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THE UNIVERSITY OF KANSAS 
SCIENCE BULLETIN 
Vol. 54, No. 7, pp. 209-256 
L/BRAR>May 31, 1991 
JUN 1 9 199, 
Classification and Phylogenetic Ref^fbriifhips of the 
Panurgine Bees: The Calliopsini and Allies 
(Hymenoptera: Andrenidae)^ 
By Luisa Ruz^ 
CONTENTS 
Abstract 210 
Introduction 210 
Materials and Methods 210 
Acknowledgments 211 
Subfamily Panurginae 212 
Key to Tribes Calliopsini, Protomeliturgini and Perditini 212 
Tribe Perditini 213 
Genus Perdita Smith 213 
Tribe Protomeliturgini 215 
Genus Protomeliturga Ducke 215 
Tribe Calliopsini 217 
Key to the Genera of the Tribe Calliopsini 218 
Genus Acamptopoeum Cockerell 219 
Genus Calliopsis Smith 222 
Key to the Subgenera of the Genus Calliopsis 223 
Subgenera Calliopsis, 225; Perissander, 226; Calliopsima, 227; Verbenapis, 228; Liopoeum, 229; 
HypomacTotera, 231; Nomadopsis, 232; Macronomadopsis, 234; Micronomadopsis, 235; Liopoeodes, new 
subgenus, 236; Ceroliopoeum, new subgenus, 237 
Genus Arhysosage Brethes 238 
Genus Spinoliella Ashmead 240 
Key to the Subgenera of the Genus Spinoliella 241 
Subgenera Spinoliella, 242; Peniella, 242 
Genus Callonychium Brethes 242 
Key to the Subgenera of the Genus Callonychium 244 
Subgenera Callonychium, 244; Paranychium, 244 
Cladistic Analysis 244 
Appendix: A New Species of Calliopsini from Argentina 253 
Literature Cited 254 
'Contribution Number 3044 from the Department of Entomology, University of Kansas, Lawrence, Kansas 
66045. 
^Laboratorio ue Zoologi'a, Universidad Catolica de Valparaiso, Av. Brasil 2950, Valparai'so, Chile. 
210 
The University of Kansas Science Bulletin 
ABSTRACT 
This is part of a taxonomic study of the genera and subgenera of the andrenid subfamily 
Panurginae. Three tribes are recognized and described here: Calliopsini (with two new 
subgenera and one new species), Protomeliturgini (Protomeliturga) and Perditini (Perdita). The 
Calliopsini includes Acamptopoeum, Calliopsis (Calliopsis s. str.), C. (Peris sander), C. (Calliop- 
sima), C. (Verbenapis), C. (Liopoeum), C. (Hypomacrotera), C. (Liopoeodes, n. subg.), C. 
(Ceroliopoeum, n. subg.), C. (Nomadopsis), C. (Macronomadopsis), C. (Micronomadopsis), Arhyso- 
sage, Spinoliella, and Callonychium. Extensive generic descriptions provide much more 
information than has been available in the past on the features of the taxa under study. Keys 
to all genera and subgenera of these tribes are included. 
A cladistic analysis has been made in order to understand the relationships among the 
genera and subgenera of the Calliopsini and closely related taxa. At least some clarification 
of lines between genera has been attained. A broad interpretation of the genus Calliopsis is 
proposed. Illustrations for each genus-group taxon are provided. 
INTRODUCTION 
The family Andrenidae contains two sub- 
families: Andreninae and Panurginae. A 
study of the Panurginae at generic and 
subgeneric levels, with a classification and 
phylogeny of the included taxa, was pre- 
sented by Ruz (1986). That study shows that 
the Panurginae is a monophyletic group and 
that several tribes can be recognized. 
The Panurginae contains numerous gen- 
era and, like the Andreninae, is found in all 
continents except Australia. Its diversity and 
abundance, however, are greatest in the 
Western Hemisphere. 
The present paper is part of the sub- 
familial study mentioned above and treats a 
strongly derived group, the Calliopsini, 
along with its closest relatives, Protomeliturga 
(Protomeliturgini) and Perdita (Perditini). 
The three tribes are restricted to the Western 
Hemisphere. 
Robertson (1922) recognized Calliopsinae 
as one of the subfamilies of his Panurgidae 
but otherwise authors have not recognized a 
higher taxon based on Calliopsis. The Calli- 
opsini, as here understood, contains five 
genera: Acamptopoeum, Calliopsis (11 subgen- 
era, two of them new), Arhysosage, Spinoliella 
(2 subgenera) and Callonychium (2 subgen- 
era). The genus Calliopsis is treated here in a 
wider sense than in the past. 
MATERIALS AND METHODS 
About 20,000 panurgine bees have been the 
study material for this and previous investigations 
(Ruz, 1986). At least three species (3 individuals 
each, or more if required) of each polytypic genus 
and subgenus have been carefully examined and 
dissected. Structures like male genitalia and asso- 
ciated sterna, sting apparatus of female, mouth- 
parts, last tergum and sterna 5 and 6, were 
cleared in a cold solution of KOH (10%) for 12 
hours or less, depending on the degree of scleroti- 
zation of the structures, then washed with water 
for several minutes and preserved in glycerol for 
later examination. 
The morphological analysis of the species se- 
lected as representatives of each genus (type 
species plus at least two additional species when 
possible) was made using a stereomicroscope and, 
for details, a compound microscope. The names 
of the species studied in detail are preceded with 
an asterisk (*) in lists of species under each taxon. 
Illustrations were made using a camera lucida, 
comparable structures being drawn at about the 
same size regardless of differences in size of the 
bees. Sternum 6 of females was drawn with the 
latero-distal margin in its ordinary, curved posi- 
tion in order to avoid breaking it; therefore the 
distal part of the sternum appears somewhat 
narrower than in a flattened sternum. Figures of 
certain structures are omitted if similar structures 
are illustrated for related taxa or if other sources 
with good drawings are available. 
In the descriptions, each character (or groups 
of morphologically associated characters) is num- 
bered. For purposes of comparison these numbers 
have been maintained within taxa of the same 
level. Certain characters (and numbers) are omit- 
ted if they are similar in related taxa. Subfamilial 
characters are omitted in the tribal, generic and 
subgeneric descriptions, unless they represent an 
exception for the group; in this case those dis- 
tinctive characters are kept in the description. 
The number of a given character used in the 
Panurginae: Calliopsini and Allies 
211 
descriptions is not coincident with that used for 
Tables 1 and 2 and the cladogram (Fig. 30). 
In order to facilitate comparisons, the number 
of each apomorphy as it appears in Table 1 and in 
Ruz (1986) has been maintained for the whole 
revision of the subfamily. The numbers of the 
characters not used in the Calliopsini and their 
close relatives are skipped in the present work. In 
order to maintain the sequence of numbered 
apomorphies, certain autapomorphies have been 
inserted with a number already used plus a letter. 
The morphological terms in the descriptions 
are mainlv based on those used by Snodgrass 
(1935, 1956), Michener (1944, 1965, 1981), 
Rozen (1951), Winston (1979), and Michener 
and Brooks (1984). Some characters whose inter- 
pretations may be difficult are explained below. 
20. Basal area and apex of labrum: As indicated 
in Figure IF 
Tentorial pit: Small, rounded, the most dor- 
sal part of the external depression associated 
with the anterior tentorial arms (Fig. 6A, D). 
39. Orbits: Inner ocular orbits, as seen in facial 
view (Fig. 6A-D). 
46. Pterostigma and prestigma lengths (Fig. 2E) 
were measured on vein Rs; their breadths 
(maximum) were measured perpendicular to 
the lengths. Prestigmal width was not meas- 
ured to the wing margin but is only the width 
of the prestigma proper. 
48. Submarginal cells (SM): For length com- 
parison, SM cell 2 (if only two SM cells 
present) has been called SM cell 2 -t- 3 assum- 
33 
, penis valve 
basal 
sclerotization 
eye 
Width 
basal 
spine -shaped 
sclerotizatlon 
lateral 
fidge 
duplication 
clypeal protuberance 
Figure 1 . Diagram to show terminology for 
Calliopsis (A, B, D-F) and Perdita (C). A, Male 
genitalia, dorsal and ventral views; B, S8 of male; 
C, D, S6 of females; E, Head, lateral view; F, 
Labrum. 
ing that vein 1st r-m (2nd transverse cubital 
vein) has disappeared. When three SM cells 
are present, cells 2 and 3 are indicated as SM 
2 and 3. 
50. Cell 2nd M of forewings: 3rd discoidal cell. 
69. Male terga: Postgradular depression width 
measured along the longitudinal axis of the 
body. 
70. Pygidial plate: Glabrous plate on distal part 
of T7 of male and T6 of female (the latter not 
considered here), laterally delimited by car- 
inae. The same area delimited only by hairs 
or only a projection of last tergum has not 
been considered as a real plate in this study. 
71. T7 of female: Hemitergite of Michener, 1944. 
In describing margins of a leg segment, the leg 
has been considered as extended laterally; the 
antenna has been considered extended forward. 
The word vertex has been used to mean the 
posterior margin of the vertex rather than the 
whole top of the head. 
In order to save space, terga and sterna are 
abbreviated as T or S plus arable numbers of the 
metasomal segments (i.e., not counting the pro- 
podeum). Thus Tl is the first metasomal (second 
abdominal) tergum. 
ACKNOWLEDGMENTS 
Acknowledgments for the study material used 
in the whole revision of the subfamily are given 
here. They may be omitted in subsequent papers 
of this series. Abbreviations in parentheses are 
used for institutions or collections through the 
rest of the work. 
This study would not have been possible with- 
out the encouragement and assistance of numer- 
ous people. I am particularly grateful to Charles 
D. Michener of the Department of Entomology, 
University of Kansas (KU), Lawrence, Kansas, 
for his generosity, advice, interest and help in the 
preparation of the manuscript and for the use of 
material in the Snow Entomological Museum 
(KU). I also thank the following persons: George 
W. Byers and the late Peter D. Ashlock (KU) for 
their suggestions in editing the manuscript; Ed- 
ward O. Wiley (KU) for his advice and willing- 
ness to discuss aspects of this investigation; Frank 
Koch of the Museum fiir Naturkunde der Hum- 
boldt-Universitat, Berlin, for lending Friese's 
types and other valuable specimens, as well as 
generously permitting study of material from the 
collection under his care when I visited the 
Museum; George R. Else of the British Museum 
(Natural History), London, England; Till Osten 
of Staatliches Museum fiir Naturkunde, Stutt- 
gart, Germany; and Mario Zunino of the Museo 
ed Istituto di Zoologia Sistematica della Univer- 
sita di Torino, Italy, also kindly lent type spec- 
imens. The late Radclyffe B. Roberts of Rutgers 
University, New Brunswick, New Jersey and 
John L. Neff of the Central Texas Melittological 
Institute, Austin (CTMI), lent and donated im- 
212 
The University of Kansas Science Bulletin 
portant specimens. Professors Salvador Peris and 
Elvira Mingo of the Institute Espanol de Ento- 
mologia of Madrid, Spain, permitted my study of 
that collection. Yu. A. Pesenko of the Zoological 
Institute, Academy of Sciences, Leningrad; 
Klaus Warncke of Ruckteschellweg, Germany; 
George C. Eickwort of the Department of Ento- 
mology, Cornell University, Ithaca, New York; 
the late Robert O. Schuster of the University of 
California, Davis, California (UCD); Paul H. 
Arnaud of the California Academy of Science, 
San Francisco, California; Wallace E. LaBerge, 
Illinois Natural History Survey, Champaign, Illi- 
nois; and William L. Overal of the Museu Para- 
ense Emilio Goeldi, Belem, Para, Brazil, lent 
important study material. Ronald J. McGinley of 
the Smithsonian Institution, Washington, pro- 
vided material from the Reed Collection then 
under his care at Harvard University, Cam- 
bridge, Massachusetts (MCZ). Robert W. Brooks 
of the Snow Entomological Museum, University 
of Kansas, Lawrence, Kansas (KU), helped in the 
preparation of diagrams, made some preliminary 
photographs and provided certain specimens for 
study. Terry Griswold of the U.S. Department of 
Agriculture, ARS-Bee Laboratory, Utah State 
University, Logan, Utah (USDA), lent study 
material, made available the panurgine collection 
of the Bee Laboratory and kindly provided hous- 
ing on my visits to Logan. Jerome G. Rozen Jr. 
of the American Museum of Natural History, 
New York (AMNH), contributed valuable ideas, 
lent many specimens for study, permitted the use 
of the bee collection of the Museum, and very 
kindly provided housing and help when I visited 
the Museum. Zine Ajmal de Toledo of the In- 
stituto de Zoologia Miguel Lillo, Tucuman, Ar- 
gentina (IZML), lent an important panurgine 
collection under her care. Haroldo Toro of the 
Universidad Catolica de Valparaiso, Chile 
(UCV), made valuable suggestions and provided 
numerous specimens. Dr. James S. Ashe (KU) 
and Francisco Saiz (UCV) provided help and let 
me use photographic equipment under their care; 
Carlos Gonzalez (UCV), Chile, and Stephen 
Reyes (KU) did some photographic work. 
I am also indebted to Rudolf Jander, Hans- 
Peter Schultze and Arturo Roig-Alsina (all from 
KU) for their help in translating materials from 
German to English. 
The drawings were mostly inked by Donna 
Stevens (KU); some of them were also prepared 
by Carmen Tobar (UCV). 
The typing of the manuscript and editorial 
assistance was by Joetta Weaver (KU). Many 
thanks also to Virginia Ashlock (KU) for her help 
in typing and editorial work. 
This study was possible thanks to the financial 
aid of the Direccion General de Investigaciones, 
Universidad Catolica de Valparaiso, Chile, and 
especially the National Science Foundation 
grants DEB82-12223 and BSR87-16817 (C. D. 
Michener, principal investigator). 
SUBFAMILY PANURGINAE 
Diagnosis. Short-tongued bees usually with non- 
metallic integument, 2-1 1 mm long; first segment 
of labial palp usually flattened; labrum (prox- 
imally) without lateral process to articulate with 
clypeus as in Andreninae; paraglossa elongate 
and somewhat tapered distally, usually as long as 
suspensorium or longer (in Andreninae wider 
distally and shorter than suspensorium); scrobal 
suture usually absent; marginal cell distally trun- 
cate (often obliquely); middle femur of female 
with ventral margin a ridge, with a comb basally 
and sparce plumose hairs distally; scopa present, 
mainly on tibia and basitarsus, moderately abun- 
dant to sparse; T2 with lateral fovea or at least a 
dark spot; gonobase absent (except 3 genera with 
a "gonobase" that may not be homologous with 
the real gonobase). 
The three panurgine tribes considered in the 
present paper, Perditini, Protomeliturgini and 
Calliopsini, share the following principal charac- 
ters: 47. Submarginal cells two (exceptionally 
three in Perditini). 66. Tibial scopa of female 
usually (at least partially) of minutely branched 
hairs. 82. Gonocoxal apodeme not inflexed. 88. 
Sting short to extremely short, with first valvula 
slightly sclerotized, without valve (except a pos- 
sible rudiment in Calliopsis (Ceroliopoeum). 
Key to Tribes Calliopsini, Protomeliturgini and Perditini 
1. Marginal cell (nearly always) about half as long as distance between its apex and wing tip 
(Fig. 2B); if longer, then lower part efface, in profile, not or scarcely divergent from eye. S6 of 
male 3 or more times as broad as long (length measured medially); distal margin medially 
almost straight or with very wide and rather shallow V-shaped emargination. Usually with 
metallic color Perditini 
—Marginal cell more than half as long as distance between its apex and wing tip. Lower part of 
face, in profile, distinctly divergent from eye. S6 of male usually less than 3 times wider than 
long; if wider, then distal margin with tapered projections and rather deeply emarginate 
medially; or if otherwise, of different shape than above. Almost never metallic 2 
2. Tentorial pit at intersection of outer subantennal and epistomal sutures. Pre-episternal 
groove extending below scrobal level. Middle femur of female with hairs of ventral margin not 
well organized in a comb Protomeliturgini 
Panurginae: Calliopsini and Allies 
213 
— Tentorial pit in outer subantennal suture (Fig. 
below scrobal level. Middle femur of female w 
ventral margin (Fig. lOD) 
6A-D). Pre-episternal groove short, absent 
ith a distinct, well-detined comb basally on 
Calliopsini 
TRIBE P1:R1)1 UNI 
This tribe contains a single enormous genus, 
restricted to North America. It has not been 
reclassified in the present work; it is diverse but 
monophyletic. In the current classification it con- 
tains only the genus Perdita, whit h is suljciivitled 
into 21 subgenera. 
The following features characterize this tribe: 
2. Integument usually metallic and with yellow 
markings. 33. Tentorial pit usually at intersection 
mca 
pr 
Pt 
Figure 2, A-E. Forewings of males. A, Proto- 
meliturga turnerae (Ducke); B, Perdita halictoides 
Smith; C, Acamptopoeum submetallicum (Spinola); 
D, Calliopsis (CeroUopoeum) laetum (Vachal); E, 
Callwpsis (HypomacTotera) callops (Cockerell and 
Porter). pt = pterostigma; pr = prestigma; 
mca = marginal cell apex; bv = basal vein. 
ot outer subantennal and epistomal sutures. 45. 
Pre-episternal groove curved, meeting scrobe. 46. 
Pterostigma large, with sides divergent; margin 
within marginal cell clearly convex. 47. Marginal 
cell usually half as long as distance between its 
ape.x and wing tip or less. 61. Hind tibia of male 
with dorsal border usually untoothed. 69. T2-5 of 
male with posterior marginal areas glabrous or 
nearly so. 72. T8 of male usually tapered distally 
and hairy. 76. S5 of female with distal margin 
broadly concave. 77. S6 of male about three times 
as broad as long (length measured on midline). 
82. Gonocoxal apodeme not inflexed. 83. Gono- 
coxites fused ventrally. 85. Volsellae weakly at- 
tached to each other medially, with denticles. 88. 
Sting extremely short, rudimentary, second val- 
vifer elongate, first valvula slightly sclerotized, 
without valve. 
Genus Perdita Smith 
(Figs. 2, 3, 30) 
Perdita Smith, 1853: 128. Type species: Perdita halictoides 
Smith (monobasic). 
The list of subgenera and synonymy for each 
can be found in Hurd (1979), with another 
subgenus in Timberlake (1954). 
I have not attempted to study the subgenera (or 
genera?) encompassed under the name Perdita. In 
the following description, however, I have at- 
tempted to indicate the variation within Perdita in 
characters useful elsewhere at the generic level. 
To shorten the description I use mostly sub- 
generic names to identify variants; in reality, 
however, I have examined primarily the species 
listed below and the use of a subgeneric name 
does not indicate that all species of that subgenus 
have the specified characters: Perdita s. str. {halic- 
toides Smith); Pentaperdita {albovittata Cockerell); 
Perditella {larreae Cockerell); Hexaperdita {ignota 
crawfordi Cockerell); Pygoperdita {interrupta Cres- 
son); Heteroperdita {trifasciata Timberlake, female); 
Glossoperdita [pelargoides (Cockerell)]; Hesperoperdita 
{trisignata Cockerell); Alloperdita (novaeangliae 
Viereck, female); Macrotera [texana (Cresson), bi- 
color Smith]; Macroterella (mortuaria Timberlake, 
male); Macroteropsis {latior Cockerell); Pseudomacro- 
tera {turgiceps Timberlake); Allomacrotera (steph- 
anomeriae Timberlake, male); Epimacrotera (ainsliei 
Crawford); Callomacrotera {maritima Timberlake, 
acapulcona Timberlake); Cockerellia {albipennis 
Cresson); Cockerellula {opuntiae Cockerell); Pro- 
cockerellia {albonotata Timberlake); Xerophasma (be- 
quaertiana Cockerell). Additional information on 
characters and their variability among and within 
subgenera of Perdita is given by Ruz (1986) and 
Danforth (1991). 
214 
The University of Kansas Science Bulletin 
Figure 3. Perdita halictoides Smith. Male: A, B, 
Genitalia, dorsal, ventral and lateral views; C, D, 
S8, dorsal, ventral and lateral views; E, S7, 
dorsal and ventral views; F, T8; G, S6, ventral 
view. Female: H, Sting; I, Sting, ventral view; J, 
S5, ventral view; K, T7; L, S6, dorsal and 
ventral views. 
Description. 1. Length 2-11 mm. 2. Integument 
metallic (at least partially), except in species of 
the subgenera Macrolera, Aiacroterella, Xerophasma, 
Perditella, Cockerellula. 5. Pubescence in general 
short, usually minute on most of metasoma, 
especially on terga. 8. Head narrower to wider 
than thorax. 11. Glossa much shorter to longer 
than prementum; usually with flabellum. 17. 
Galea! comb of 0-15 setae. 20. Labrum less than 
twice as broad as long to more than twice as long; 
basal area usually glabrous, sometimes almost 
fully pilose; distal margin of basal area a ridge; 
labrum somewhat protuberant, with apex in- 
flexed or not. 25. Mandible of male simple to 
bidentate, or with prebasal tooth-like projection. 
30. Inner subantennal suture usually curved, 
sometimes almost straight. 33. Tentorial pit usu- 
ally at intersection of outer subantennal and 
epistomal sutures (sometimes in outer suture 
though close to the epistomal suture). 34. Anten- 
na! sockets below middle of face [e.g., P. (P.) 
halictoides Smith] to slightly above as in Xe- 
rophasma. 36. Antennal flagellum of male un- 
modified, shorter to longer than head. 37. Lower 
mesal paraocular area rather flattened or slightly 
convex as in P. (P.) halictoides (sometimes some- 
what protuberant). 39. Orbits usually sub- 
parallel, sometimes convergent below as in 
Callomacrotera, or divergent below as in male of 
Macrotera. 45. Pre-episternal groove usually 
curved, reaching scrobe, sometimes absent (e.g., 
Macrotera, Macroteropsis, Cockerellia, Pentaperdita) . 
