S3-S6. 38, 5. lusoria, S3-S6. 39, 5. aberrans, genital capsule, ventral view. 40, 5. lusoria, genital capsule, ventral view. 41,5. potosi, ocellar area.

42, 5. antiochensis, flagellum. 43, 5. potosi, flagellum. 44, 5. lusoria, flagellum. 45, 5.

potosi, propodeum. 46, 5. lusoria, propodeum. 47, 5. danforthi, genital capsule, ventral

view. 48, 5. potosi, genital capsule, ventral view. 49, 5. antiochensis, Tl. 50, 5. lusoria,

Tl.

52
Appendix 2

Onagraceae Taxonomy and Synonymy

Floral records associated with museum specimens and reports in the literature often represent names of Onagraceae that have been synonymized. The following list presents

these older names or combinations followed by (=) the currently recognized nomenclature. This information was kindly provided by Warren Wagner, Department of Botany, National Museum of Natural History, Smithsonian Institution.

Gaura nealleyi = G. suffulta nealleyi

Oenothera cespitosa var. marginata - O. caespitosa ssp. marginata

O. cespitosa var. montana = O. caespitosa ssp. cespitosa

O. clavaeformis var. aurantiaca = Camissonia claviformis ssp. aurantica

O. clavaeformis var. clavaeformis = Camissonia claviformis ssp. claviformis

O. clavaeformis var. cruciformis = Camissonia claviformis ssp. cruciformis

O. clavaeformis var. integrior = Camissonia claviformis ssp. integrior

O. drummondii = O. drummondii ssp. drummonii

O. greggii — Calylophus hartwegii pubescens

O. hartwegii = Calylophus hartwegii

O. hookeri = O. elata

O. missouriensis, Megapterium missouriense = O. macrocarpa ssp. macrocarpa

O. runcinata = O. pallida ssp. runcinata

O. strigosa = O. villosa strigosa

O. tanacetifolia = Camissonia tanacetifolia

O. tetragona = O. fruticosa glauca

O. trichocalyx = O. pallida ssp. trichocalyx

53
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