46. Pterostigma longer than and twice as broad 
(or more) as prestigma; margin basal to vein r 
clearly diverging from costa (slightly so in Mac- 
rotera and Macroteropsis); margin within marginal 
cell usually convex [except in Macrotera and in P. 
(Macroteropsis) latior]. 47. Marginal cell broadly 
truncate, very short, usually nearly half as long as 
distance from apex to wing tip (Fig. 2B) (not as 
short in Xerophasma and Macrotera). 48. Submargi- 
nal cells two except in Xerophasma and Alloperdita, 
which have three, but second small and petiolate 
anteriorly. 50. Cell second M sometimes absent 
or weak. 51. First recurrent vein usually close to 
first transverse cubital, sometimes meeting it. 55. 
Propodeal triangle with no visible hairs (some- 
times with sparse, minute hairs [e.g., P. (Al- 
lomacrotera) stephanomenae, Epimacrotera ainsliei, and 
P. (Glossoperdita) pelargoides]. 57. Basitarsus 1 (both 
sexes) from six to nine times longer than broad; 
tarsomeres 2-4 unmodified. 58. Femur 2 of fe- 
male with comb on ventral margin basally not 
well defined (not easily differentiated from other 
hairs). 59. Middle tibial spurs finely toothed, 
slightly curved (or almost straight), somewhat 
shorter than basitarsus 2; strongly curved at apex 
in Macrotera. 61. Tibia 3 of female somewhat less 
than twice as long as basitarsus 3, with kei- 
rotrichia on most of inner surface, sometimes 
very sparse toward the middle, only at ends, or 
apparently absent; male with dorsal margin of 
tibia 3 usually untoothed, with keirotrichia usu- 
ally on most of surface, sometimes sparse. 62. 
Tibial scopa usually of rather short hairs (long in 
some species); in most cases apparently simple 
but with minute branches, with rather long alter- 
nate branches in P. (Callomacrotera) maritima Tim- 
berlake; scopal hairs usually sparse or moderately 
abundant, dense in Macrotera and Callomacrotera. 
63. Hind tibial spurs usually slightly curved, 
strongly curved at apices as in Macrotera; outer 
spur usually shorter than inner. 64. Basitibial 
plate of male well defined. 66. Claws bifurcate. 
67. Metasoma in male usually wider than to 
sometimes narrower than thorax. 70. Pygidial 
plate of male absent, though T7 at apex usually 
with median projection (projecting area truncate, 
bifurcate, or tapered). 71. T7 of female as in 
Figure 3K. 72. T8 of male generally somewhat 
tapered at apex, distinctly hairy (Fig. 3F). 74. S4 
of male with distal margin broadly and gently 
concave or almost straight. 75. S5 of male with 
distal margin widely and gently concave (some- 
times almost straight). 76. S5 of female with no 
median sclerotized area proximally and no gra- 
dulus; distal margin broadly and shallowly con- 
cave (Fig. 3J). 77. S6 of male three or more times 
wider than long (length measured medially), dis- 
tally with a very wide V-shaped emargination or 
margin almost straight (Fig. 3G). 78. S6 of 
female with no proximal laminar lobes, basal 
Panurginae: Cai.i.iopsini and Allies 
215 
sclerotization fully fused to sternum and not 
spine-shaped (similar to that of Protoineliturga) , 
lateral margin with ridge almost straight or 
curved, longitudinal basal carina absent, duplica- 
tion membranous (sclerotized area below it), 
distal margin distinctly concave or narrowly 
emarginate medially, S6 with small sparse hairs 
proximallv, forming dense patch at both sides of 
midline distally (Figs. IC, 3L). 79. S7 of male 
with two distal lateral lobes and with shallow to 
deep V-shaped or concave apical emargination 
(Fig. 3E). 80. S8 of male cross-like with distal 
part wide to narrow, truncate, rounded or ta- 
pered; basal part wider to narrower than distal 
projection (Fig. 3C, D), bilobed, or bifurcate in 
some species. 82. Gonocoxal apodeme well or 
usually strongly developed (laterally or medially 
in ventral view), not infle.xed (Fig. 3A, B). 83. 
Gonocoxites elongate, connected by slightly scle- 
rotized cuticle to fully fused ventrally. 84. Gono- 
stvlus fused to gonocoxite, well developed, 
elongate. 85. Volsellae attached to each other by 
small membranous area; with denticles. 86. Penis 
valves elongate, simple, usually tapered apically, 
fused to each other generally rather extensively 
(sometimes free only at apices). 87. Penis mem- 
branous, usually almost as long as and wider than 
valve, fused to valves in great part. 88. Sting 
short (sometimes truncate, e.g., P. halictoides), not 
reaching stylus, first valvifer (triangular plate) 
rather elongated (not triangular), first valvula 
little sclerotized, valve absent (Fig. 3H, I). 
Comments. This is the largest genus of the 
subfamily, containing approximately 500 species, 
grouped in 21 subgenera and several species 
groups. 
Discussion. The genus Perdita, in spite of all the 
diversity that it presents, has several apomorphies 
that make it a distinctive group. According to the 
cladogram (Fig. 30) this genus is the sister group 
of the Calliopsini and Protomeliturga together. 
Rozen (1966) has found that Perdita has distinct 
larval characters different from those of any other 
Panurginae. 
The pollen balls in the cells of several species of 
Perdita are coated with a secreted, cellophane-like 
layer, a synapomorphy shared with the Calliop- 
sini (Rozen, 1967, and personal communication). 
li Protomeliturga does not cover the pollen ball, this 
fact may mean that Perdita, hot Protomeliturga, is 
the sister group of Calliopsini. This was the result 
that I obtained in a preliminary cladistic analysis 
using a smaller set of characters. 
Distribution. This genus occurs in southern 
Canada, the United States, Mexico and Guate- 
mala (Hurd, 1979). 
TRIBE PROTOMELITURGINI 
The main characters of this monotypic tribe 
are the following: 20. Labrum of female with 
basal area hairy laterally. 33. Tentorial pit at 
intersection of outer subantennal and epislomal 
sutures. 45. Pre-episternal groove extending be- 
low scrobal level. 46. Pterostigma with sides 
diverging; margin within marginal cell convex. 
61. Hind tibia of male with dorsal border a 
carina. 69. T2-5 of male with posterior marginal 
areas pilose laterally forming a hair band. 70. T7 
of male with a strong lateral projection. 76. S5 of 
female with distal margin almost straight. 82. 
Gonocoxal apodeme not inflexed. 83. Gonocox- 
ites short, fused ventrally. 85. Volsellae well sepa- 
rate from each other, without denticles. 87. Penis 
extremely wide and completely fused to penis 
valves. 88. Sting very short, first valvula little 
sclerotized, without valve. 
Genus Protomeliturga Ducke 
(Figs. 2A, 4, 5, 30) 
Protomeliturga Ducke, 1912: 63, 90. Type species: Calli- 
opsis turnerae Ducke, 1907 (monobasic). 
Diagnosis. Segments 3 and 4 of labial palp at 
right angle to segment 2. Basal vein (forewing) 
strongly curved. Terga with distal hair bands 
laterally. Male. Antennal flagellum much shorter 
than length of head. Tibia 3 with dorsal margin a 
sharp carina. T7 at apex strongly bent down and 
forward, with a conspicuous projection laterally. 
G \-y w H 
Figure 4. Protomeliturga turnerae (Ducke). Male: 
A, B, Genitalia, dorsal, ventral and lateral views; 
C, S7, dorsal and ventral views; D, S8, lateral 
view; E, T7; F, Leg 3, outer view; G, S8, dorsal 
and ventral views; H, S8, ventrolateral view, 
tc = tibial carina. 
216 
The University of Kansas Science Bulletii 
Female. Middle tibial spur finely and densely 
toothed. Tibia 3 more than twice as long as 
basitarsus 3, with scopa mostly of long, minutely 
branched hairs. Inner hind tibial spur conspic- 
uously curved. 
Description. 1 . Length 5-8 mm. 3. Lower part of 
face (clypeus of male, small spot in lower paraoc- 
ular of female) and areas of thorax and legs 
yellow. 4. Metasoma with no yellow marks. 5. 
Pubescence in general short, mostly appressed, 
rather abundant, inconspicuous in most of dor- 
sum of thorax. 6. Integument microareolate, dull 
on most of head and thorax, shiny on metasoma. 
7. Punctures fine, mostly dense. 8. Head wider 
than long and narrower than thorax (sometimes 
as broad as thorax). 11. Glossa somewhat longer 
than prementum. 13. Segment 1 of labial palp 
longer than 2-4 together. 14. Segment 2 of labial 
palp about three times longer than 3. 15. Seg- 
ment 3 of labial palp articulated at angle to 
segment 2 so that 3 and 4 project at angle to 1 and 
2 (as in long-tongued bees). 16. Maxillary blade 
more than twice as long as prepalpal part of 
galea. 17. Galeal comb of about 11 bristles. 20. 
Labrum somewhat wider than long; basal area 
densely hairy and with distal margin a strongly 
projecting ridge; labral apex strongly inflexed as 
in Pseudopanurgus (Fig. 5B, C). 27. Clypeus 
slightly more than twice as wide as long in male, a 
little less than three times in female; moderately 
protuberant (about 1/3 width of eye in lateral 
view); distal margin with rounded projection near 
lateral margin of labrum (Fig. 51, K, L). 30. 
Inner subantennal suture almost straight. 31. 
Subantennal area wider than half length of inner 
suture and as wide as socket. 33. Tentorial pit at 
intersection of epistomal and outer subantennal 
sutures (Fig. 51). 34. Antennal sockets in middle 
of face. 36. Flagellum of male slightly clavate 
(Fig. 5J), much shorter than head; Hagellomere 1 
about twice as long as 2, longer than broad. 37. 
Lower mesal paraocular area slightly convex. 38. 
Facial fovea narrow and shallow in male, wider 
and well-marked in female. 39. Orbits convergent 
below in male, subparallel in female. 40. Ocelli 
mostly above orbital tangent. 41. Vertex convex. 
42. Gena of male (lateral view) narrower than 
eye, of rather uniform width. 43. Pronotum with 
dorsolateral preapical lamella, without dorsal 
ridge medially. 44. Mesepisternum with almost 
no flat area facing anteriorly. 45. Pre-episternal 
groove rather shallow, punctate, extending below 
scrobe, more clearly marked above. 46. Ptero- 
stigma longer and wider than prestigma; side 
basal to vein r clearly diverging from costa, that 
within marginal cell convex. 47. Marginal cell 
rather pointed at apex (obliquely truncate. Fig. 
2A); length little shorter to somewhat longer than 
distance from its apex to wing tip. 48. Submargi- 
nal cells two. 49. Submarginal cell 1 about as long 
as cell 2-1-3. 50a. Basal vein strongly curved. 51 . 
First recurrent vein distant from first transverse 
cubital. 52. Forewing with cu-v longer than 
second abscissa of M-f Cu. 53. Hind wing with 
cu-v 1/4 to 1/5 as long as second abscissa of 
M-I-Cu. 54. Dorsal surface of propodeum about 
as long as metanotum. 55. Propodeal triangle 
densely pilose, especially on lateral areas, with no 
striae. 57, Basitarsus 1 about six times longer 
than broad in male and more than three times 
longer in female; tarsomeres 2-4 unmodified. 58. 
Femur 2 of female on ventral margin basally with 
comb not well developed and not clearly defined. 
59. Middle tibial spur of both sexes slightly more 
than half as long as basitarsus 2 with fine, minute 
and dense teeth. 60. Basitarsus 2 of both sexes 
about as long as 1 and shorter than 3. 61. Tibia 3 
of female more than twice as long as basitarsus 3 
(Fig. 5H), with keirotrichia on most of inner 
surface except close to dorsal and ventral mar- 
gins; male tibia 3 with dorsal margin a strong, 
sharp carina (as in Pseudopanurgus) (Fig. 4F), with 
keirotrichia as in female. 62. Tibial scopa mostly 
of long, branched hairs (some with branches only 
distally, a few on dorsal margin simple). 63. Hind 
tibial spurs with minute, fine teeth; outer some- 
what shorter than inner and slightly curved at 
Figure 5. Protomeliturga turnerae (Ducke). Male: 
A, 86, ventral view. Female: B, Labrum; C, 
Labrum, lateral view; D, S6, dorsal and ventral 
views; E, T7; F, Sting; G, S5, ventral view; H, 
Leg 3, outer view; I, Head, frontal view (hairs 
and labrum omitted). Male: J, Antenna; K, L, 
Head, frontal and lateral views (hairs omitted, 
labrum only in K). tp = tentorial pit; cdp = distal 
clypeal projection. 
Panurginae: Calliopsini and Allies 
217 
apex; inner more ccmspiciiously cui\ed, espe- 
ciallv in female. 64. Basitibial plate of male well 
defined, margins carinate. 65. Tarsus 3 with no 
modifications except basitarsus of female with 
small apical projection. 66. Claws deeply cleft; 
rami subequal in male, inner shorter than outer 
in female. 67. Metasoma in male somewhat 
broader than thorax. 69. T2-5 of male with 
gradulus posterolaterally absent and with 
postgradular depression rather shallow; posterior 
marginal areas of Tl-5 in male and 'ri-4 in 
female pilose, forming bands laterally (band less 
distinct on T5 of male and T4 of female); lateral 
fovea of T2 (both sexes) rather narrow, gently 
depressed. 70. Pygidial plate of male absent; T7 
at apex strongly curved down and forward, with a 
conspicuous projection laterally (Fig. 4E). 71. T7 
of female as in Figure 5E. 72. T8 of male 
trapezoidal. 73. Metasomal sterna of male mostly 
with hairs straight, short, appressed, except on S6 
obliquely directed toward midline. 73a. SI -5 of 
female with hairs as in male but longer and 
branched on premarginal areas. 74. S4 of male 
with distal margin slightly and broadly convex. 
75. S5 of male with margin straight medially. 76. 
S5 of female with neither median proximal sclero- 
tized area nor gradulus; distal margin broadly 
and gendy concave (Fig. 5G). 77. S6 of male 
broadly and gently concave medially (Fig. 5A). 
78. S6 of female similar to that of Pseudopanurgus 
except longitudinal basal carina absent; duplica- 
tion well sclerotized; distal margin slightly con- 
cave medially; premarginal area fully hairy but 
sparsely so medially; rest of sternum with hairs 
minute (Fig. 5D). 79. S7 of male wide, distally 
with a V-shaped emargination and a lateral, short 
projection; proximal arms forming a U (Fig. 4C). 
80. S8 of male wide, large, strongly carinate 
laterally and much exposed distally (surpassing 
T7) (Fig. 4D, G, H). 82. Gonocoxal apodeme 
well developed (dorsal view), rather wide, not 
inflexed (suture almost invisible) (Fig. 4A, B). 83. 
Gonocoxites much shorter than penis valve, fused 
medially. 84. Gonostylus fused to gonocoxite, 
slightly shorter than half length of gonocoxite. 85. 
Volsella strongly lateral in position, free mesally, 
fused to gonocoxite proximally, without denticles. 
86. Penis valves wide, completely fused to each 
other and to penis. 87. Penis long (slightly sur- 
passing penis valve), extremely wide, with two 
mesal longitudinal sclerotized areas. 88. Sting 
short, not reaching stylus, first valvula little scle- 
rotized, valve absent (Fig. 5F). 
Comments. This genus is known only from one 
species, Prolomeliturga lurnerae (Ducke). 
The material available for this study was: the 
type of P. turnerae (lectotype female and lec- 
toallotype male) from the Museu Paraense Emilio 
Goeldi (Belem, Para, Brazil) (not dissected) and 
two specimens (one male and one female) from 
the Berlin Museum which, although labeled 
"Typus," are not the real types. The specimens 
from the two museums are certainly conspecific. 
Discussion. The cladistic analysis shows this 
species to be the sister group of the Calliopsini. It 
presents a series of derived features, although at 
first sight the genus suggests a rather primitive 
panurgine. Probably because of its lateral hair 
bands on the metasomal terga, Ducke (1907) 
erroneously placed it in Calliopsis. 
Distribution. Prolomeliturga is known only from 
Sao Luiz de Maranhao, Brazil. 
TRIBE CALLIOPSINI 
This tribe, found only in the western hemi- 
sphere, is the most derived group of the Pan- 
urginae. I list below a series of its characters, 
using the same series of character numbers as in 
the generic descriptions, in order to avoid repeti- 
tion in the descriptions of the genera as well as to 
emphasize the unity of the tribe. 
30. Inner subantennal suture about as long as 
antennal socket diameter or shorter. 33. Tentorial 
pit in outer subantennal suture. 45. Pre-epister- 
nal groove rather short, absent below scrobal 
level. 46. Pterostigma basally nearly as wide as 
distally or little narrower; side within marginal 
cell usually straight. 48. Submarginal cells two. 
55. Propodeal triangle glabrous. 58. Middle 
femur of female, on ventral margin, with a 
distinct, well-defined comb. 61. Tibia 3 of female, 
on inner surface, with keirotrichia present toward 
dorsal border and at base and apex, sometimes 
only at the two ends; male tibia 3 with dorsal 
border untoothed. 62. Tibial scopa, on outer 
surface, of sparse to extremely sparse, mostly 
minutely branched and simple hairs. 69. Tl-5 of 
male and T 1-4 of female with posterior marginal 
area usually completely or partially hairy (short 
hairs). 70. Pygidial plate of male usually present. 
72. Tergum (T)8 of male usually about as wide as 
long and roughly hexagonal but with lateral 
angles rather rounded. 73. Sl-5 of male with 
hairs mostly directed straight backward. 76. S5 of 
female with a median sclerotized area between 
proximal margin and gradulus except in Callo- 
nychium (Fig. lOE); gradulus usually short except 
in Callonychium long and recurved; distal margin 
broadly convex medially. 78. S6 of female with 
proximal laminar lobes; with basal spine-like 
sclerotization except of different shape in some 
Calliopsis (Nomadopsis) and absent in Callonychium; 
basal sclerotization free from disc of sternum 
distally (except fused on Arhysosage); lateral mar- 
gin with a strong, usually almost straight ridge; 
longitudinal basal carina absent; duplication usu- 
ally at least somewhat sclerotized except thin in 
some Calliopsis s. 1.; most of surface with minute 
sparse hairs except apically with well-defined 
rows of dense hairs forming a lateral patch or a 
continuous, usually curved, band of hairs. 80. S8 
of male with distal projection well developed; 
218 
The University of Kansas Science Bulletin 
basal body with a lateral acute projection except 
in Acamptopoeum and Calliopsis (Verbenapis), which 
have no projection or only a small convexity, 
respectively. 82. Gonocoxal apodeme (ventral 
view) conspicuous, not inflexed. 83. Gonocoxites 
globose, connected with one another by weakly 
sclerotized cuticle to completely fused and highly 
sclerotized. 84. Gonostylus inconspicuous or ab- 
sent. 85. Volsella usually well developed and 
without denticles [except with denticles in C. 
(Nomadopsis)]; volsella sometimes apparently 
absent or rudimentary (completely fused to gono- 
coxite?) as in Arhysosage, Spinoliella and Callonych- 
lum. 86. Penis valves dorsally fused through a 
small, narrow bridge. 88. Sting usually short or 
greatly reduced (except surpassing stylus in 
Acamptopoeum); second valvifer (triangular plate) 
usually unmodified; first valvula only slightly 
sclerotized, valve absent [except a rudiment of 
valve in Calliopsis (Ceroliopoeum)]. 
cdp 
Figure 7, A, B. Heads of males, ventral views. 
A, Spinoliella (Peniella) maculata (Spinola); B, Callo- 
nychium (Callonychium) mandibulare Brethes. 
cdp = distal clypeal projection; es = epistomal 
suture; c = clypeus, lateral area; p = paraocular 
area. 
Key to the Genera of the Tribe Calliopsini 
1. Male 2 
— Female 6 
2. Orbits convergent below. Tentorial pit clearly below median point of outer subantennal 
suture (Fig. 6A, B) 3 
B 
D 
Figure 6, A-D. Heads of males, frontal views. A, Acamptopoeum submetallicum (Spinola); B, Calliopsis 
(Calliopsis) andreniformis Smith; C, Arhysosage ochracea Brethes; D, Callonychium (Paranychium) chilense 
(Friese). tp = tentorial pit; ff= facial fovea. 
Panurginae: Calliopsini and Allies 
219 
— Orbits subparallcl or divergent below. Tentorial pit at nietiian point of outer subantennal 
suture or nearly so (Fig. 6C, D) 4 
3. T7 dist.illv with a median smooth area, delimited by hairs, tapered at apex. S4 with distal 
maigin straight medially. S5 on distal margin without median projection or any other 
modification Acamptopueum 
— T7 with pygidial plate rounded or truncate at ape.x (sometimes poorly delimited by ridge or 
carina laterally). S4 with distal margin usually produced in middle, broadly concave mesally 
(Fig. IIG). S5 with well-developed median projection on distal margin, though sometimes 
inconspicuous (Figs. IIC, 18E, 20H) Calliopsis 
4. Orbits strongly divergent below. Lower paraocular area flat. Most of body yellow. Hind 
tibial spurs strongly curved (similar to those of female in Fig. 26C). Metasoma much wider 
than head. Total length about 1 cm or nearly so Arhysosage 
— Orbits usually subparallel, if not so, lower paraocular area very swollen on inner area. Body 
with yellow markings but usually predominantly black. Inner hind tibial spur almost straight, 
outer straight or somewhat curved (similar to those of female in Fig. 27G). Metasoma as 
broad as head. Total length 4-7 mm 5 
5. Metasoma at apex slightly curved or straight. Paraocular area yellow only on lower part. 
Clypeus (ventral view) distally with projection beside lateral part of labrum; epistomal suture 
laterally sinuous (Fig. 7A). Antennal socket usually at lower 1/3 of face Spinoliella 
— Metasoma at apex strongly curved downward and foreward. Paraocular area with yellow 
surpassing antennal socket level, usually narrowly following orbit. Clypeus (ventral view) 
with margins of lateral areas usually almost straight (Fig. 7B). Antennal socket (lower 
margins) usually at lower 1/4 of face Callonychium 
6. Labrum with basal area well excavated, distal part convex, protuberant in lateral view. 
Orbits generally convergent below or if not, lower paraocular area not swollen on inner corner 
7 
— Labrum flat or with smooth, rounded, nearly transverse ridge, distal area flat, not inflexed. 
Orbits subparallel or divergent below 8 
7. Labrum with basal area at least laterally pilose (Fig. 9F, G). Hind tibia with keirotrichia 
widespread on most of inner surface (Fig. lOF) Acamptopoeum 
— Labrum with basal area usually glabrous; if pilose, also flat (without ridge). Hind tibia with 
keirotrichia widespread but absent toward ventral margin to completely absent between dense 
patch at each end Calliopsis 
8. Orbits divergent below. Metasoma wider than thorax. Middle and hind tibial spurs strongly 
curved at apices (Fig. 26C). Lower paraocular area convex Arhysosage 
— Orbits subparallel. Metasoma about as wide as thorax or narrower. Middle and hind tibial 
spurs slightly curved (Fig. 27G). Lower paraocular area strongly convex only on inner corner 
(Fig. 7A, B) 9 
9. Gena black. Paraocular area on lower part with yellow spot not narrowed along orbit. 
Antennal sockets at lower 1/3 of face. Claws bifurcate. Facial fovea not linear Spinoliella 
— Gena with longitudinal yeUow band to completely yellow (Fig. 8D). Paraocular area with 
lower yellow spot narrowed along orbit (Fig. 6D). Antennal sockets (lower margins) usually at 
lower 1/4 of face. Claws simple. Facial fovea linear Callonychium 
Genus Acamptopoeum Cockerell 
(Figs. 2C, 6A, 9, 10, 30) 
Friesea Schrottky, 1902: 418 (preoccupied). Type spe- 
cies; Friesea brasiliensis Schrottky, \902= Acamptopoeum 
prinii (Holmberg, 1884) (monoba.sic). 
Acamptopoeum CockcTcW, 1905: 320. Type species: Camp- 
topoeum trifasciatum Spinola, 1851, sensu Cockerell, 
1905 = Acamptopoeum submelallicum (Spinola, 1851) 
(monobasic and original designation). 
Parafriesea Schrottky, 1906: 118 (new name for Friesea 
Schrottky). Type species: Friesea brasiliensis Schrottky, 
1902 = Acamptopoeum prinii (Holmberg, 1884) (auto- 
basic). 
Diagnosis. Close to Calliopsis; distinguishing 
characters are as follows: Pubescence of thorax 
similar in length and density to that of Colletes. 
Orbits convergent below. Labrum with strong 
transverse salient. Hair bands (sometimes absent 
in male) on premarginal areas of metasomal 
terga. Male. S4 and S5 with distal margins almost 
straight. Female. Middle tibial spur coarsely 
toothed. Sting surpassing stylus, truncate at 
apex. 
Description. 1. Length 8-11 mm. 2. Integument 
dull or metallic. 3. Lower half of face (extending 
up beside orbit), pronotum (sometimes) and legs 
yellow (spots reduced in female). 4. Metasoma 
220 
The University of Kansas Science Bulletin 
Figure 8, A-D. Calliopsine males, lateral views. A, Acamptopoeum submetallicum (Spinola); B, 
Calliopsis (Calliopsis) andreniformis Smith; C, Arhysosage ochracea Brethes; D, Callonychium (Paranychium) 
chilense (Friese). 
with no yellow marks. 5. Pubescence relatively 
long on head and thora.x, denser on thorax 
(similar to that of Colletes), much shorter on 
metasoma. 7. Punctures in general fine, usually 
dense on thorax and metasoma, sparser on sterna 
of male. 8. Head distinctly broader than long and 
broader than thorax. 11. Glossa less than half as 
long as prementum, slender. 13. Segment 1 of 
labial palp almost twice as long as 2-4 together or 
somewhat longer. 16. Maxillary blade less than 
twice as long as prepalpal part of galea. 1 7 . Galeal 
comb of 18-24 bristles. 20. Labrum less than 
twice as broad as long; basal part partially or 
almost completely pilose; distal margin of basal 
area strongly salient, hiding inflexed apical part 
of labrum (Fig. 9F, G). 25. Mandible with upper 
margin rather conspicuously medially produced. 
27. Clypeus somewhat less than three times 
broader than long and somewhat protuberant 
(about 1/3 width of eye in lateral view); distal 
margin with a usually conspicuous acute or 
rounded projection near lateral margin of la- 
brum. 30. Inner subantennal suture angulate. 31. 
Subantennal area wider than inner suture length 
and than socket. 33. Tentorial pit in outer suban- 
tennal suture, but close to epistomal suture. 34. 
Antennal sockets approximately at middle of face. 
36. Antennal flagellum of male unmodified, 
somewhat longer than head; flagellomere 1 about 
as long as 2 and about as long as broad. 37. 
Lower mesal paraocular area rather flattened or 
slightly convex. 38. Facial fovea oval, shallow. 39. 
Orbits strongly convergent below. 40. Ocelli 
above dorsal orbital tangent. 41. Vertex convex. 
42. Gena of male (lateral view) somewhat vari- 
able (usually narrower than eye but sometimes 
wider; wider dorsally than ventrally. 43. Pro- 
notum with dorsal preapical ridge rounded. 44. 
Mesepisternum with anterior surface sometimes 
reduced, slightly convex. 45. Pre-episternal 
groove curved, reaching scrobal level, not extend- 
ing below. 46. Pterostigma somewhat longer and 
somewhat wider than prestigma; margin basal to 
vein r subparallel to costa; that within marginal 
cell straight (or slightly convex). 47. Marginal cell 
obliquely and rather broadly truncate at apex, 
much longer than distance from its apex to wing 
tip. 49. Submarginal cell 1 shorter (at least 
Panurginae: Calliopsini and Allies 
221 
slightly) than cell 2 + 3. 51. First recurrent vein 
tar from hrst transverse cubital. 52. Forewing 
with cu-\' two or three times longer than second 
abscissa \I + Cai. 53. Hind wing with cu-v 1/4 to 
1/5 as long as M + Cu. 54. Dorsal surface of 
propodeum about as long as metanotum. 55. 
Propodeal triangle very smooth, depressed medi- 
ally. 57. Basitarsus 1 fi\e to seven times longi-r 
than broad in male, about four times longer than 
broad in female; tarsomeres 2-4 unmodified. 59. 
Middle tibial spur of female about as long as 
basitarsus 2 or somewhat shorter, with coarse 
teeth (Fig. lOG); in male rather longer than half 
basitarsal length and with hne, dense teeth. 60. 
Basitarsus 2 of male longer than 1 or 3 (2 and 3 
somewhat variable in female); tarsomeres 2-4 
unmodified. 61. Tibia 3 of female somewhat less 
than twice as long as basitarsus 3, with kei- 
rotrichia on inner surface e.xcept toward ventral 
and dorsal margins (Fig. lOF); male tibia 3 with 
keirotrichia as in female. 62. Tibial scopa rather 
sparse, though dense on dorsal margin; outer 
surface with branched and simple hairs. 63. Hind 
tibial spurs (both se.xes) with fine and dense teeth 
(appearing untoothed), slightly curved toward 
apices, outer somewhat shorter than inner. 64. 
Basitibial plate of male well defined. 65. Tarsus 3 
with no modifications e.xcept basitarsus of female 
with apical process. 66. Claws bifurcate; rami 
subequal in male, inner much shorter than outer 
in female. 67. Metasoma in male somewhat wider 
than thorax. 69. T2-5 of male with gradulus 
posterolaterally not surpassing anterior half of 
tergum, not carinate, with postgradular depres- 
sion on T2 and T3 well marked and narrow. 
Figure 9. Acamptopoeum submetallicum (Spinola). 
Male: A, B, Genitalia, dorsal, ventral and lateral 
views; C, D, S7 and 8, dorsal and ventral views; 
E, S6, ventral view; F, Labrum; G, Labrum, 
lateral view; H, T8. 
shallower on posterior terga; posterior marginal 
areas of T 1-5 in male and Tl-4 in female pilose, 
forming hair bands (sometimes weak in male); 
lateral fovea of T2 (both sexes) oval, weak (bor- 
ders difiicult to see). 70. Pygidial plate of male 
absent, represented only by an apically acute, 
smooth and bare area. 71. T7 of female squared 
(Fig. IOC). 72. T8 of male as in Figure 9H. 73. 
Metasomal sterna of male with hairs directed 
caudad, mostly on premarginal areas and not 
appressed, sparse or absent on midline. 73a. Sl-5 
of female with hairs short, dense and appressed, 
mixed with hairs somewhat longer, sparser, and 
not appressed (Fig. IDE). 74, 75. S4 and 5 of male 
with distal margins almost straight medially. 77. 
S6 of male with small apical emargination (Fig. 
9E). 78. S6 of female with basal sclerotization 
long and very acute, duplication fully and 
strongly sclerotized, distal margin concave medi- 
ally, S6 with patch of dense hairs on premarginal 
area, interrupted medially (Fig. lOH). 79. S7 of 
male with distal median projections short, prox- 
imal arms widely open (Fig. 9C). 80. S8 of male 
with median distal projection long, clavate, 
abruptly separated from a transversely rectan- 
gular body (Fig. 9D). 83. Gonocoxites globose, 
ventrally completely fused to each other. 84. 
Gonostylus fused to gonocoxite, short (almost 1/3 
length of gonocoxite in ventral view). 85. Volsel- 
lae well developed, fused to each other forming a 
highly sclerotized plate, without denticles. 86. 
Penis valves with folds, somewhat intricate, 
rather rounded at apices; dorsally fused to each 
other by small and narrow bridge; laterally with a 
long row of hairs (Fig. 9A, B). 87. Penis usually 
reaching apex of penis valve or nearly so, nar- 
rower than valve, largely sclerotized, clearly sepa- 
rated from penis valve. 88. Sting elongate, 
truncate at apex, surpassing stylus (Fig. lOA, B). 
Comments. Cockerell indicated that the type 
species of this genus is Camptopoeum trifasciatum 
Spinola, 1851 (now in Liopoeum). His description, 
however, corresponds to Camptopoeum submetal- 
licum Spinola, 1851. Because of this misidentifica- 
tion the latter species has been considered for 
many years as the type species of Acamptopoeum. 
According to the International Code of Zoological 
Nomenclature, Article 70c, cases of misidentified 
type species have to be referred to the Interna- 
tional Commission on Zoological Nomenclature 
for resolution. I will therefore request the Inter- 
national Commission to designate Camptopoeum 
submetallicum Spinola ( = C. trifasciatum of Cocker- 
ell, not Spinola, 1851) as the type species of 
Acamptopoeum. 
If the request is denied, then the name used by 
Cockerell in designating the type species must be 
The University of Kansas Science Bulletin 
Figure 10. Acamptopoeum submetallicum (Spin- 
ola). Female: A, B, Sting, ventral and lateral 
views; C, T7; D, Femur 2 (hairs on inner surface 
omitted); E, S5, ventral view; F, Tibia 3, inner 
view; G, Spur, leg 2; H, 86, dorsal and ventral 
views. fc = femoral comb; ke = keirotrichia; 
mps = median proximal sclerotization. 
maintained. In that case Acamptopoeum would 
contain the species here placed in Calliopsis (Lto- 
poeum); Liopoeum would be a junior synonym. 
Parafriesea would stand as the generic name for the 
species now placed in Acamptopoeum. 
The South American genus Acamptopoeum is 
represented by the following species: *A. argenti- 
/zuOT (Friese, 1906a), *A. pnVzn (Holmberg, 1884), 
A. inauratum (Cockerell, 1926), A. colombi- 
ensis Shinn, 1965, A. mgritarse (Vachal, 1909), A. 
vagans (Cockerell, 1926), A. maculatum (Smith, 
1853) (see Shinn, 1965), and *A. submetallicum 
(Spinola, 1851). 
The type locality (east Florida) given for A. 
maculatum is not within the otherwise known 
range of the genus. It seems very unlikely that 
Florida is its real habitat (Shinn, 1967; Mitchell, 
1960, under Calliopsis). 
Discussion. Acamptopoeum is a distinctive genus 
although in the past its position has been con- 
fused. At first its species were considered as 
Camptopoeum, together with European and other 
South American species (Spinola, 1851); later it 
was treated as synonym of Liopoeum (Michener, 
1944; Rozen, 1951, 1958). 
Shinn (1967) noted that according to Moure 
(1956, personal communication to Shinn) Acamp- 
topoeum constitutes "a clearcut group of closely 
related species." On the other hand, Shinn 
(1967) agrees with Michener (1944: 246) in con- 
sidering the possibility that '' Parafriesea = Acampto- 
poeum ... is probably a mere subgenus of Calliop- 
sis. 
My study, however, does not support the idea 
of including Acamptopoeum in any other genus. 
The cladistic analysis indicates it to be the sister 
group of Calliopsis s. 1. The cladogram (Fig. 30) 
shows ihai Acamptopoeum, in spite of the superficial 
similarity to some subgenera of Calliopsis, pres- 
ents a series of apomorphies which support it as a 
good genus (see discussion in Cladistic Analysis 
section). 
Distribution. This genus occurs in Argentina, 
Peru, Paraguay, Uruguay, Brazil, Colombia, and 
Chile (Moure, in litt.; Schrottky, 1903, 1913; 
Friese, 1906a, b; Holmberg, 1844, Shinn, 1965). 
Genus Calliopsis Smith 
Diagnosis. Superficially similar to Acamptopoeum 
but differing as follows: Pubescence of thorax 
usually shorter and less dense than in Colletes. 
Labrum usually with transverse ridge but not 
strongly salient. Male. S4 with distal margin 
produced medially (sometimes inconspicuously). 
S5 usually with well-developed median projection 
distally. Female. Sting not reaching stylus, tapered 
at apex. 
Description. 3. Head, thorax, and legs usually 
with yellow marks. 8. Head broader than long 
(sometimes as broad as long in female). 20. 
Labrum with basal area glabrous. 33. Tentorial 
pit in outer subantennal suture, close to epistomal 
suture. 39. Orbits at least slightly convergent 
below, usually more strongly so in male; or in 
female sometimes subparallel. 41. Vertex at least 
slightly conve.x. 45. Pre-episternal groove not 
reaching scrobal level. 74. S4 of male with distal 
margin produced in the middle, sometimes incon- 
spicuously so. 75. S5 of male with distal median 
projection, usually well developed (inconspicuous 
in C. (Verbenapis). 80. S8 of male with two short, 
lateral projections between body and distal pro- 
jection, sometimes very small. 84. Gonostylus 
absent. 
Discussion. The genus Calliopsis is treated here 
in a new broad sense to include not only the 
subgenera Calliopsis s. str. , Perissander, Calliopsima 
and Verbenapis, traditionally included in the 
genus, but also Nomadopsis s. str., Macronomadop- 
sis, Micronomadopsis, Hypomacrotera, Liopoeum and 
two new subgenera Liopoeodes and Cerolwpoeum. 
This new concept of the genus Calliopsis is based 
on the following facts: 
Panurginae: Calliopsini and Allies 
223 
1 . The similarity between some complex geni- 
talic structures oi Nomadopsis (in the former sense, 
as a genus) and those of some species of Calliopsis 
(in the usual sense) suggests that some species of 
Nomadopsis are more closely related to some Calli- 
opsis than to other Nomadopsis. The cladogram 
(Fig. 30), however, still shows Nomadopsis (usual 
sense) as a monophyletic group, although the 
synapomorphies are not strong. Small ridges on 
the lateral part of the penis valve of males occur in 
Micronomadopsis, Hypomacrotera, Calliopsis s. str. , 
Perissander, Liopoeodes and Ceroliopoeum. This fact 
may indicate a close relationship among these six 
taxa. 
2. Chemosystematic studies indicate that, looking 
at the Dufour's gland lipids, the Callwpsis-Nomadop- 
sis- Hypomacrotera group ("paraffin bees") appears to 
be monophyletic (Cane, 1983). The cladogram of 
this analysis shows one species of Calliopsis (usual 
sense) as being more closely related to one species of 
Nomadopsis than to two other species of Calliopsis. 
3. Lack of strong characters to diagnose the 
traditional genera. Calliopsis and Nomadopsis (in the 
old sense) have been characterized mainly by the 
presence of hair bands and of yellow integumental 
marks on the metasomal terga respectively. Both 
characters appear independendy in other Pan- 
urginae. Larval characters also do not differentiate 
these groups (Rozen, 1966). 
4. Existence of C (Liopoeodes) xenopous n. sp., a 
species that does not fit within taxa already estab- 
lished. Maintaining generic rank for Calliopsis, No- 
madopsis and Hypomacrotera leads to proliferation of 
genera difficult to separate from one another. 
5. There is litde morphological differentiation 
among the subgenera except for Liopoeodes and 
Verbenapis, which appear in the cladogram as having 
more apomorphies than the rest of the subgenera. 
There is no constant apomorphy for C. (Microno- 
madopsis), which therefore may be paraphyletic. 
6. The cladistic analysis indicates that Calliopsis s. 
1. is a natural group. The cladogram shows six 
synapomorphies (62, 81, 84, 85, 111, 115) that 
make this group monophyletic. The strongest char- 
acters are: 62, 81, 84, and 115. 
Key to the Subgenera of the Genus Calliopsis 
1. Males 2 
— Females 12 
2. Pubescence of basitarsus 3 on dorsal margin (at least in part) about as long as basitarsus or 
longer (Fig. 16A). Apical area of labrum clearly convex, almost glabrous except margin; 
labrum with no defined ridge separating basal and apical areas (Fig. 15H) Liopoeum 
— Pubescence of basitarsus 3 on dorsal margin about half as long as basitarsus or shorter. Apical 
area of labrum hairy, rather flattened; labrum with ridge or carina separating basal and apical 
areas (Fig. IF) [except entire labrum flat and pilose in C. (H.) subalpina] 3 
3. T2-5 at least laterally with premarginal hair bands 4 
— T2-5 without premarginal hair bands 7 
4. Subantennal area black. Metanotum laterally without velvet area. Propodeal triangle 
basally smooth, concave medially Verbenapis 
— Subantennal area yellow. Metanotum laterally with area (sometimes reduced to a small and 
narrow strip) of dense velvety hairs. Propodeal triangle basally rugose or at least slightly 
roughened 5 
5. Clypeus (lateral view) clearly protuberant (Figs. IE, 8B). Metanotum laterally with 
conspicuous patch of velvet-brown hairs Calliopsis s. str. 
— Clypeus (lateral view) almost flat. Metanotum laterally with small patch of velvet-white hairs 
. . . ' '. 6 
6. Labrum with basal area delimited by strong carinate ridge. Basitarsus 2 somewhat shorter 
than 3. Propodeal triangle basally with strong striae; striated basal part delimited by strong 
transverse ridge Calliopsima 
— Labrum with basal area delimited by weak ridge. Basitarsus 2 much longer than 3. Propodeal 
triangle basally with weak striae, posteriorly not delimited by strong transverse ridge .... 
Perissander 
7. Metasoma without yellow marks 8 
— Metasomal terga usually with interrupted or complete yellow bands 10 
8. Antennal scape robust (Fig. 24F). Pterostigma (maximum width between costa and Rs) 
more than twice as broad as prestigma, side within marginal cell clearly convex (Fig. 2D) 
Ceroliopoeum 
224 The University of Kansas Science Bulletin 
— Antennal scape normal. Pterostigma less than twice as broad as prestigma, side within 
marginal cell straight or nearly so (Fig. 2E) 9 
9. Inner orbits subparallel. Marginal cell about twice as long as submarginal cell 2 + 3 or 
longer. Tarsus 3 with tarsomeres 2-5 unmodified Hypomacrotera 
— Inner orbits convergent below (Fig. 22H). Marginal cell less than twice as long as 
submarginal cell 2 + 3. Tarsus 3 with basitarsus and tarsomeres 2-4 asymmetrical; distitarsus 
widened medially (Fig. 22K) Liopoeodes 
10. Tarsi 1, 2 with tarsomeres slender. Tarsus 3 often with tarsomeres 2-4 broadened laterally, 
asymmetrical (Fig. 211) Micronomadopsis 
— Tarsi 1-3 with tarsomeres broadened distally, symmetrical 11 
11. Tibia 3 about twice as long as basitarsus 3. S4 clearly produced in middle of distal margin. 
S5 with well-developed median projection on distal margin (Fig. 18E) Nomadopsis 
— Tibia 3 somewhat longer than basitarsus 3. S4 inconspicuously produced in middle of distal 
margin. S5 distally with short median projection (Fig. 20H) Macronomadopsis 
12. Pterostigma (maximum width between costa and Rs) more than twice as broad as 
prestigma, margin within marginal cell clearly convex. Middle femur with ventral comb less 
than half length of femur Ceroliopoeum 
— Pterostigma less than twice as broad as prestigma, margin within marginal cell straight or 
nearly so. Middle femur with ventral comb about half length of femur 13 
13. T2-4 with premarginal hair bands at least laterally 14 
— T2-4 without premarginal hair bands 19 
14. Integument black (with some slight blue but no yellow). Wing venation mostly dark. Inner 
orbits subparallel (Fig. 23B). Tl-2 polished, scarcely punctate Liopoeodes 
— Integument with at least some yellow marks. Wing venation mostly yellowish. Orbits 
convergent below. Terga with rather abundant punctation 15 
15. Middle tibial spur with coarse teeth distally (Fig. 16F). Tibia 3 with keirotrichia on inner 
surface at base and at apex only (Fig. 16D). S6 distally with oblique patch of hairs beside 
midline (Fig. 16G) Lwpoeum (part) 
— Middle tibial spur with teeth somewhat coarse distally (Fig. 14K). Tibia 3 with keirotrichia on 
most of inner surface or absent toward ventral border. S6 distally with hairs forming a curved 
fringe 16 
16. Tarsus 1 on inner surface with hairs sparse, mostly stiff, curved and not tapered at apices 
(Fig. 14H). Metanotum laterally without patch of white, velvety hairs Verbenapis 
— Tarsus 1 on inner surface with hairs dense, unmodified. Metanotum laterally with small 
(sometimes inconspicuous) patch of white, velvety hairs 17 
17. Clypeus (lateral view) protuberant 1/3 width of eye or more. Paraocular area with lowest 
part noticeably wider than at level of antennal sockets, usually swollen on lower inner corner 
Calliopsis s. str. 
— Clypeus (lateral view) protuberant about 1/4 to 1/9 width of eye. Paraocular area with lowest 
part almost as wide as at level of antennal sockets, slightly convex 18 
18. Propodeum mostly punctate, at base conspicuously rugose and triangle delimited by strong 
ridge or carina. Metanotum laterally with patch of white velvety hairs clearly visible Calliopsima 
— Propodeum posteriorly with extensive impunctate area, at base usually little or not very 
strongly rugose, triangle delimited by rather rounded ridge. Metanotum laterally with 
inconspicuous (difficult to see) patch of white, velvety hairs Perissander 
19. Middle tibial spur with most teeth much longer than those of hind tibial spurs (Fig. 16F). S6 
distally with oblique patch of hairs beside midline (Fig. 16G) Lwpoeum (part) 
— Middle tibial spur with most teeth only slightly longer than those of hind tibial spurs, fine or 
somewhat coarse (Fig. 171). S6 distally with hairs forming a curved fringe (Fig. 17H) 20 
20. Propodeal triangle polished. Metasoma with no yellow markings Hypomacrotera 
— Propodeal triangle at least slightly rugose basally. Metasoma usually with complete or 
interrupted yellow integumental bands 21 
21. Body length less than 10 mm. Middle tibial spur with 4 teeth on distal half (Fig. 21H) 
Micronomadopsis 
— Body length 10 mm or longer. Middle tibial spurs with more than four (coarse or fine) teeth on 
distal half (Figs. 19D, 20J) Macronomadopsis and Nomadopsis 
Panurginae: Calliopsini and Allies 
225 
Subgenus Calliopsis Smith s. str. 
(Figs. lA, B, D-F; 6B. 8B. 11, 12, :5()) 
Calliopsis Smith, 1853: 128. Type species: Calliopsis 
andreniformis Smith, 18,53 (bv desitrnation of Ash- 
mead,' 1899: 85). 
Diagnosis. Glossa somewhat shorter than pre- 
mentum. Metasomal terga black, vvithdistal hair 
bands. Axilla and rnctanotiun laterally with dense 
patches ot \el\et\' hairs. Male. Basitarsus 2 some- 
what longer than 1 and 3. 
Description. 1. Length 5-8 mm. 3. Lower half of 
face, some spots on thorax (pronotum dorsally, 
pronotal lobe, tegula) and legs yellow in male, 
yellow much reduced in female. 4. Metasoma 
black. 5. Pubescence short, especially short and 
appressed on metasoma, forming premarginal 
bands on terga. 7. Punctures fine, well marked, 
dense on thorax. 8. Head broader than long and 
broader than thorax. 1 1 . Glossa somewhat 
shorter than prementum. 13. Segment 1 of labial 
palp longer than 2-4 together. 16. Maxillary 
blade longer than prepalpal part of galea. 17. 
Galeal comb of 17-26 bristles. 20. Labrum 
slightly protuberant, about twice as broad as 
long; basal area depressed, its distal margin a 
rounded ridge; labral apex somewhat inflexed. 
Figure 1 1 . Calliopsis (Calliopsis) andreniformis 
Smith. Male: A, B, Genitalia, dorsal, ventral and 
lateral views; C, 85, ventral view; D, Penis valve; 
E, F, S7 and 8, dorsal and ventral views; G, 
Metasomal tip, ventral view; H, S6, ventral view. 
pvr = penis valve ridges. 
27. Clypeus more than twice as broad as long, 
protuberant (about 1/3 width of eye in lateral 
view); distal margin with a rather small, pointed 
projection near lateral margin of labrum in male, 
rounded and bigger in female. 30. Inner suban- 
tennal suture curved. 31. Subantennal area wider 
than inner suture length and than antennal 
socket. 34. Antennal socket below middle of face. 
36. Antennal flagellum of male unmodified (Fig. 
12G); length variable (shorter than head as in C. 
andreniformis or longer); flagellomere 1 longer than 
2 (almost twice as long or less) and about as long 
as broad. 37. Lower mesal paraocular area 
slightly convex to strongly swollen. 38. Facial 
fovea clearly visible, very narrow (almost linear). 
40. Ocelli just above dorsal orbital tangent. 42. 
Gena of male (lateral view) narrower than eye, 
broader dorsally than ventrally. 43. Pronotum 
dorsally with preapical ridge rounded, developed 
only laterally, not strong. 43a. Axilla of male 
(lateral to scutellum) with patch of velvety, brown 
hairs. 43b. Metanotum laterally with velvety 
hairs forming a conspicuous brown patch in 
males, and small white patch in females. 44. 
Mesepisternum with anterior flat area reduced. 
45. Pre-episternal groove deep, narrow, punctate. 
46. Pterostigma somewhat longer than and about 
as wide as prestigma; side basal to vein r sub- 
parallel to costa, that within marginal cell 
straight. 47. Marginal cell narrowly truncate at 
apex, little longer than distance from its apex to 
wing tip. 49. Submarginal cell 1 about as long as 
cell 2-1-3 (or inconspicuously longer). 51. First 
recurrent vein not meeting first transverse cubital 
but sometimes very close to it. 52. Forewing with 
cu-v about three times longer than second ab- 
scissa M -I- Cu. 53. Hind wing with cu-v 1/4 to 1/5 
as long as second abscissa M-I-Cu. 54. Dorsal 
surface of propodeum shorter than metanotum. 
55. Propodeal triangle striate basally, not delim- 
ited by ridge or carina posteriorly. 57. Basitarsus 
1 about seven to eight times longer than broad in 
male and about four to five times in female; 
tarsomeres 2-4 unmodified. 59. Middle tibial 
spur with minute and fine teeth, almost straight 
and longer than half length of basitarsus in 
female, slightly curved at apex and about 1/3 as 
long in male. 60. Basitarsus 2 of male somewhat 
longer than 1 and 3; in female basitarsus 1 as long 
as 2 and 3, tarsomeres 2-4 unmodified. 61. Tibia 
3 of female about twice as long as basitarsus 3, 
inner surface with keirotrichia except along ven- 
tral half; male tibia 3 with keirotrichia on most of 
inner surface. 62. Tibial scopa of medium-sized, 
sparse, and mostly simple hairs. 63. Hind tibial 
spurs toothed similarly to middle one, somewhat 
curved, outer shorter than inner. 64. Basitibial 
plate of male well defined. 65. Basitarsus 3 of 
female with moderate apical projection; tarso- 
meres 2-4 unmodified. 66. Claws deeply cleft 
with rami subequal in male, inner ramus much 
shorter than outer in female. 67. Metasoma in 
male wider than thorax. 69. T2-5 of male with 
226 
The University of Kansas Science Bulletin 
gradulus posterolaterally reaching to middle of 
postgradular area at most, not strongly carinate 
and with postgradular depression rather shallow, 
narrow; posterior marginal areas of Tl-5 in male 
and Tl-4 in female each with hair band; lateral 
fovea of T2 (both sexes) slightly or well de- 
pressed, small and narrow in male, larger and 
wider in female. 70. Pygidial plate of male pre- 
sent. 71. T7 of female rather square, with no 
large dorsal, proximal expansion (Fig. 12B). 72. 
T8 of male similar to that oi Callwpsima. 73. SI -5 
of male with hairs mostly directed posteriorly; S6 
pilose only on distal projections, mostly directed 
outward. 73a. Sl-5 of female with hairs short, 
simple, appressed proximally; longer, branched, 
not appressed on premarginal and marginal areas 
medially. 75. S5 of male with a long and acute 
median apical projection, densely pilose at apex 
(Fig. IIC). 77. S6 of male distally with deep 
median emargination between two tapered pro- 
jections (Fig. IIH). 78. S6 of female with basal 
sclerotization almost completely free from ster- 
num; duplication thin, attached to a distal hard- 
ened area of sternum; distal margin produced in 
middle; S6 with distal hairs forming dense, some- 
what curved band (Fig. 12C). 79. S7 of male with 
two distal (almost transverse), elongate projec- 
tions (complex at base); proximal arms forming a 
V (Fig. HE). 80. S8 of male with distal projection 
almost rectangular, with two small laterodistal 
lobes; body robust with two lateral curved and 
acute projections distally and with a weak median 
longitudinal ridge dorsally (Fig. IIF). 83. Gono- 
coxites globose, distally (ventral view) connected 
to one another by weakly sclerotized cuticle. 85. 
Volsellae attached to one another by slightly 
sclerotized cuticle, with no denticles. 86. Penis 
valves complex (with projections, folds, etc.), 
tapered distally, with minute ridges laterally (ven- 
Figure 12. Calliopsis (Calliopsis) andreniformis 
Smith. Female: A, S5, ventral view; B, T7; C, S6, 
dorsal and ventral views; D, Sting; E, Head, 
frontal view. Male: F, Head, frontal view; G, 
Antenna. 
tral view) (Fig. 11 A, D), fused dorsally by a 
narrow bridge. 87. Penis much shorter and nar- 
rower than valve, clearly separated from penis 
valve, somewhat sclerotized. 88. Sting not reach- 
ing stylus apex (Fig. 12D). 
Comments. Calliopsis s. str. contains 12 described 
species (Shinn, 1967). The species that I studied 
in detail were: C. (C.) andreniformis Smith and C. 
(C.) hondurasica Cockerell. 
Discussion. Shinn (1967) indicated that Calliop- 
sis s. str. is most closely related to Perissander. This 
has been confirmed by the cladistic analyses 
presented here. 
Distribution. This subgenus occurs from Pan- 
ama to Canada, and from the eastern United 
States to Utah, Nevada, California, and Baja 
California Sur. 
Subgenus Perissander Michener 
(Figs, similar to 11, 12; 30) 
Perissander M\cheT\ev, 1942: 275. Type species: Calliopsis 
anomoplera Michener, 1942 (monobasic and original 
designation). 
Diagnosis. Similar to Calliopsis s. str. but differ- 
ing as follows: Glossa much shorter than premen- 
tum. Only metanotum with narrow patch of 
dense hairs laterally. Propodeal triangle delimited 
by rounded ridge posteriorly. Male. Forewing 
sometimes brown at apex. Basitarsus 2 much 
longer than 1 or 3. 
Description. As in Calliopsis s. str. except: 3. 
Wing tip of male sometimes brown as in male of 
Hypomacrotera. 4. Metasoma sometimes fully or 
partially reddish. 7. Punctures of metasoma of 
male somewhat denser than those of Calliopsis s. 
str. 11. Glossa much shorter than prementum. 
13. Segment 1 of labial palp about as long as 2-4 
together. 16. Maxillary blade tapered abruptly at 
apex, slightly shorter than prepalpal part of galea. 
17. Galeal comb of 11 bristles. 20. Labrum of 
male sometimes almost flat [e.g., in C. (P.) 
anomoptera] with basal area narrow and with distal 
margin a weak ridge. 27. Clypeus about three 
times broader than long, little protuberant (about 
1/5 or 1/6 width of eye in lateral view); distal 
margin with a rounded projection near lateral 
margin of labrum, little developed in male. 30. 
Inner subantennal suture angulate. 31. Suban- 
tennal area about as wide as antennal socket. 36. 
Antennal flagellum of male as long as or slightly 
shorter than head; flagellomere 1 about as long as 
2. 37. Lower mesal paraocular area somewhat 
convex. 38. Facial fovea of variable shape. 43b. 
Metanotum with narrow patch of dense hairs 
laterally (usually hidden by base of hind wing). 
45. Pre-episternal groove shallow. 46. Ptero- 
stigma about as long as prestigma. 47. Marginal 
cell about as long as to much longer than distance 
from its apex to wing tip. 51. First recurrent vein 
not as close to first transverse cubital as in C. 
Panurginae: Calliopsini and Allies 
227 
andreniformis Smith. 52. Ii)n_\Ning with c u-\ abmii 
half to twice as long as M + Cu. 54. Dorsal 
surface of propodcum longer than nietanotuni. 
55. Propodeal triangle basally with weak striae, in 
middle delimited by ridge posteriorly. 59. Middle 
tibial spur with few and sparse teeth in female and 
about 1/5 as long as basitarsus 2 in mak-. bO. 
Basitarsus 2 of male much longer than 1 and 3. 
61. Tibia 3 of female less than twice as long as 
basitarsus 3. 63. Hind tibial spurs generally with 
fine teeth (sometimes somewhat coarse). 66. 
Claws in male with rami short, subequal on legs 1 
and 3, inner shorter than outer on leg 2. 75. S5 of 
male with distal, median projection sometimes 
somewhat rounded at ape.x. 78. S6 of female with 
distal margin concave in the middle. 80. S8 of 
male with only one small median lobe at apex. 85. 
\'olsellae fully fused to each other distally, con- 
nected proximally by weakly sclerotized cuticle. 
Comments. There are seven described species in 
this subgenus (Shinn, 1967). I studied in detail: 
C. (P.) anomoptera Michener, C. (P.) gilva Shinn 
and C. (P.) rogen Shinn. 
Discussion. The subgenus Perissander, as noted 
above and in Figure 30, is closely related to 
Calliopsis s. str. and both together are the sister 
group of the subgenera Liopoeum, Hypomacrotera, 
MicTonomadopsis, Macronomadopsis, Nomadopsis s. 
str., Ceroliopoeum, and Liopoeodes. The metanotal 
velvet area of males, so well developed in Calliop- 
sis s. str., is small, less dense, and rather difficult 
to see, although still present, at least in the species 
that I studied. This is in disagreement with Shinn 
(1967) who has stated that this pilose area is 
absent in this subgenus. 
Distribution. Perissander is found in the south- 
western United States and northwestern Mexico, 
including Baja California (Shinn, 1967). 
Subgenus Calliopsima Shinn 
(Figs. 13, 30) 
Calliopsima Shinn, 1967: 834. Type species: Calliopsis 
Tozeni Shinn, 1967 (original designation). 
Diagnosis. Differs from Calliopsis s. str. as fol- 
lows: Glossa longer than prementum. Clypeus 
flattened. Labrum with distal margin of basal 
area a strong carina. Metanotum basilaterally 
with narrow patch of dense hairs. Basitarsus 2 
shorter than 3. Male. Labral apex short. 
Description. Agrees with description of Calliopsis 
s. str. except: 1. Length 7-8 mm. 5. Pubescence 
denser on metasoma. 7. Punctures dense on 
metasoma of both sexes. 11. Glossa longer than 
prementum. 13. Segment 1 of labial palp about 
twice as long as 2-4 together. 16. Maxillary blade 
longer than prepalpal part of galea. 17. Galeal 
comb of 13 bristles. 20. Labrum of male with 
basal area large and slightly depressed; distal 
margin of basal area (both sexes) usually a strong 
Figure 13. Calliopsis (Calliopsima) rozeni Shinn. 
Male: A, B, Genitalia, dorsal, ventral and lateral 
views; C, D, S6 and 5, ventral views; E, S8, 
dorsal and ventral views; F, T8; G, S7, dorsal and 
ventral views. Female: H, T7; I, Sting; J, S6, 
dorsal and ventral views. 
carina; apex of labrum very narrow in male, 
wider in female. 27. Clypeus flattened (about 1/9 
width of eye in lateral view); distal margin with 
projection near lateral margin of labrum 
rounded, similar to that oi Perissander. 31. Suban- 
tennal area about twice as wide as inner suture 
length and slightly wider than antennal socket. 
36. Antennal flagellum of male somewhat shorter 
than head; flagellomere 1 somewhat longer than 2 
and slightly wider than long. 37. Lower mesal 
paraocular area rather flat or slightly convex. 45. 
Pre-episternal groove rather wide and shallow. 
46. Pterostigma less than twice as broad as 
prestigma. 47. Marginal cell longer than in Calli- 
opsis s. str. 49. Submarginal cell 1 somewhat 
longer to shorter than cell 2-1-3. 51. First recur- 
rent vein clearly distant from first transverse 
cubital. 52. Forewing with cu-v longer than 
second abscissa M-I-Cu. 54. Dorsal surface of 
propodeum longer than metanotum. 55. Propod- 
eal triangle striate laterally, rugose medially, pos- 
teriorly delimited by carina. 57. Basitarsus 1 
about six to seven times longer than broad in 
male and about four times in female. 59. Middle 
tibial spur about half as long as basitarsus 2. 60. 
Basitarsus 2 in both sexes longer than 1 and 
shorter than 3. 61. Tibia 3 of female less than 
228 
The University of Kansas Science Bulletin 
twice as long as basitarsus. 62. Tibial scopa of 
medium-sized, dense, branched hairs. 69. T2-5 
of male with gradulus long and distinctly cari- 
nate; postgradular depression rather deep on 
T2-3; fovea of T2 clearly visible, in female only 
slightly bigger and not much wider than that of 
male. 71. T7 of female rather squared (Fig. 13H). 
75. S5 of male with median apical projection, 
rather short and clavate (Fig. 13D). 77. S6 of 
male with distal margin broadly produced medi- 
ally; with two short distal mesal lobes (Fig. 13C). 
79. S7 of male with distal area trianglar, laterally 
with a weakly sclerotized elongate projection; 
proximal arms forming a V (Fig. 13G). 80. S8 of 
male with distal projection long and narrow 
except arrowhead-shaped at apex; body of S8 
with latero-distal projections acute but very small 
(Fig. 13E). 85. Volsellae strongly developed (of 
different shape and much larger than in Calliopsis 
s. str. and Perissander), fused to each other prox- 
imally, free distally, with no denticles. 86. Penis 
valves long, rather simple, angulate at apex, 
attached to each other dorsally by reduced mem- 
branous area. 87. Penis long, narrower than 
valve. 88. Sting even shorter than in Calliopsis s. 
str. (much shorter than stylus) (Fig. 131). 
Comments. The subgenus Calliopsima contains 
15 species (Shinn, 1967). C. (C.) rozeni Shinn and 
C (C.) coloradensis Cresson were examined in 
detail. 
Discussion. The cladogram (Fig. 30) shows that 
the subgenus Calliopsima is the sister group of the 
other subgenera of Calliopsis s. 1. (based on male 
characters only) but is more closely related to 
Verbenapis than to Calliopsis s. str. and Perissander 
together. The relationships among these subgen- 
era agree with those stated by Shinn (1967). 
Distribution. According to Shinn (1967) Calliop- 
sima occurs from Canada to southern Mexico. 
Subgenus Verbenapis Cockerell and Atkins 
(Figs. 14, 30) 
Verbenapis Cockerell and Atkins, 1902: 44. Type species: 
Calliopsis verbenae Cockerell and Porter, 1899 (mono- 
basic). 
Diagnosis. Genitalic and associated sterna more 
similar to those of Calliopsima than to those of 
Calliopsis and Perissander. It differs from these 
subgenera as follows: Metanotum laterally with- 
out velvety hairs. Subantennal and supraclypeal 
areas black. Glossa as in Calliopsima. Male. Basi- 
tarsus 2 shorter than 3 (as in Calliopsima). S6 with 
median projection with minute median distal 
emargination. Female. Tarsus 1 on inner surface 
with rigid hairs, curved at blunt apices. 
Description. Agrees with that of Calliopsis s. str. 
except: 1. Length 7-8 mm. 3. Face with yellow 
marks restricted to clypeus and lower paraocular 
area; thorax black; legs with yellow reduced, 
Figure 14. Calliopsis (Verbenapis) verbenae Cock- 
erell and Porter. Male: A, B, Genitalia, dorsal, 
ventral and lateral views; C, S6, ventral view; D, 
E, S8 and 7, dorsal and ventral views; F, T8. 
Female: G, S6, dorsal and ventral views; H, 
Tarsus 1, inner view; I, T7; J, Sting; K, Spur, leg 
2. 
although extensive in male. 5. Tergal hair bands 
less dense in male. 7. Punctures sparser on thorax 
and metasoma. 11. Glossa longer than premen- 
tum as in Calliopsima. 13. Segment 1 of labial palp 
more than three times longer than 2-4 together. 
16. Maxillary blade more than twice as long as 
prepalpal part of galea. 17. Galeal comb of 18 
bristles. 20. Labrum of male similar to that of 
Calliopsima; in female, basal area well depressed 
and with weaker distal ridge. 27. Clypeus some- 
what protuberant (about 1/4 width of eye in 
lateral view); distal margin with a small rounded 
projection near lateral margin of labrum as in 
Perissander. 36. Antennal flagellum of male similar 
to that oi Calliopsima. 37. Lower mesal paraocular 
area somewhat convex as in Perissander. 38, 45, 
46, 47. Facial fovea, pre-episternal groove, 
pterostigma, and marginal cell as in Calliopsima. 
49. Submarginal cell 1 somewhat longer than cell 
2-1-3. 51. First recurrent vein somewhat distant 
from first transverse cubital. 52. Forewing with 
cu-v longer (about twice) than second abscissa 
M-I-Cu. 55. Propodeal triangle polished, shiny, 
with deep median concavity. 57. Basitarsus 1 
about eight times longer than broad in male and 
about seven times in female, inner surface with 
Panurginae: Calliopsini and Allies 
229 
cun'ed hairs in leniiilc; tarsoiiieres ot female 
(inner surface) with hairs as on basitarsus. 58. 
Femur of female with comb of hairs shorter than 
in Callwpsis s. str. 59. Middle tibial spur with 
teeth serrate (Fig. 14K), sparser than in Calliopsis 
s. str., similar to that of Perissander. 60. Basitarsus 
2 of male as in Calliopsvna; female with basitarsus 
2 as long as 1 and 3 . 61, 62 . Tibia 3 of female and 
tibial scopa similar to those of Calliopsima but 
scopa of sparser hairs with shorter branches. 67. 
Metasoma in male similar to that of Calliopsis s. 
str. but only slightly wider than thorax. 69. T2-5 
of male with gradulus short; that of T2 (both 
sexes) indistinct, difficult to see. 70. Pygidial plate 
of male present. 71. T7 of female as in Figure 
141. 72. T8 of male as in Figure 14F. 74. S4 of 
male with distal margin inconspicuously pro- 
duced in middle (difficvilt to see). 75. S5 of male 
with distal margin little produced medially, with 
no projection as in the other subgenera of Calliop- 
sis. 11 . S6 of male with a small median emargina- 
tion (Fig. 14C). 78. S6 of female similar to 
Calliopsis but duplication somewhat sclerotized 
(Fig. 14G). 79. S7 of male similar to that of 
Calliopsima but distal area shorter and with no 
lateral elongated projection (Fig. 14E). 80. S8 of 
male with distal projection elongate, clavate; 
proximal area almost rectangular with no projec- 
tions but small bumps (Fig. 14D). 85. Volsellae 
almost fully fused, well sclerotized medially, with 
no denticles. 86. Penis valves similar to those of 
Calliopsima but wider (especially in lateral view), 
dorsally fused to each other through a narrow 
sclerotized bridge. 87. Penis long, but somewhat 
wider than in Calliopsima, narrower than valve. 
88. Sting extremely short (Fig. 14J). 
Comments. The subgenus Verbenapis comprises 
four species (Shinn, 1967; Hurd, 1979). C. (V.) 
verbenae Cockerell and Porter, C. (V.) nebraskensis 
Crawford and C. (V.) michenen Shinn were avail- 
able for this study. 
Discussion. Verbenapis is, according to the clado- 
gram (Fig. 30), the sister group of the remaining 
subgenera of Calliopsis. On one hand it is the most 
closely related to Calliopsima; on the other hand it 
is the closest relative of Acamptopoeum. This is in 
disagreement with Shinn (1967), who stated that 
Calliopsima has a mixture of the characters found 
in Calliopsis s. str. and Acamptopoeum. 
Verbenapis, however, is the subgenus that pos- 
sesses the most characters agreeing with those of 
Acamptopoeum. Verbenapis has the propodeal tri- 
angle smooth, concave medially, and lacks the 
metanotal pad of hairs, as in Acamptopoeum. At the 
same time other characters seem transitional in 
Verbenapis. For example, S6 in Acamptopoeum has a 
very small distal, median emargination; in Verben- 
apis the same area is slightly sclerotized, with two 
small, weakly differentiated lobes laterally (Fig. 
14C), while in Calliopsima the lateral lobes of S6 
are well developed and the apical margin clearly 
concave. Finally, in the subgenera Calliopsis, Peris- 
sander, Liopoeum, Hypomacrotera, Micronomadopsis, 
Ceroliopoeum and Liopoeodes, S6 has elongate, acute 
lateral projections and a strongly concave margin 
medially. A similar gradual progression is shown 
by S8. 
Distribution. United States (except southeast 
and west) and Mexico (Shinn, 1967). 
Subgenus Liopoeum Friese 
(Figs. 15, 16, 30) 
Liopoeum Friese, 1906b: 176. Type species: Camplopoeum 
hirsutulum Spinola, 1851 (by designation of Sand- 
house, 1943: 564). 
Diagnosis. Close to Hypomacrotera but differing 
as follows: Pubescence long, especially on legs. 
Male. Tarsus 3 with tarsomeres 2-4 expanded 
laterally. Terga with or without yellow marks. S5 
with distal median projection long, truncate or 
tapered at apex. Female. Middle tibial spur with 
sparse and usually coarse teeth. Tibia 3 on inner 
Figure 15. Calliopsis (Liopoeum) hirsutulum 
(Spinola). Male: A, B, Genitalia, dorsal, ventral 
and lateral views; C, S8, apex, ventral view (hairs 
omitted); D, E, S7 and 8, dorsal and ventral 
views; F, S5, ventral view; G, S6, dorsal and 
ventral views; H, Labrum; I, T7 apex. 
230 
The University of Kansas Science Bulletin 
surface with keirotrichia dense at proximal and 
distal ends, sometimes present toward dorsal 
margin but sparse. S6 with distal patch of oblique 
hairs (in well-organized rows) at each side of 
produced median area. 
Description. 1. Length 5-8 mm. 3. Lower half of 
face, pronotum (sometimes) and legs with yellow 
marks; spot on paraocular area (both sexes) 
extending up beside orbit as in Acamptopoeum 
(male); spots of female usually more reduced on 
face than in male, absent on thorax. 4. Metasoma 
with or without yellow marks on terga. 5. Pubes- 
cence usually long on head and thorax and 
especially so on legs; usually shorter on meta- 
soma. 7. Punctures fine, usually much sparser on 
metasomal terga. 8. Head wider than thorax 
(about as wide as thorax in female of C. (L). 
trijasciata (Spinola) and an undescribed Chilean 
species from Bio-Bio area). 11. Glossa longer 
than half length of prementum. 13. Segment 1 of 
labial palp usually slightly longer than 2-4 to- 
gether. 14. Segment 2 of labial palp less than 
twice as long as 3. 16. As in Calliopsis s. str. 17. 
Galeal comb of 16-19 bristles. 20. Labrum sim- 
ilar to that of Calliopsis s. str. but with no well- 
defined ridge (Fig. 15H). 27. Clypeus more than 
twice as broad as long and somewhat to slightly 
protuberant (1/3 to 1/6 width of eye in lateral 
view); distal margin with an acute or rounded 
projection near lateral margin of labrum. 30. 
Inner subantennal suture angulate or strongly 
curved. 31. As in Calliopsis s. str. 34. Antennal 
socket about middle of face in female, below 
middle in male. 36. Antennal flagellum of male 
unmodified, longer than head; flagellomere 1 
about as long as 2 and about as long as broad 
(slightly longer in an undescribed Chilean spe- 
cies). 37. Lower paraocular area similar to that of 
Acamptopoeum. 38. Facial fovea variable (oval with 
borders not well delimited or narrow and well 
defined in an undescribed Chilean species). 40. 
Ocelli above dorsal orbital tangent except tangent 
crossing the middle ocellus in undescribed 
Chilean species. 42. Gena of male (lateral view) 
about as broad as, or broader than eye, dorsally 
wider than ventrally. 43, 44. As in Calliopsis s. str. 
45. Pre-episternal groove weak (sometimes seen 
only as a smooth strip). 46. Pterostigma longer 
than and slightly less than twice as broad as 
prestigma; otherwise as in Calliopsis s. str. 47. 
Marginal cell somewhat obliquely truncate at 
apex, much longer than distance from apex to 
wing tip. 49. Submarginal cell 1 longer than cell 
2-1-3. 51 . As in Calliopsis s. str. 52. Forewing with 
cu-v longer than second abscissa M-I-Cu. 53. 
Hind wing with cu-v about 1/3 as long as second 
abscissa M-(-Cu. 54. Dorsal surface of propod- 
eum about as long as metanotum. 55. Propodeal 
triangle basally smooth, microareolate, or with 
striae (sometimes depressed medially). 57. Basi- 
tarsus 1 five to six times longer than broad in 
male and about four times in female; tarsomeres 
2-4 unmodified. 59. Middle tibial spur of female 
with sparse and usually coarse teeth (Fig. 16F) 
(not very coarse in undescribed Chilean species), 
about as long as basitarsus 2 or slightly shorter; in 
male with very fine and dense teeth, about 1/2 
basitarsal length. 60. Basitarsus 2 of both sexes 
about as long as 1 and somewhat shorter than or 
as long as 3; tarsomeres 2-4 unmodified. 61. 
Tibia 3 of female less than twice as long as 
basitarsus 3, inner surface with keirotrichia at 
proximal and distal ends (almost absent medi- 
ally); male tibia 3 with keirotrichia on most of 
inner surface. 62. Similar to that of Calliopsis s. 
str. 63. Hind tibial spurs (both sexes) with fine 
teeth (denser in male), almost straight or clearly 
curved toward apex, outer slightly shorter than 
inner. 64. Basitibial plate of male distinguishable, 
though margins not strongly developed (weaker 
in undescribed Chilean species). 65. Basitarsus 3 
unmodified; tarsomeres 2-4 in male expanded 
laterally (at least slighdy), in female unmodified. 
66, 67. Similar to those of Calliopsis s. str. 69. 
T2-5 of male with gradulus posterolaterally short 
and weak (or almost absent) and postgradular 
depression narrow, rather shallow; posterior mar- 
ginal areas of Tl-5 in male and Tl-4 in female 
usually scarcely pilose; terga of female sometimes 
with distal hair bands; lateral fovea of T2 (both 
sexes) narrow, slightly depressed. 70. Pygidial 
plate of male recognizable, short, not well delim- 
ited laterally, truncate or rounded at apex (Fig. 
151). 71.T7offemaleasinFigure 16C. 72.T8of 
male similar to that of Hypomacrotera. 73. SI -5 of 
male with hairs directed posteriorly, very short 
(except longer distally), present mostly on lateral 
areas; on S6 mostly directed obliquely outward, 
on distal projections only. 73a. SI -5 of female 
with hairs as in Acamptopoeum. 75. S5 of male, on 
distal margin, with long median projection; pro- 
jection truncate or tapered and densely pilose 
apically (Fig. 15F). 77. Similar to that of Calliopsis 
s. str. 78. S6 of female as in Calliopsis s. str. but 
with distal hairs forming a dense oblique patch 
(several well-organized rows), absent on midline 
(Fig. 16G). 79. S7 of male with distal lateral 
projections intricate, with minute, dense hairs at 
apex and small brush of longer hairs medially; 
proximal arms forming a V (Fig. 15D). 80. S8 of 
male with distal projection with apical long hairs, 
deeply concave ventrally (Fig. 15C, E). 83. Gono- 
coxites globose, in ventral view fused to each 
other proximally, connected by membrane dis- 
tally; in dorsal view, proximally with a well- 
developed mesal projection. 85. Volsellae well 
developed, fused to each other by a narrow firm 
bridge distally, by membrane proximally, with 
few denticles. 86. Penis valves complex (intricate, 
with folds), tapered toward apices; dorsally fused 
to each other by small and narrow bridge. 87. 
Similar to that of Calliopsis s. str. 88. Sting 
elongate, reaching but not surpassing apex of 
stylus (Fig. 16B). 
Comments. The subgenus Liopoeum contains a 
few described species, most of which have been 
Panurginae: Calliopsini and Allies 
231 
Figure 16. Calliopsis (Liopoeum) hirsutulum 
(Spinola). Male: A, Tarsus 3. Female: B, Sting; C, 
T7; D, Tibia 3, inner view; E, S5, ventral view; 
F, Spur, leg 2; G, S6, dorsal and ventral views. 
placed erroneously in Camptopoeum (see Spinola, 
1851). {Camptopoeum is palearctic and not in the 
Calliopsini.) Friese (1908) segregated Liopoeum as 
a subgenus of Camptopoeum for some South Amer- 
ican species. However, they were still mixed with 
species of other genera. 
The described species now recognized as be- 
longing to the subgenus Liopoeum are: *C. (Lio- 
poeum) argentina (Jorgensen), *hirsutula (Spinola), 
*mendocina (Jorgensen), and *lrifasciata (Spinola) 
(all new combinations). 
Discussion. Among the subgenera of Calliopsis, 
Liopoeum is one of the most distinctive. Some 
species in this subgenus have hair bands on the 
metasomal terga of females, while others have 
tergal yellow marks, and no hair bands. This fact 
indicates that the presence or absence of these 
characters may not be useful or diagnostic at the 
subgeneric level. 
Distribution. C. (Liopoeum) occurs in Argentina 
and Chile. 
Subgenus Hypomacrotera Cockerell and Porter 
(Figs. 2E, 17, 30) 
Hypomacrotera Cockerell and Porter, 1899: 418. Type 
species; Hypomacrotera callops Cockerell and Porter, 
1899 (original designation). 
Diagnosis. Metasoma without yellow marks [red 
in C. (H.) subalpina (Cockerell)]. Pilosity short. 
Marginal cell narrowly truncate and much longer 
than distance from its apex to wing tip. 
Ptcrostigina with sides parallel. Male. Tarsus 3 
unmodified. Forewing tip dark brown. S5 distally 
with median elongate tapered projection. Female. 
Middle tibial spur with fine and dense teeth. S6 
distally with rounded fascia of dense hairs. 
Description. Agrees with description of subgenus 
Liopoeum except: 1. Length 5-8 mm. 3. Lower 
part of face (approximately 1/3) yellow (or 
whitish); apex of forewing in male brown. 4. 
Metasoma without yellow marks [red in C. (H.) 
subalpina[. 5. Pubescence in general short; much 
shorter on metasoma than on rest of body. 7. 
Punctures rather dense on metasomal terga. 13. 
Segment 1 of labial palp somewhat longer than 
2-4 together. 17. Galeal comb of 10-16 bristles. 
20. Labrum less than twice as broad as long; male 
with basal area and ridge in C. (H.) callops 
(Cockerell and Porter), similar to that oi Liopoeum 
but flat, in C. (H.) subalpina more extensively 
pilose; female with well-developed ridge. 27. 
Clypeus width more than three to five times 
broader than long, rather flattened (lateral view). 
30. Inner subantennal suture angulate. 31. Sub- 
antennal area broader than inner suture length 
and about as wide as (or slightly wider than) 
Figure 17. Calliopsis (Hypomacrotera) callops 
(Cockerell and Porter). Male: A, B, Genitalia, 
dorsal, ventral and lateral views; C, S8, tip (hairs 
omitted); D, S8, dorsal and ventral views; E, T8; 
F, S5, ventral view; G, S7, dorsal and ventral 
views; L, S6, ventral view. Female: H, S6, dorsal 
and ventral views; I, Spur, leg 2; J, Sting; K, T7. 
232 
The University of Kansas Science Bulletin 
socket. 34. Antennal socket appro.ximately at 
middle of face. 36. Flagellomere 1 slightly longer 
than broad (or about as wide as long). 37. Lower 
paraocular area similar to that of Acamptopoeum. 
38. Facial fovea oval, weakly differentiated. 39. 
Orbits convergent below, slightly so in C. (H). 
subalpina. 40. Ocelli just above dorsal orbital 
tangent. 42. Gena of male (lateral view) nearly as 
wide as eye; somewhat wider dorsally than ven- 
trally. 45. Pre-episternal groove weak, short, not 
reaching scrobal level. 46. Pterostigma longer 
than and about as wide as prestigma (or some- 
what less than twice as broad as prestigma). 47. 
Marginal cell narrowly and transversely truncate 
at apex, longer than distance from its ape.x to 
wing tip. 51. First recurrent vein rather close to 
first transverse cubital. 53. Hind wing with cu-v 
1/4 to 1/5 as long as second abscissa M + Cu. 54. 
Dorsal surface of propodeum similar to that of 
Liopoeum but somewhat variable. 55. Propodeal 
triangle basally smooth, depressed medially, 
glabrous. 57. Basitarsus 1 three times longer than 
broad in female. 59. Middle tibial spur of female 
slightly more than half length of basitarsus 2, with 
fine and dense teeth (Fig 171); of male toothed as 
in female but less than half basitarsal length. 60. 
Basitarsus 2 of male longer than 1 and somewhat 
shorter than 3; of female as in male but some- 
times basitarsus 2 about as long as 3. 61. Tibia 3 
of female on inner surface with keirotrichia except 
toward ventral margin. 62. Tibial scopa of sparse, 
rather short and apparently simple hairs on outer 
surface, inconspicuously branched and somewhat 
longer on margins. 63. Hind tibial spurs (both 
sexes) almost straight. 65. Tarsus 3 with no 
modifications. 69. Male with postgradular de- 
pression deep on T2, shallow on T3-5; posterior 
marginal areas of Tl-5 in male and Tl-4 in 
female pilose; terga (both sexes) with no hair 
bands; lateral fovea of T2 (both sexes) weakly 
depressed, somewhat wider in female. 70. Pygid- 
ial plate of male well developed in C (H.) sub- 
alpina, in C (H.) callops only distinguishable at 
apex. 71. T7 of female rather squared, with 
dorsal, proximal area not greatly expanded (Fig. 
17K). 75. S5 of male with median elongate 
projection tapered (Fig. 17F). 78. S6 of female 
with distal hairs longer than in Liopoeum and 
arranged in well-defined, curved band of dense 
hairs (Fig. 17H). 79. S7 of male with no patch of 
hairs forming a distal brush and with median 
small spine-like projection between proximal long 
arms (Fig. 17G) [absent in C. (77). subalpina]. 80. 
S8 of male with distal part ventrally concave only 
at base, with hairs shorter than in Liopoeum (Fig. 
17C, D). 83. Gonocoxite dorsally, on proximal 
area, with no mesal projection in C. (H.) callops 
[but present in C. (H). subalpina]. 86. Penis valves 
with ridges medially on outer margin (ventral 
view) (Fig. 17A). 
Comments. Only two species of this subgenus 
are recognized: *C. (H.) callops (Cockerell and 
Porter) and *C. (H.) subalpina (Cockerell) (new 
combinations), each with two subspecies (Hurd, 
1979). Because of size and metasomal color, the 
two species are superficially very different. None- 
theless, they are closely related. 
Until now Hypomacrotera has been poorly char- 
acterized, the major diagnostic character having 
been the infuscated area at the apex of the 
forewing in males. This character is not strong, 
since a brown spot on the apex of the forewing 
also appears in males of some species of Calliopsis 
s. str. , C. (Perissander), and in C. (Liopoeodes) 
xenopous, n. sp. The latter species has characters 
that are a mixture of those of Hypomacrotera, 
Liopoeum, and Calliopsis s. str. 
The cladogram (Fig. 30) shows that the sub- 
genus Hypomacrotera is more closely related to 
Liopoeum than to other subgenera of Calliopsis. 
Rozen (1970) indicates that Hypomacrotera coats 
pollen masses with a waterproof substance as do 
other Calliopsis. 
Distribution. C. (Hypomacrotera) occurs in the 
southwestern United States (Colorado, New 
Mexico, Arizona, California) and Mexico (Baja 
California and Sonora) (Hurd, 1979). 
Subgenus Nomadopsis Ashmead 
(Figs. 18, 19, 30) 
Nomadopsis Ashmead, 1898: 285. Type species: "Per- 
dila" zonalis Cresson, 1879 (monobasic and original 
designation), lapsus for Calliopsis zonalis Cresson, 
1879. 
Spinoliella subg. Claremontiella Cockerell, 1933: 25. Type 
species: Spinoliella euxanlha Cockerell, 1933 = Calliopsis 
zonalis Cresson, 1879 (monobasic and original desig- 
nation). 
Diagnosis. Metasomal terga with yellow bands. 
Sterna with hairs mostly short, straight, ap- 
pressed. Differs from Macronomadopsis as follows: 
Male. Tarsus 1 and 2 with tarsomeres 2-4 wid- 
ened distally. Tarsus 3 similar to 1 and 2, sym- 
metrical. S4 and S5 with distal produced area 
well developed. 
Description. 1. Length 7-12 mm. 3. Lower part 
of face, some areas of thorax (pronotum, tegula), 
and legs with yellow marks, reduced in female. 4. 
Metasoma with complete or interrupted yellow 
bands. 5. Pubescence in general short; dense, 
mostly very short and appressed on sterna. 7. 
Punctures mostly fine, well marked, smaller and 
usually dense on metasoma. 8. Head nearly as 
broad as thora.x, sometimes narrower in female. 
11. Glossa longer than prementum (flabellum 
present in several species). 13. Segment 1 of labial 
palp more than two to five times longer than 2-4 
together. 16. Maxillary blade more than twice as 
long as prepalpal part of galea. 17. Galeal comb 
of 18-23 bristles. 20. Labrum broader than long; 
basal area depressed; distal margin of basal area a 
rounded ridge; labral apex convex, inflexed. 27. 
Panurcinae: Calliopsini and Allies 
233 
Figure 18. Calliopsis (Nomadopsis) zonalis 
Cresson. Male: A, B, Genitalia, dorsal, ventral 
and lateral views; C, D, S8 and 7, dorsal and 
ventral views; E, F, S5 and 6, ventral view; G, 
T8; H, Tarsus 3. 
Clypeus more than twice as broad as long, pro- 
tuberant (usually 1/2 width of eye in lateral view 
in male and usually 1/3 in female, except almost 
flat in both sexes of C. (N.) edwardsii Cresson; 
distal margin with a rounded projection beside 
lateral margin of labrum. 30. Inner subantennal 
suture curved or somewhat angulate. 31. Suban- 
tennal area about as wide as inner suture length 
and at least slightly wider than antennal socket. 
34. Antennal socket somewhat below middle of 
face. 36. As in Calliopsis s. str. but flagellomere 1 
longer than 2, about as long as broad or little 
longer. 37. Lower mesal paraocular area some- 
what swollen to almost flat. 38. Facial fovea 
rather deep, narrow. 39. Orbits sometimes sub- 
parallel in female. 40. Ocelli fully or mostly above 
dorsal orbital tangent. 42. Gena of male (lateral 
view) little wider to little narrower than eye; 
somewhat widened dorsally. 43. As in Calliopsis s. 
str. 44. Mesepisternum with area facing anteri- 
orly rather reduced and slightly convex. 45. Pre- 
episternal groove narrow, shallow, punctate, not 
reaching scrobal level or at least not extending 
below it. 46. Pterostigma about as long as or 
longer than and about twice as wide as prestigma 
(sometimes less than twice); otherwise as in Calli- 
opsis s. str. 47. Marginal cell obliquely and 
broadly truncate, clearly longer than distance 
from its apex to wing tip. 49. Submarginal cell 1 
at least slightly longer than cell 2-f-3. 51. As in 
Calliopsis s. str. 52. Forewing with cu-v somewhat 
longer to three times longer than second abscissa 
M-I-Cu. 53. Hind wing with cu-v about 1/4 as 
long as second abscissa M -I- Cu. 54. As in Calliop- 
sis s. str. 55. Propodeal triangle finely striate 
basally. 57. Basitarsus 1 five to nine times longer 
than broad in male, two to five times in female; 
tarsomeres moderately widened distally in male. 
59. Middle tibial spur with fine, rather sparse 
teeth; straight and about half as long as (or longer 
than) basitarsus 2 in female; slightly curved at 
apex, usually less than half as long as basitarsus 
3, in male. 60. Basitarsus 2 of male longer than 1 
and about as long as 3; shorter than 1 and 3 in 
female; male tarsomeres 2-4 widened distally. 61. 
Tibia 3 of female less than twice as long as 
basitarsus 3, inner surface with keirotrichia 
scarce dorsally between patches near ends, absent 
ventrally; male tibia 3 with keirotrichia on most 
of inner surface of tibia. 62. As in Calliopsis s. str. 
63. Hind tibial spurs with fine teeth, inner spur 
longer than outer; slightly curved at apices. 64. 
As in Calliopsis s. str. 65. Basitarsus 3 unmodified, 
that of female gradually and slightly tapered 
toward apex; tarsomeres 2-4 similar to those of 
tarsi 1 and 2 (although shorter) in male (Fig. 
18H), narrower in female. 66, 67, 68. As in 
Calliopsis s. str. 69. T2-5 of male with gradulus 
posterolaterally short, usually not strongly cari- 
nate and postgradular depression shallow, very 
narrow; posterior marginal areas of Tl-5 in male 
and Tl-4 in female pilose; terga (both sexes) with 
no hair bands; lateral fovea of T2 (both sexes) 
slightly depressed, narrow in male, much wider 
in female. 70. Pygidial plate of male long, with 
well-defined margins. 71. T7 of female with 
dorsal, proximal area not greatly expanded (Fig. 
19B). 72. T8 of male as in Figure 18G. 73. Sl-5 
of male with hairs dense, mostly directed back- 
ward and appressed, usually longer on distal 
area; S6 with hairs only on distal projections. 
73a. Sl-5 of female with hairs similar to those of 
male. 75. S5 of male with a median, narrow, 
medium-sized projection on distal margin; pro- 
jection densely pilose at apex (Fig. 18E). 77. S6 of 
male with distal margin greatly projected medi- 
ally; projection with small distal emargination 
(Fig. 18F). 78. S6 of female with proximal lami- 
nar lobes sometimes with very shallow emargina- 
tion between them; basal sclerotization usually 
spine-shaped [widened at apex in C. (N.) zonalis 
Cresson, tapered in other species], mostly free 
from sternum; duplication somewhat sclerotized, 
attached to a distal hardened area of sternum; 
distal margin produced medially; S6 with distal 
hairs curved, forming a dense curved band (Fig. 
19A). 79. S7 of male distally rectangular, usually 
distally emarginate and with two short lateral 
lobes; proximal arms forming a V (Fig. 18D). 80. 
S8 of male with distal projection elongate, cla- 
vate; body of sternum widened medially, with a 
small, tapered projection at each side of the distal 
projection (Fig. 18C). 83. Gonocoxites globose, 
234 
The University of Kansas Science Bulletin 
Figure 19. Calliopsis (Nomadopsis) zonalis 
Cresson. Female: A, S6, dorsal and ventral views; 
B, T7; C, Sting; D, Spur, leg 2. 
ventrally connected by membranous area. 85. 
Volsellae large, connected by slightly sclerotized 
cuticle, with numerous denticles. 86. Penis valves 
with some folds, tapered distally, fused dorsally 
through a rather narrow sclerotized bridge. 87. 
Penis completely sclerotized except at apex (un- 
like any other Calliopsis, J. Rozen, personal com- 
munication), shorter and narrower than valve, 
clearly separated from valves. 88. See Calliopsis s. 
str. and Figure 19C. 
Comments. This subgenus contains eight de- 
scribed species (Rozen, 1958). I studied C. (N.) 
linsleyi (Rozen), puellae (Cockerell), and zonalis 
Cresson (the first two are new combinations). The 
females of this subgenus do not present features 
that distinguish them as a group from Macrono- 
madopsis and Micronomadopsis (Rozen, 1958). 
Discussion. Although Nomadopsis (usual sense) 
was confused with Spinoliella for a long time, there 
is no doubt that they belong to two different 
lineages among the Calliopsini. A similar state- 
ment was also made by Shinn (1967). Rozen 
(1951, 1958) and Shinn (1967) agree about the 
close relationship among the traditional Nomadop- 
sis, Calliopsis, Hypomacrotera, Liopoeum and 
Acamptopoeum, the latter being, in my analyses, a 
distinct genus, the sister group of Calliopsis. 
The cladogram (Fig. 30) shows that Microno- 
madopsis is the sister group of Nomadopsis s. str. 
and MacTonomadopsis. According to Rozen (1958), 
however, based on male genitalia and S7-8, 
Nomadopsis s. str. " ... is quite distinct from 
Macronomadopsis and Micronomadopsis and can not 
be affiliated more closely with one or the other." 
The male genitalia of Nomadopsis resemble those 
of Macronomadopsis (especially in shape of penis 
valve and volsella), more than those oi Microno- 
madopsis. The first two share some apomorphies 
that indicate that they are more closely related to 
one another than to Micronomadopsis. Further- 
more, the shape of the sterna of some Nomadopsis 
s. str. and Macronomadopsis is transitional between 
the two subgenera. 
Distribution. Western United States, southern 
British Columbia, and northern Mexico. 
Subgenus Macronomadopsis Rozen 
(Figs. 20, 30) 
Macronomadopsis Rozen, 1958: 93. Type species: Nomad- 
opsis michenen Rozen, 1958 (original designation). 
Diagnosis. Differs from Nomadopsis as follows: 
Male. Tarsus 1 and 2 with tarsomeres 2-4 con- 
spicuously widened distally. S4 with distal median 
projection inconspicuous. S5 with median projec- 
tion on distal margin small, slightly sclerotized. 
Description. Agrees with description oi Nomadop- 
sis except: 1. Length 8-12 mm. 3. Pronotum 
sometimes with no yellow marks. 13. Segment 1 
of labial palp somewhat longer than to twice as 
long as 2-4 together. 16. Maxillary blade longer 
than prepalpal part of galea. 17. Galeal comb of 
about 30 bristles. 20. Labrum usually broader 
than long [about twice as broad as long in C. (M.) 
zebrata Cresson]. 27. Clypeus usually more than 
twice as broad as long [little less than twice as 
broad in C. (M.) anthidia Fowler]; in lateral view 
protuberant 1/3 to 1/5 width of eye. 31. Suban- 
tennal area slightly wider than inner suture 
length. 34. Antennal socket about middle of face. 
37. Lower mesal paraocular area swollen to 
slightly convex. 38. Facial fovea shallow. 42. 
Gena of male (lateral view) somewhat narrower 
than eye, slightly widened dorsally. 45. Pre- 
Figure 20. Calliopsis (Macronomadopsis) zebrata 
Cresson. Male: A, B, Genitalia, dorsal, ventral 
and lateral views; C, S6, ventral view; D, E, S7 
and 8, dorsal and ventral views; G, S8, distal 
projection, ventral view (hairs omitted); H, S5, 
ventral view; I, Tarsus 3. Female: F, T7; J, Spur, 
leg 2. 
Panurginae: Calliopsini and Allies 
235 
episternal groove very short and weak, difficult to 
see. 46. Pterostigma less than twice as broad as 
prcstigma. 51. First recurrent vein never very 
close to first transverse cubital. 53. Hind wing 
with cu-v about 1/3 or 1/4 as long as second 
abscissa M + Cu. 54. Dorsal surface of propod- 
eum about as long as metanotuin. 57. Basitarsus 
1 six to seven times longer than broad in male and 
about four times in female: tarsomeres 2-4 of all 
tarsi distinctly expanded distally in male, slightly 
so in female. 59. Middle tibial spur with fine teeth 
as in Somadopsis but clearly longer than half 
length of basitarsus 2 in female, about half as long 
in male. 60. Basitarsus 2 of male shorter than 1 
and 3; about as long as 1 and shorter than 3 in 
female; tarsomeres 2-4 widened distally, more 
conspicuously so in male than in female. 61. 
Tibia 3 of female about twice as long as basitarsus 
3. 65. Tarsus 3 with tarsomeres 2-4 somewhat 
expanded distally (Fig. 201). 71. T7 of female 
rather square (Fig. 20F). 74. S4 of male incon- 
spicuously produced on distal margin medially. 
75. S5 of male with median projection on distal 
margin small, rather tapered at apex, slightly 
sclerotized (Fig. 20H). 77. S6 of male with a 
narrow, median distal emargination and 2 short, 
mesal projections (Fig. 20C). 78. S6 of female 
similar to that of Nomadopsis but with deep V- 
shaped emargination between proximal laminar 
lobes, basal sclerotization spine-like. 79. S7 of 
male similar to that of Nomadopsis but distal 
margin more deeplv emarginated and lateral 
lobes better defined (Fig. 20D). 80. S8 of male 
with distal projection narrowed toward apex (Fig. 
20E, G). 85. Volsellae connected by slightly 
sclerotized cuticle, with no denticles. 86. Penis 
valves strongly curved at apices (Fig. 20A). 87. 
Penis partially sclerotized, about as wide as valve. 
Comments. This subgenus contains five species 
(Rozen, 1958; Hurd, 1979). I studied C. (Macro- 
nomadopsis) micheneri (Rozen) and zebrata Cresson 
(only C. zebrata illustrated) (new combinations). 
Distribution. United States west of South Da- 
kota, Colorado, and New Mexico (Hurd, 1979). 
Subgenus Micronomadopsis Rozen 
(Figs. 21, 30) 
Micronomadopsis Rozen, 1958: 107. Type species: No- 
madopsis Jracta Rozen, 1958 (original designation). 
Diagnosis. Similar to but usually smaller than 
Nomadopsis and Macronomadopsis, from which it 
differs as follows: Male. Tarsus 1 with tarsomeres 
2-4 very thin. Tarsus 3, in several species, with 
tarsomeres 2-4 widened laterally, asymmetrical. 
Penis valve (in most species) as in Figure 21 A. 84 
and S5 as in Nomadopsis. Female. Middle tibial 
spur with only 4 teeth on distal half. 
Description. Agrees with description of Nomadop- 
sis except: 1. Length 5-10 mm. 8. Head broader 
than thorax in male, about as broad in female. 1 1 . 
Glossa somewhat longer than prementum to 
Figure 21. Calliopsis (Micronomadopsis) scutellaris 
Fowler. Male: A, Genitalia, dorsal and ventral 
views; B, S7, dorsal and ventral views; C, S6, 
ventral view; D, 88, dorsal and ventral views; I, 
Tarsus 3. Female: E, Sting; F, 86, dorsal and 
ventral views; G, T7; H, Spur, leg 2. pvr = penis 
valve ridges. 
much shorter. 13. Segment 1 of labial palp 
slightly shorter to about three times longer than 
2-4 together. 16. Maxillary blade length as in 
Macronomadopsis. 17. Galeal comb of about 14 
bristles. 27. Clypeus sometimes twice as broad as 
long, protuberant almost 1/3 to 1/7 width of eye. 
31. Subantennal area wider than inner suture 
length, about as wide as socket (sometimes 
wider). 34. Antennal sockets about middle of face 
as in Macronomadopsis . 36. Antennal flagellum of 
male as long as head or longer. 42. Gena of male 
(lateral view) narrower than eye, broadest in the 
middle. 43. Pronotum similar to that o{ Nomadop- 
sis, although ridge laterally less developed. 45. 
Pre-episternal groove short, sometimes indistinct. 
46. Pterostigma longer than prestigma. 51. First 
recurrent vein as in Macronomadopsis. 52. Fore- 
wing with cu-v usually slightly longer than second 
abscissa M + Cu. 53. Hind wing with cu-v as in 
Macronomadopsis. 54. Dorsal surface of propod- 
eum usually somewhat longer than metanotum, 
shorter in C. (Micronomadopsis) helianthi (Swenk 
and Cockerell). 55. Propodeal triangle weakly 
striate, sometimes smooth. 57. Basitarsus 1 six to 
seven times longer than broad in male, three to 
four times longer in female; tarsomeres 2-4 un- 
modified. 59. Middle tibial spur about 1/2 to 1/4 
as long as basitarsus 2 in male. 60. Basitarsus 2 of 
male at least slightly longer than 1, somewhat 
variable in relation to 3; of female as in Macrono- 
madopsis. 61. Tibia of female similar to that of 
Nomadopsis except with keirotrichia of inner sur- 
face denser dorsally in C. (M.) helianthi. 62. Tibial 
scopa sometimes of hairs with very short 
branches. 65. Basitarsus 3 unmodified, some- 
times very short; tarsomeres 2-4 from un- 
236 
The University of Kansas Science Bulletin 
modified to widened and asymmetrical in male of 
several species (Fig. 211), slender, unmodified in 
female. 67. Metasoma narrower to slightly wider 
than thora.x. 70. Pygidial plate of male sometimes 
shorter and wider than in Nomadopsis [e.g., in C. 
(M.) helianthi]. 71. T7 of female square (Fig. 
21G). 75. S5 of male with distal median projec- 
tion longer than in Nomadopsis (tapered or trun- 
cate at ape.x). 77. S6 of male with distal margin 
deeply emarginate medially, with two elongate, 
tapered projections (Fig. 21C). 78. S6 of female 
similar to that oi Macronomadopsis. 79. S7 of male 
with distal area usually with lateral, almost hori- 
zontal expansions (Fig. 2 IB); sometimes with 
short, median, spine-shaped projection on prox- 
imal margin. 80. S8 of male somewhat similar to 
that oi Macronomadopsis (Fig. 21D). 85. Volsella 
similar to that oi Macronomadopsis. 86. Penis valve 
complex, similar to that of Liopoeum, Hypomacro- 
tera, Calliopsis, Perissander, Liopoeodes and Cerolio- 
poeum, with fine, minute ridges laterally (ventral 
view) as in Hypomacrotera, Calliopsis, Perissander, 
Liopoeodes and Ceroliopoeum (Fig. 21 A). 87. Penis 
very short, partially sclerotized, much narrower 
than valve. 88. Sting as in Figure 2 IE. 
Comments. Micronomadopsis contains 20 species 
(Rozen, 1958; Hurd, 1979). I studied C. (Micro- 
nomadopsis) helianthi (Swenk and Cockerell) 
[ = euphorbiae (Cockerell)], fracta (Rozen) (new 
combinations), and scutellaris Fowler. 
Rozen's (1958) species of uncertain position, 
C. boharti (Rozen), smithi (Rozen) and xenus 
(Rozen), were not assigned to any subgenus. 
Some of their characters fit with Macronomadopsis 
and others with Micronomadopsis. 
Distribution. This subgenus is widespread in the 
western part of the United States. 
Liopoeodes, new subgenus 
(Figs. 22, 30) 
Type species: Calliopsis (Liopoeodes) xenopous, new species 
(see Appendix). 
Diagnosis. Similar to Liopoeum except: Integu- 
ment mostly dark. Hairs of head and thorax 
rather brownish and erect. Male. Forewing tip 
dark as in Hypomactrotera. Hind tarsus with all 
segments modified. Female. Black, metasomal 
terga with apical white hair bands. 
Description. As in Liopoeum except: 1. Length 
6-8 mm. 3. Lower half of face (reduced spots) 
and forelegs of male with yellow marks; female 
without yellow. 4. Metasoma without yellow 
areas. 5. Pubescence darkened; on head and 
thorax mostly erect and clearly longer than that of 
metasoma. 7. Punctures especially fine and dense 
on metasomal terga. 14. Segment 2 of labial palp 
slightly longer than 3. 17. Galeal comb of 23 
bristles. 20. Labrum less than twice as broad as 
long; basal area with distal margin a well marked 
ridge. 27. Clypeus somewhat protuberant (1/2 to 
fe- ■^-'^j^.'-- 
Figure 22. Calliopsis (Liopoeodes) xenopous n. sp. 
Male: A, B, Genitalia, dorsal, ventral and lateral 
views; C, S5; D, S6; E, F S7 and 8, dorsal and 
ventral views; G, T8; H, Head, frontal view 
(hairs omitted); I, Antenna; J, Mandible; K, 
Tarsus 3. 
1/3 width of eye in lateral view); distal margin 
with well-developed projection near lateral mar- 
gin of labrum. 34. Antennal socket somewhat 
below middle of face in both sexes. 36. Antennal 
flagellum of male with flagellomere 1 slightly 
longer than broad. 37. Lower paraocular area 
slightly convex. 38. Facial fovea well defined, 
longer and broader in female than in male. 42. 
Gena of male (lateral view) narrower than eye. 
46. Pterostigma with margin within marginal cell 
slighdy convex. 49. Submarginal cell 1 somewhat 
shorter than cell 2-1-3. 51. First recurrent vein 
rather close to transverse cubital vein. 53. Hind 
wing with cu-v about 1/5 as long as second 
abscissa M-I-Cu. 55. Propodeal triangle basally 
microareolate, depressed medially, with fine and 
weak striae. 57. Basitarsus 1 about seven times 
longer than broad in male and almost six times in 
female. 59. Middle tibial spur of female with 
several well-defined and clearly separated teeth, 
slightly shorter than basitarsus 2; in male about 
1/3 basitarsal length. 60. Basitarsus 2 of male less 
than twice as long as 1 and 3; basitarsus 2 of 
female slightly longer than 1 and 3.61. Tibia 3 of 
female about twice as long as basitarsus 3, inner 
surface with keirotrichia at proximal and distal 
ends and toward upper margin. 63. Hind tibial 
Panurginae: Cali.iopsini and Allies 
237 
spurs with tine teeth, ahiiost straight in male, 
outer one somewhat curved at apex in female. 64. 
Basitibial plate of male clearly distinguishable. 
65. Basitarsus 3 of male modified distally; tarso- 
meres 2-4 asymmetrical; distitarsus modified 
(Fig. 22K). 66. Claws with rather short rami in 
male, especially on tarsus 3. 69. Tl-T) of male and 
Tl-4 of female with posterior marginal areas 
apparently glabrous; terga of female with distal 
hair bands; lateral fovea of T2 widened and 
slightly depressed, almost indistinguishable and 
ventral in position rather than lateral in male. 70. 
Pvgidial plate of male apparently absent. 71. T7 
of female as in Figure 23D. 72. T8 of male as in 
Figure 22G. 73. Sl-5 of male with hairs sparse, 
mostly on middle line. 75. S5 of male with 
median projection shorter than in Liopoeum; api- 
cal hairs of projection longer than in Liopoeum 
(Fig. 22C). 79. S7 of male with tapered projection 
between pro.ximal arms as in Hypomacrotera (Fig. 
22E). 80. SB of male with distal projection some- 
what convex ventrally (Fig. 22F). 83. Gonocox- 
ites, in dorsal view, without mesal proximal 
projection. 85. Volsellae with no denticles. 88. 
Sting rather short, not reaching stylus apex (Fig. 
23C). 
Comments. The subgenus Liopoeodes contains a 
single species from Argentina. The subgeneric 
name is slightly modified from one used by J. S. 
Moure on some 30-year-old identification labels 
found in the collection of the Snow Entomological 
Museum (KU). 
Discussion. The mosaic of features exhibited by 
this new subgenus (some of them also present in 
other calliopsine genera) is one more evidence 
that it and its close relatives should be treated as 
species of the genus Calliopsis. 
The presence of a darkened area at the apex of 
the forewing of the male suggests C. (Hypomac- 
rotera) and some species of C. (Perissander). On the 
other hand, because of other characters indicated 
above, Liopoeodes appears to be more like C. 
(Liopoeum). 
Etymology. The name of this subgenus is based 
on Liopoeum plus the Greek suffix -odes, indicating 
similarity to Liopoeum. 
Ceroliopoeum, new subgenus 
(Figs. 2D, 24, 30) 
Type species: Camptopoeum laelum Vachal, 1909. 
Diagnosis. Appearance similar to that of Lio- 
poeum but clypeus convex medially and with 
marginal projection (beside labrum) broadly 
rounded; labrum with distal margin of basal part 
a strong ridge, almost a carina; pubescence short 
and rather sparse. Male. Antennal scape stout. 
Basitarsus 2 very narrow, longer than basitarsus 3 
(at least slightly). Tarsus 3 with tarsomeres 2-4 
narrow, apices not expanded laterally. Female. 
Tibial spur 2 as long as basitarsus 2, sinuate, with 
small, well-separated teeth. 
Figure 23. Calliopsis (Liopoeodes) xenopous n. sp. 
Female: A, S5, ventral view; B, Head, frontal 
view (hairs omitted); C, Sting; D, T7; E, S6, 
dorsal and ventral views; F, Antenna. 
Figure 24. Calliopsis (Ceroliopoeum) laeta 
(Vachal). Male: A, Genitalia, dorsal and ventral 
views; B, C, S7 and 8, dorsal and ventral views; 
D, S6, ventral view; E, Head, frontal view; F, 
Antenna. Female: G, Head, frontal view; H, 86, 
dorsal and ventral views; I, T7; J, Sting, 
v = valve-like structure. 
238 
The University of Kansas Science Bulletin 
Description. Agrees with that of Liopoeum except: 
1. Length 5-6 mm. 3. Male with legs mostly 
yellow; female with yellow spots on pronotum (as 
in male), tibiae and tarsi. 5. Pubescence rather 
short, mostly very short and appressed on meta- 
soma. 8. Head about as wide as thorax. 17. 
Galeal comb of 12 bristles. 20. Labrum with 
distal margin of basal part a strong ridge, almost 
a carina; apical area flattened. 27. Clypeus some- 
what protuberant (1/3 width of eye in lateral 
view), with distal margin (frontal view beside 
labrum) broadly rounded; central area convex. 
34. Male with flagellomere 1 about half as long as 
2. 35. Antennal scape robust. 38. Facial fovea 
narrow, well delimited. 42. Gena of male (lateral 
view) about 1/2 as wide as eye. 46. Pterostigma 
somewhat more than twice as broad as prestigma; 
margin within marginal cell slightly convex. 52. 
Forewing with cu-v about as long as second 
abscissa of M + Cu. 55. Propodeal triangle micro- 
areolate, depressed medially and with very fine 
longitudinal striae. 57. Basitarsus 1 almost eight 
times longer than broad in male, about six times 
in female. 59. Middle tibial spur of female as long 
as basitarsus 2, sinuose, with fine, well-separated 
teeth. 60. Basitarsus 2 about as long as 1 and 
slightly longer than 3 in male, and about as long 
as 1 and 3 in female. 61. Tibia 3 of female on 
inner surface with keirotrichia dense at proximal 
and distal ends, sparser toward dorsal margin. 
62. Tibial scopa of sparse and minutely branched 
hairs. 65. Tarsus 3 unmodified. 67. Metasoma in 
male narrower than thorax. 69. Tl-5 of male and 
Tl-4 of female with posterior marginal area 
scarcely pilose; hairs somewhat denser on T4 of 
female; lateral fovea of T2 (both sexes) indis- 
tinguishable. 70. Pygidial plate of male clearly 
delimited laterally, rather truncate at apex. 71. 
T7 of female as in Figure 241. 73a. SI -5 of female 
with hairs sparser than in Lwpoeum and 
Acamptopoeum and mostly not appressed. 75. S5 of 
male with median projection tapered apically (as 
in Fig. 1 IG). 79. S7 of male with patch of small, 
sparse hairs medially on latero-distal projections 
(Fig. 24B). 80. S8 of male with distal projection 
convex ventrally (Fig. 24C). 83. Gonocoxites, in 
dorsal view, without proximal mesal projection. 
85. Volsella without denticles. 88. First valvula of 
sting with a minute expanded area, perhaps a 
rudiment of valve, and a very short, curved hair 
behind this structure (Fig. 24J). 
Comments. C. (Ceroliopoeum) laeta (Vachal) is the 
only known species of this subgenus. 
Discussion. In the cladogram (Fig. 30), Cerolio- 
poeum is the sister group of Hypomacrotera, Lio- 
poeum, Nomadopsis, Alacronomadopsis and Microno- 
madopsis together. 
Distribution. Argentina. 
Etymology. The name of the subgenus refers to 
the distinctive antennal flagellum and the likeness 
to Lwpoeum. It is from the Greek keraos, horned, 
plus Liopoeum. 
Genus Arhysosage Brethes 
(Figs. 6C, 8C, 25, 26, 30) 
Arhysosage Brethes, 1922: 121. Type species: Arhysosage 
johnsoni Brethes, 1922 = Camptopoeum ochraceum Friese, 
1908 (monobasic). 
Ruiziella Timberlake, 1952: 105 (preoccupied). Type 
species: Camptopoeum ochraceum Friese, 1908. 
Ruziapis Timberlake, 1952: 528 {lapsus lor Ru!:apis, and 
replacement lor Rmziella). 
Rutzapis Timberlake, 1953: 598 (emendation of 
Ru:tapis). 
Diagnosis. Related to Spinoliella and Callonych- 
ium, sharing with them the low position of the 
antennal sockets. It can be differentiated by the 
following characters: Orbits distinctly divergent 
below. Hind tibial spurs distinctly curved toward 
apices. Male. Body almost completely yellow. 
Mandible strongly curved and elongate, with 
preapical tooth and projection on upper margin. 
Description. 1. Length about 10 mm. 3. Most of 
head and thorax of male yellow; size of yellow 
areas of female variable. 4. Metasoma with yellow 
bands very extensive to largely incomplete. 5. 
Pubescence in general short; appressed hairs on 
most of dorsum of thorax and metasoma. 7. 
Punctures very fine and dense. 8. Head broader 
than long and broader than thorax. 11. Glossa 
longer than prementum, slender. 12. Paraglossa 
shorter than suspensorium. 13. Segment 1 of 
Figure 25. Arhysosage ochracea Brethes. Male: A, 
B, Genitalia, dorsal, ventral and lateral views; C, 
S8, dorsal and ventral views; D, Mandible; E, 
Labrum; F, S6, ventral view; G, S7, dorsal and 
ventral views. 
Panurginae: Calliopsini and Allies 
239 
labial palp less than twice as long as 2-4 together. 
16. Ma.xillary blade longer than prepalpal part of 
galea. 17. Galeal comb absent. 20. Labrum less 
than twice as broad as long, partially or fully 
pilose; flat in male, with slight transverse ridge 
(almost flat) in female. 25. Mandible of male 
strongly arcuate, upper margin with prebasal 
projection and also with preapical tooth (some- 
times very conspicuous) (Figs. 6C, 25D). 27. 
Clypeus more than four times broader than long 
and somewhat protuberant (1/3 width of eye, 
lateral view) in male; about three times broader 
than long and distinctly protuberant (about 1/2 
width of eye, lateral view) in female; distal mar- 
gin with distinct projection (variable in size) near 
lateral margin of labrum. 30. Inner subantennal 
suture angulate. 31 . Subantennal area wider than 
length of inner suture and than antennal socket. 
33. Tentorial pit almost at middle of outer suban- 
tennal suture. 34. Antennal socket far below 
middle of face (on lower third in male, somewhat 
higher in female). 36. Antennal flagellum of male 
unmodified, much shorter than head; flagello- 
mere 1 about as long as 2 or little longer and 
about as long as broad. 37. Lower mesal paraocu- 
lar area slightly conve.x. 38. Facial fovea deep, 
narrow. 39. Orbits strongly divergent below in 
male, slightly so in female. 40. Middle ocellus 
below dorsal orbital tangent, lateral ocelli above. 
41. Vertex convex. 42. Gena of male (lateral 
view) wider than eye, slightly wider dorsally than 
ventrally. 43. Pronotum with dorsal preapical 
ridge rounded, strong in male, weak in female. 
44. Mesepisternum with flattened area facing 
anteriorly rather reduced. 45. Pre-episternal 
groove only distinguishable above scrobal level, 
continued downward as black line (difficult to see 
on dark integument). 46. Pterostigma longer than 
and slightly wider than prestigma; side basal to 
vein r subparallel to costa, that within marginal 
cell straight. 47. Marginal cell obliquely and 
broadly truncate at apex, longer than distance 
from apex to wing tip (slightly so in female). 49. 
Submarginal cell 1 about as long as cell 2 -I- 3 or 
longer. 51. First recurrent vein clearly distant 
from first transverse cubital. 52. Forewing with 
cu-v as long as or longer than second abscissa 
M-I-Cu. 53. Hind wing with cu-v little less than 
1/2 to 1/3 as long as second abscissa M -I- Cu. 54. 
Dorsal surface of propodeum slightly longer than 
metanotum. 55. Propodeal triangle basally with 
fine striae, depressed medially. 57. Basitarsus 1 
about five times longer than broad in male and 
four times longer in female; tarsomeres 2-4 un- 
modified. 59. Middle tibial spur about half as 
long as basitarsus 2 or somewhat longer, dis- 
tinctly curved apically, finally serrate (apical teeth 
larger than those at base). 60. Basitarsus 2 about 
as long as 1 and shorter than 3, tarsomeres 2-4 
unmodified. 61. Hind tibia of female somewhat 
longer than basitarsus 3; inner surface with kei- 
rotrichia in patch at base and apex, sparse or 
absent toward dorsal margin, absent ventrally 
(Fig. 26C); male tibia 3 with keirotrichia on most 
of inner surface but sparser ventrally. 62. Tibial 
scopa of moderately dense and apparently simple, 
but minutely branched hairs. 63. Hind tibial 
spurs strongly curved toward apices, outer about 
as long as inner or somewhat longer; teeth small. 
64. Basitibial plate of male with margins well 
defined. 65. Tarsus 3 unmodified. 66. Claws 
deeply cleft, rami subequal in male, inner ramus 
much shorter than outer in female. 67. Metasoma 
in male wider than thorax. 69. T2-5 of male with 
gradulus posterolaterally long (surpassing middle 
of each tergum), strongly carinate (especially on 
T2-4) and with postgradular depression narrow, 
rather shallow; posterior marginal areas of Tl-5 
in male and Tl-4 in female pilose (minute hairs); 
terga (both sexes) with no hair bands; lateral 
fovea of T2 (both sexes) slightly depressed, small 
in male, larger in female. 70. Pygidial plate of 
male well developed, abruptly elevated and cari- 
nate laterally on distal part, apex slightly emargi- 
nate. 71. T7 of female not expanded dorsally but 
with a strong ventral proximal projection (Fig. 
26D). 72. T8 of male hexagonal (Fig. 26F). 73. 
Sl-6 of male with most of hairs minute, dense, 
appressed, and directed posteriorly, longer dis- 
tally. 73a. SI -5 of female pilose as in male but 
hairs somewhat longer and denser. 74, 75. S4 and 
5 of male with distal margins slightly and broadly 
concave medially. 77. S6 of male distally bilobed, 
with small median V-shaped emargination (Fig. 
25F). 78. S6 of female with basal spine-like 
sclerotization fused to sternum; lateral margin 
with strong curved ridge; duplication thin, at- 
tached to distal hardened area of sternum; distal 
margin concave medially; S6 apically with a well- 
defined, curved and dense band of curved hairs 
(Fig. 26B). 79. S7 of male distally with 2 rather 
short, finger-like lateral projections; proximal 
arms long and forming a U (Fig. 25G). 80. S8 of 
male with a median long distal projection clavate 
at apex, abruptly separated from basal part which 
has weak median ridge dorsally (Fig. 25C). 83. 
Gonocoxite short, squared, completely fused both 
dorsally and ventrally (Fig. 25A, B). 84. Gono- 
stylus short, finger-like, fused to gonocoxite. 85. 
Volsella apparently absent? (or completely fused 
to gonocoxite). 86. Penis valves long, with some 
folds, distally bent downward at right angle, 
tapered toward apex, dorsally fused to each other 
by small, narrow bridge. 87. Penis almost reach- 
ing apices of penis valves, proximally wider and 
fused to valve; distal half well sclerotized ven- 
trally. 88. Sting very short (not attaining stylus 
apex) (Fig. 26E). 
Comments. This genus contains three known 
species: A. *flava Moure, *germana Moure, 
*ochracea (Friese). 
Discussion. Arhysosage is, according to the clado- 
gram (Fig. 30), the sister group of Spinoliella and 
Callonychium. 
Distribution. Argentina. 
240 
The University of Kansas Science Bulletin 
Figure 26. Arhysosage ochracea Brethes. Male: A, 
T7, dorsal view; F, T8. Female: B, S6, dorsal and 
ventral views; C, Tibia 3, inner view; D, T7; E, 
Sting; G, S5, ventral view, hts = hind tibial spurs; 
pp — pygidial plate. 
Genus Spinoliella Ashmead 
(Figs. 7A, 27, 28, 30) 
Diagnosis. Closer to Callonychium than io Arhyso- 
sage. It can be differentiated from Callonychium as 
follows; Gena black. Antennal sockets on lower 
third of face or nearly so. Clypeus with lateral 
part (beside mandibular articulation) rather hori- 
zontal. Male. Basitibial plate a swollen and shiny 
area, not delimited by carina. Female. S6 nar- 
rowed and with patch of hairs distally. Sting 
short, not reaching stylus. 
Description. 1 . Length 4-9 mm. 3. Lower part of 
face (below antennal socket) and some areas of 
legs with yellow marks. Thorax black (sometimes 
with small yellow spots). 4. Metasomal terga with 
yellow laterally or forming bands. 5. Pubescence 
short, especially short and appressed on meta- 
soma. 7. Punctures in general fine. 8. Head 
broader than long and as broad as thorax or 
nearly so. 11. Glossa approximately as long as 
prementum or longer. 13. Segment 1 of labial 
palp about twice as long as 2-4 together, or 
somewhat longer. 16. Maxillary blade about 
twice as long as prepalpal part of galea. 17. Galeal 
comb of about 14-17 bristles. 20. Labrum less 
than twice as broad as long; basal part glabrous; 
distal margin of basal area a weak transverse 
ridge; labral apex flattened (Fig. 27D). 25. Man- 
dible with well-developed, pointed, basal projec- 
tion on upper margin [mandible strongly curved 
in male of ^. nomadoides (Spinola)] (Fig. 27F). 27. 
Clypeus variable in width (in males more than 
three to five times broader than long, in females 
less than three to four times broader); distinctly 
protuberant (lateral view); distal margin with 
strong projection near lateral margin of labrum. 
30. Inner subantennal suture angulate or strongly 
curved. 31. Subantennal area slightly wider than 
inner suture length and than antennal socket. 33. 
Tentorial pit approximately at midpoint of outer 
subantennal suture. 34. Antennal socket on lower 
third of face or nearly so. 36. Antennal flagellum 
of male unmodified, much shorter than head; 
flagellomere 1 about as long as 2 or little longer, 
and slightly longer than broad. 37. Lower paraoc- 
ular area strongly swollen mesally. 38. Facial 
fovea narrow, well marked. 39. Orbits subparallel 
or slightly divergent below. 40. Middle ocellus 
below dorsal orbital tangent. 41. Vertex convex. 
42. Gena of male (lateral view) generally wider 
than eye (about as wide as eye in S. psamita Toro 
and Ruz), of uniform width or somewhat wider in 
the middle. 43. Pronotum with dorsal preapical 
ridge usually more prominent laterally than me- 
dially. 44. Mesepisternum with surface facing 
anteriorly extremely reduced and slightly convex. 
45. Pre-episternal groove weak, almost reaching 
Figure 27. Spinoliella nomadoides (Spinola). Male: 
A, B, Genitalia, dorsal, ventral and lateral views; 
C, T8; D, Labrum; E, Metasomal tip showing 
S6; F, Mandible. Female: G, Leg 3, outer view. 
gn = gonostylus. 
Panurginae: Calliopsini and Allies 
241 
scrobal level. 46. Pterostigma longer than (at least 
slightly) and somewhat less than twice as wide as 
prestigma; side basal to vein r subparallel to 
costa, that within marginal cell straight or nearly 
so. 47. Marginal cell rather wiciely and somewhat 
obliquely truncate at apex, variable in length. 49. 
Subniarginal cell 1 longer than cell 2 + 3. 51 . F"irst 
recurrent vein distant from first transverse 
cubital. 52. Forewing with cu-v longer than 
second abscissa M + Cu. 53. Hind wing with cu-v 
less than 1/2 as long as second abscissa M + Cu. 
54. Dorsal svnface of propodeum at least twice as 
long as metanotum. 55. Propodeal triangle ba- 
sally depressed medially, microareolate, with no 
striae, glabrous. 57. Basitarsus 1 about seven 
times longer than broad in male, five times longer 
in female; tarsomeres 2-4 unmodified. 59. Mid- 
dle tibial spur almost straight, finally serrate; 
usually about half as long as basitarsus 2 (some- 
times somewhat longer). 60. Basitarsus 2 usually 
as long as 1 (sometimes somewhat longer) and 
shorter than 3, tarsomeres 2-4 unmodified. 61. 
Tibia 3 of female (Figs. 27G, 28D) almost twice 
as long as basitarsus 3; keirotrichia on inner 
surface forming patches at both ends (sometimes 
with few hairs between patches); male tibia 3 with 
keirotrichia rather dense but absent medially. 62. 
Tibial scopa of medium-sized, simple, sparse 
hairs (especially on outer surface). 63. Inner hind 
tibial spur with fine and dense teeth, almost 
straight; outer with teeth sparser than on inner. 
64. Basitibial plate of male a swollen shiny area 
without defined margins; in female, rounded with 
margin a weak ridge. 65. Basitarsus 3 of female 
and tarsomeres unmodified. 66. Claws (in both 
sexes) bifurcate as in Arhysosage. 67. Metasoma in 
male usually slightly narrower than thorax. 69. 
T2-5 of male with gradulus posterolaterally 
short, not strongly carinate and with postgradular 
depression narrow (about 1/10 length of tergum) 
and shallow except somewhat deeper on T2; 
posterior marginal areas of Tl-5 in male and 
Tl-4 in female pilose; terga (both sexes) with no 
hair bands; lateral fovea of T2 (both sexes) 
slightly depressed, usually oval, somewhat vari- 
able in size. 70. Pygidial plate of male well 
developed with margins carinate. 71. T7 of fe- 
male trapezoidal (Fig. 28C) or squared. 72. T8 of 
male as in Figure 27C. 73. Metasomal sterna of 
male with hairs mostly very short, more abun- 
dant laterally, sparse or absent on midline, di- 
rected posteriorly (some di'^ected obliquely on 
S6). 73a. Metasomal sterna of female with hairs 
short, dense, directed caudad, not appressed, 
longer only on distal sternal margins. 74, 75. 
Similar to Arhysosage. 11 . S6 of male with distal 
margin produced medially (Fig. 27E). 78. S6 of 
female with proximal laminar lobes surpassing 
the apodemes; duplication thin (attached to hard- 
ened area of sternum distally); S6 with distal 
margin medially produced or concave; with pre- 
marginal fringe or patch of hairs at apex (Fig. 
28E). 79. S7 of male distally with two small 
projections; proximal arms forming a V (Fig. 
28B). 80. SB of male with body gradually sepa- 
rated from median distal projection; projection 
tapered toward apex and short-pilose (Fig. 28A). 
83. Gonocoxites rather elongate, though some- 
what wider in middle except S. psamita, rather 
squared (seen from ventral side); ventrally com- 
pletely fused on midline. 84. Gonostylus vestigial 
(about 1/8 length of gonocoxite in ventral view) 
and fused to gonocoxite (Fig. 27A, B). 85. Volsel- 
lae indistinct, apparently absent (perhaps repre- 
sented only by sclerotized area fused to 
gonocoxite and gonostylus). 86. Penis valve api- 
cally bilobed (simple in undescribed Argentine 
species not placed as to subgenus). 87. Penis 
completely membranous, somewhat shorter and 
wider than valve, and fused to valves basally. 88. 
Sting short (not reaching stylus apex) (Fig. 28F). 
Discussion. In the cladogram (Fig. 30) Spinoliella 
is the sister of Callonychium; it has few unique 
apomorphies. 
The subgenera Spinoliella s. str. and Peniella 
Toro and Ruz are not redescribed here. A key to 
these two subgenera, however, is given mainly to 
note some additional characters that separate 
them and to correct a mistake in the key by Toro 
and Ruz (1972). 
Key to the Subgenera of Spinoliella 
1. Males 2 
— Females 3 
2. Basitarsus 3 on outer surface with hairs 
short, dense. Facial fovea at least as 
Figure 28. Spinoliella nomadoides (Spinola). Male: 
A, B, S8 and 7, dorsal and ventral views. Female: 
C, T7; D, Tibia 3, inner view; E, S6, dorsal and 
ventral views; F, Sting. 
242 
The University of Kansas Science Bulletin 
broad as one-third minimum width of 
scape. S6 with distal projection gradually 
separated from rest of sternum and no- 
ticeably surpassing lateral areas 
Spinoliella s. str. 
— Basitarsus 3 on outer surface with hairs 
long, sparse. Facial fovea approximately 
as broad as one-fifth minimum width of 
scape. S6 with distal projection rather 
abruptly separated from rest of sternum 
and slightly surpassing lateral areas . . 
Peniella 
3. Outer hind tibial spur about half length 
of inner, distinctly curved at ape.x (Fig. 
27G). S6 distally much narrower than 
proximally; hairs at apex dense, forming 
a patch at each side of midline 
Spinoliella s. str. 
— Outer hind tibial spur about 2/3 length 
of inner, almost straight. S6 with distal 
area slightly narrower than proximally; 
hairs at apex ordered in 3 well-defined 
rounded rows Peniella 
Subgenus Spinoliella Ashmead s. str. 
Spinoliella Ashmead, 1899: 84. Type species: Camp- 
topoeum nomioides (sic) SpmoXa. — Camptopocum 
nomadoides Spinola, 1851 (monobasic and original 
designation. 
This subgenus is easily distinguished by the 
characters given in the above key and in Toro and 
Ruz (1972). 
Comments. Spinoliella s. str. contains two species: 
*S. (S.) nomadoides (Spinola) and * psamita Toro 
and Ruz. 
Distribution. Chile. 
Subgenus Peniella Toro and Ruz 
Peniella Toro and Ruz, 1972: 146. Type species: Camp- 
topoeum maculalum Spinola, 1851 (original designa- 
tion). 
This subgenus is distinguished by the charac- 
ters given in the above key and by Toro and Ruz 
(1972). 
Comments. Peniella includes four species: S. (P.) 
*maculata (Spinola), *herbsti (Friese), *rozeni Toro 
and Ruz and * rufiventris Toro and Ruz. 
Distribution. Chile. 
Genus Callonychium Brethes 
(Figs. 6D, 7B, 8D, 29, 30) 
Diagnosis. Close to Spinoliella, distinguished as 
follows: Gena with yellow. Antennal sockets 
(lower margins) usually at lower fourth of face. 
Lateral part of clypeus strongly bent posteriorly. 
Male. Basitibial plate flattened, delimited by car- 
Figure 29. Callonychium mandibulare Brethes. 
Male: A, B, Genitalia, dorsal, ventral and lateral 
views; C, S6, ventral view; D, E, S7 and 8, dorsal 
and ventral views; F, Labrum; G, T8; H, Mandi- 
ble. Female: I, S5, ventral view; J, Sting, lateral 
view; K, 86, dorsal and ventral views; L, T7; M, 
Sting, ventral view. 
ina. Metasomal apex strongly bent down and 
forward. Sterna with modifications (ridges, pro- 
jections, carinae). Female. S6 distally almost as 
broad as basally, distal margin slightly concave 
with curved distal band of curved and dense 
hairs. 
Description. 1. Length 3-7 mm. 3. Head and 
thorax with several to extensive yellow areas, 
more abundant in male than in female. Yellow 
area of lower part of face generally prolonged 
onto dorsal part along orbit, yellow also on gena. 
Thorax with yellow at least on pronotum, meta- 
notum and legs. 4. Metasoma with yellow marks 
laterally only or forming complete or fragmented 
bands. 5. Pubescence short and sparse on head 
and thorax, shorter and appressed on metasoma. 
6. Integument in general distinctly microareolate, 
especially on head and thorax, slightly so on 
metasoma. 7. Punctures mostly fine, sparse on 
head and thorax, smaller on metasoma e.xcept on 
sterna of male, dense on sterna of female. 8. Head 
somewhat broader than long and usually about as 
broad as thorax. 11. Glossa about as long as 
prementum or longer. 13. Segment 1 of labial 
palp almost three times longer than 2-4 together 
Panurcjinae: Calliopsini and Allies 
243 
or longer. 16. Ma.xillary l)lacic about twice as long 
as prepalpal part of galea. 17. CJaleal comb 
absent. 20. Labrum about as long as broati or 
broader; basal area glabrous; distal margin ot 
basal area in male usually appearing Battened or 
with weak ridge, of female with ridge stronger 
than in male; labral apex not inflexed in either 
sex. 25. Mandible of male with moderate to 
strong process on upper margin. 27. Clypeal 
width variable (almost three to five times broader 
than long), usually distinctly protuberant (about 
1/3 of eye width in lateral view), lateral part 
strongly bent posteriorly, distal margin usually 
with no projection near lateral margin of labrum. 
28. Epistomal suture, below intersection of outer 
subantennal suture, usually almost straight. 30. 
Inner subantennal suture angulate just below 
antennal socket (Fig. 6D). 31. Subantennal area 
wider than inner suture length and wider (or 
much wider) than socket. 33. Tentorial pit at 
midpoint of outer subantennal suture or nearly 
so. 34. Antennal sockets (lower margins) usually 
at lower fourth of face (sometimes at lower third). 
36. Antennal flagellum of male unmodified, 
much shorter than head; tiagellomere 1 as long as 
2 or longer, about as long as broad. 37. Lower 
mesal paraocular area often conspicuously pro- 
tuberant; lateral part of lower paraocular area 
strongly bent posteriorly. 38. Facial fovea well 
marked, narrow (almost linear). 39. Orbits gen- 
erally subparallel, sometimes slightly convergent 
below in male. 40. Ocelli variable in position 
(middle ocellus above or below dorsal orbital 
tangent). 41. Vertex convex. 42. Gena of male 
(lateral view) somewhat variable in width, of 
uniform width from above to below or somewhat 
wider medially. 43. Pronotum with dorsal pre- 
apical ridge prominent only laterally (weak or 
absent medially). 44. Mesepisternum with area 
facing anteriorly usually reduced and slightly 
convex. 45. Pre-episternal groove shallow, ap- 
pearing only as a line (as in Arhysosage) if extend- 
ing below scrobal level. 46. Pterostigma longer 
than and about twice as wide as prestigma; side 
basal to vein r subparallel to costa [except clearly 
divergent in C. mandibulare (Friese)], that within 
marginal cell straight. 47. Marginal cell obliquely 
and widely truncate at apex, shorter than distance 
from its apex to wing tip. 48. Submarginal cells 
two. 49. Submarginal cell 1 longer than cell 2 4-3. 
51. First recurrent vein entering 2nd submarginal 
cell but variable in position, not meeting first 
transverse cubital. 52. Vein cu-v shorter than 
second abscissa M -I- Cu. 53. Hind wing with vein 
cu-v 1/4 to 1/6 as long as second abscissa M -I- Cu. 
54. Dorsal surface of propodeum usually about 
twice as long as metanotum or longer. 55. Pro- 
podeal triangle basally with no special modifica- 
tions, glabrous. 57. Basitarsus 1 five to seven 
times longer than broad in male and four to seven 
in female; tarsomeres 2-4 unmodified. 59. Mid- 
dle tibial spur somewhat curved, finely toothed 
and about half as long as basitarsus 2 or nearly so. 
60. Basitarsus 2 longer than basitarsus 1 (at least 
slightly) and shorter than 3. 61. Tibia 3 of female 
about twice as long as basitarsus 3 or nearly so; 
inner surface with keirotrichia limited to patch at 
each end; male tibia 3 with keirotrichia forming 
narrow band of sparse hairs on inner surface 
between patches at ends. 62. Tibial scopa of 
medium or small, scattered and inconspicuously 
branched hairs on outer surface; longer and 
denser on dorsal margin. 63. Hind tibial spurs 
appearing untoothed but with minute teeth diffi- 
cult to see; outer shorter than inner, slightly 
curved or almost straight, inner straight. 64. 
Basitibial plate of male well defined, sometimes 
with smooth borders. 65. Tarsus 3 unmodified. 
66. Claws deeply cleft with rami subequal in 
male, simple in female. 67. Metasoma in male 
narrower than thorax (except about as broad as 
thorax in C. mandibulare Friese). 69. T2-5 of male 
with gradulus posterolaterally short (weak) or 
absent and postgradular depression narrow, shal- 
low, except deep on T2 of C. mandibulare; poste- 
rior marginal areas of T 1-5 in male and Tl-4 in 
female pilose; terga (both sexes) without hair 
bands; lateral fovea of T2 (both sexes) with shape, 
length and depth variable. 70. Pygidial plate of 
male well developed, rounded or bifurcate at 
apex. 71. T7 of female rather squared, with 
dorsal and proximal area not greatly expanded 
(Fig. 29L). 72. T8 of male somewhat variable but 
usually similar to an elongated trapezoid (Fig. 
290). 73. SI -5 of male with hairs mostly directed 
posteriorly, variable in length and density, mostly 
short, somewhat oblique on S6; sternal surface 
with modifications such as protuberances, projec- 
tions, ridges or carinate areas; metasoma strongly 
curved down and forward at apex. 73a. SI -5 of 
female with hairs mostly short, dense, appressed. 
74. S4 of male with distal margin almost straight, 
somewhat concave or emarginate (except widely 
convex in C. mandibulare). 75. S5 of male with 
distal margin almost straight medially (except 
widely convex in C. mandibulare). 76. S5 of female 
with long and recurved gradulus and no median 
sclerotized area between gradulus and proximal 
margin (Fig. 291). 77. S6 of male distally with 
emargination (deep or shallow) between lateral 
and median areas (Fig. 29C). 78. S6 of female 
with proximal laminar lobes widely separated 
from lateral apodemes; basal sclerotization ab- 
sent; lateral margin with strong, somewhat 
curved ridge; duplication heavily sclerotized; dis- 
tal margin concave medially with dense curved 
hairs (Fig. 29K). 79. S7 of male transverse, bar- 
like, with a short laterodistal projection (Fig. 
29D). 80. S8 of male with elongate, slender, 
median projection abruptly separated from a 
proximally emarginate rectangular body (Fig. 
29E). 83. Gonocoxites short, rather square, ven- 
tral and dorsal sides completely fused. 84. Gono- 
stylus rudimentary, fused to gonocoxite 
(recognizable only by the few, small hairs). 85. 
Volsellae indistinct, apparently absent (or repre- 
sented only by a sclerotized transverse area com- 
pletely fused to each other and to gonocoxite). 86. 
244 
The University of Kansas Science Bulletin 
Penis valves usually complex; dorsally, not at- 
tached to each other by bridge but attached 
basally to penis by membrane. 87. Penis mem- 
branous (variable in shape), shorter and wider 
than valve, with special internal sclerotization. 
88. Sting rudimentary (reduced to a small sclero- 
tized area), triangular plate modihed, elongate 
(Fig. 29J, M). 
Comments. The subgenera Callonychium Brethes 
and Paranychium Toro recognized by Toro (1989) 
have not been redescribed in this study, although 
a key for subgenera by Toro and Herrera (1980) 
is here provided. The type species of this genus, 
C. argentinum, was not available for study. 
Discussion. Callonychium is clearly differentiated 
by its apomorphies and is more closely related to 
Spinoliella than to Arhysosage. The relationship 
among these three genera has been also noted by 
Toro and Herrera (1980). 
Rozen (personal communication) has studied 
the larvae oi Spinoliella herbsti and a new species of 
Callonychium. His preliminary observations indi- 
cate that both species present "peculiar thoracic 
tubercles ..." that "may represent a synapo- 
morphy. "" He has also found that the low anten- 
nal papillae of most panurgines are somewhat or 
strongly projecting in these species; they are 
presumably synapomorphic. On the other hand 
he stated that "the recessed labial maxillary 
region of Callonychium and the projecting one of 
Spinoliella seem incongruous with the idea of the 
close relationship." 
In my opinion the first two characters indicated 
by Rozen are likely to be good synapomorphies, 
which would reinforce the close cladistic rela- 
tionship indicated by adult characters between 
Spinoliella and Callonychium. The third character, 
however, may not be useful to establish rela- 
tionships. 
The similarity between the flabellum of Callo- 
nychium and that of Perdita that Michener and 
Brooks (1984) observed does not lead to the 
conclusion that these two genera are closely re- 
lated, and as those authors concluded, must result 
from convergence. 
Key to the Subgenera of C.m.lonychium 
1 . Axilla yellow, acute. First metasomal 
sternum of male without premarginal 
process Callonychium s. str. 
— Axilla black, depressed. First metasomal 
sternum of male with premarginal proc- 
ess Paranychium 
Subgenus Callonychium Brethes s. str. 
Callonychium Brethes, 1922: 120. Type species: Callo- 
nychium art^enlmum Brethes, 1922 (monobasic). 
This subgenus is distinguished by the charac- 
ters given in Toro and Herrera (1980). 
Comments. The subgenus Callonychium s. str. 
contains six species: C. argentinum Brethes, 
brasiliense (Ducke), *jiaviventre (Friese), luteimacula- 
turn (Strand), *mandibulare (Friese) and petuniae 
Cure and Wittmann and others (undescribed) 
from eastern South America. 
Distribution. Argentina, Paraguay, Brazil. 
Subgenus Paranychium Tore 
Paranychium Toro and Herrera, 1980: 213 (not valid 
because no type species was designated). 
Paranychium Toro, 1989: 231. Type species: Camptopoeum 
chilense Friese, 1906 (original designation). 
This subgenus is distinguished by the charac- 
ters given by Toro and Herrera (1980) and by 
Toro (1989). 
Comments. The subgenus Paranychium contains 
five species: C. (P.) *chilense (Friese), *aricense 
Toro and Herrera, * atacamense Toro and Herrera, 
* coquimbense Toro and Herrera and minutum 
(Friese) as well as undescribed species from Ar- 
gentina. 
Distribution. Argentina, Chile. 
CLADISTIC ANALYSIS 
This is part of a larger study of the subfamily 
Panurginae. A preliminary hypothesis of rela- 
tionships among genera of Panurginae was used 
to identify the close relatives of the three tribes 
Perditini, Protomeliturgini, and Calliopsini 
(Ruz, 1986). The tribe Meliturgini was regarded 
as the sister to the three tribes mentioned above, 
and these four were in turn related to the tribe 
Panurgini. This outgroup information was used 
to construct a hypothetical ancestor using the 
Maddison, Donoghue and Maddison algorithm 
(1984). This ancestor is characterized in the 
matrix (Table 2) and its characters were coded 
according to codes for the entire subfamily. Thus 
"1" may be apomorphic at the subfamily level, 
but plesiomorphic at the present level of analysis. 
This was done so that the several studies I plan 
(including Ruz, 1986) can be examined together. 
When a character is variable within a taxon, 
the code for the plesiomorphic state (0) was used 
in the analysis. An exception was character 72 
(Fig. 30) for which the derived state in C. (Micro- 
nomadopsis) was used to demonstrate an apparent 
convergence with C. (Liopoeodes). Omission of 
character 72 did not change the topology of the 
cladogram. 
Panurginae: Calliopsini and Allies 245 
lablc 1. List ol cliaracters for genera and subgenera ol panurgine tribes Calliopsini, Protomeliturgini 
and Perditini. (For the Panurginae as a whole, (0) is considered plesiomorphic but for the groups 
treated here it is in a few cases apomorphic.) 
3. Metasoma with yellow markings (1). Ahhough the presence of yellow on the mctasoma is widespread 
among bees, including some genera of Panurginae, it is lacking almost completely in the outgroups, 
which indicates that its presence should be considered as apomorphic. Lack of yellow was coded (0). 
5. Metasomal terga with distal hair bands (1). Several groups of bees exhibit this feature, which 
apparently has arisen independently in different lineages. Among the Panurginae, however, it is 
present onh' in Protomeliturga, Acamptupoeum, Calliopsis s. str. , C (Perissander), C. (Calltopsirna), and C. 
(Verbenapis). Since this character is also present in various Andreninae and colletid bees (both 
outgroups) it is possible that the lack of these hair bands (coded 0) is not plesiomorphic but is 
apomorphic. The genera of Panurginae mentioned above, however, because of other sets of characters, 
are considered among the most derived genera. Therefore the presence of these hair bands is likely to 
be apomorphic. 
6. Glossa more or less as long as prementum (1). This condition was considered as apomorphic for the 
subfamily, since a shorter glossa (0) is present in most genera of the Andreninae and Colletinae. This 
study, however, shows that this condition (1) may be present in the ancestor of Calliopsini and is 
widespread among the Calliopsini and other Panurgini; it therefore is considered as a plesiomorphy for 
Calliopsini. 
Glossa shorter than prementum (coded as in Table 2) is an apomorphy for the present analysis and 
appears as a reversal in the cladogram (Fig. 30) for C. (Perissander) and Perdita (although the glossa is 
long in some groups of Perdita). 
8. Paraglossa shorter than suspensoriurn (1). An autapomorphy for Arhysosage, a highly derived genus. 
Although the paraglossa is short in Andreninae (outgroup), this feature seems to be autapomorphic for 
Arhysosage, a genus with many derived characters. In other Panurginae the paraglossa is longer than or 
about as long as the suspensorium (0), and for this subfamily this condition is presumably 
plesiomorphic. Both shortness of the paraglossa in Arhysosage and its strong elongation among the long- 
tongued bees seem to be apomorphic. 
9. Segment 2 oj labial palpus about three times longer than 3(1). Autapomorphic, unique for Protomeliturga. 
Segment 2 of the labial palpus in other Panurginae and in the outgroups is usually about as long as 3, 
or if longer, less than two times longer than segment 3 (0). 
1 1. Segments 3-4 of labial palpus articulating preapically and projecting at about 90° to the axis of 1-2 {\). An 
autapomorphy for Protomeliturga, unique among the short-tongued bees, in almost all of which the 
articulation of the labial palpal segments 2-4 is apical (0). The preapical articulation between labial 
palpal segments 2 and 3 is present in long-tongued bees and is certainly derived. 
12. Mentum about 1/5 length of prementum (1). A few Panurginae present the apomorphic condition: 
Callonychium and two Old World genera, Camptopoeum and Melitturga. Short-tongued bees usually have a 
mentum shorter to much shorter than 1/5 the length of the prementum, a condition which is 
plesiomorphic and coded (0). 
13. Galeal comb absent (1). Many Panurginae and the outgroups have a well-developed galeal comb 
(0). This comb is absent in Arhysosage, Callonychium (also in some Perdita) and in several Old World 
panurgines. The evolutionary trend goes from comb well developed, reduced, to absent; the latter 
condition is clearly derived. 
15. Labrum of male with weak transverse ridge (1). Labrum of male flat (2). One or the other of the derived 
states of this character appears occasionally among Panurginae. A weak ridge is present in Spinoliella 
and Callonychium, while a flat labrum characterizes some Old World and North American genera, and 
Arhysosage. A labral ridge (sometimes strong) or carina (delimiting the basal area) (0) is widespread 
among bees including the outgroups except in Megandrena (Andreninae) in which the labrum is flat, 
without a ridge. 
16. Labrum of male with basal area pilose (1). Generafly when a labral ridge or carina exists, hairs 
appear lateral and distal to the ridge or carina. This is also the case among many Panurginae and in the 
outgroups. The genera that present the derived feature are indicated in the cladogram, except for 
Perdita in which only some species have developed the derived condition. Lack of such hairs is coded 
17. Labrum of female with apex not inflexed (1). Among females of Panurginae and the outgroups, this 
feature is present only in Arhysosage, Spinoliella, and Callonychium. Otherwise the female labrum usually 
246 The University of Kansas Science Bulletin 
shows the plesiomorphic condition, that is, with the apex at least sHghtly inflexed (0). 
18. Mandible of male simple (1). This is an apomorphic character for the Panurginae since nearly all 
colletids and most Andreninae have bidentate mandibles. Among the Calliopsini, a preapical tooth is 
present only in Arhysosage. This condition (coded as in Table 2) also appears in many Perditini and 
should be considered as apomorphic for the tribes here considered. The cladogram shows this as a 
reversal in Perdita and Arhysosage. 
22. Tentorial pit of male at intersection of outer subantennal and epistomal sutures (1 ). Tentorial pit of male in outer 
subantennal suture (close to epistomal suture) (2). Tentorial pit oj male near middle of outer subantennal suture (3). In 
colletids (second outgroup for Panurginae), even though they possess only the inner subantennal 
suture, the tentorial pit is in the epistomal suture sometimes considerably below the subantennal suture 
(0). In Andreninae (first outgroup for Panurginae) this pit is sometimes in the intersection between the 
epistomal and the outer subantennal suture, as in (1). The migration of the tentorial pit into the outer 
subantennal suture is unique to Panurginae and obviously apomorphic. Condition (1) should be 
considered plesiomorphic in this study, while (2) and (3) are sequential apomorphic states and uniquely 
derived characters in the Calliopsini. 
23. Antennal sockets of male (lower margins) on lower 1/3 or 1/4 of face (1). In most Panurginae and 
outgroups the antennal sockets are placed more or less in the middle of the face (0). 
24. Antennal scape of male robust (1). Apomorphy for some Old World genera and Calliopsis 
(Ceroliopoeum). A derived feature among Panurginae, absent in the outgroups. A slender scape is coded 
(0). 
28. Lower paraocular area strongly swollen mesally (1). This character is rarely present among bees. The 
only Panurginae with this feature are Spinoliella and Callonychium, although it is approached in some 
Calliopsis and Perdita. The rest of the genera have the lower paraocular area flattened or slightly convex 
(0). In the outgroups this area is usually completely flat. 
37. Pronotal carina or lamella of male present (1). This structure is present only in three genera of 
Panurginae, one of which is Protomeliturga, although it seems that it appears independently in several 
other groups of bees. Many bees, including the panurgine outgroups, show the plesiomorphic 
condition, i.e., the lack of a carina or lamella on the dorsal part of pronotum (0); instead, at least 
laterally, a rounded ridge is developed. 
39. Pre-epistcrnal groove short, not extending below scrobal level (1 ). A short pre-episternal groove is present 
in all the tribe Calliopsini, in some other panurgine genera and some Andreninae. The Colletinae 
(second outgroup), however, show a very long groove that always extends below the scrobe (0). This 
fact indicates that a long groove is plesiomorphic and a short one apomorphic. 
40. Pre-episternal groove curved, meeting scrobe (1). Some Andreninae and Perdita and Acamptopoeum 
among the Panurginae present this feature, which is clearly apomorphic. In Colletinae the pre- 
episternal groove is straight and does not meet the scrobe (0), which indicates that this is the primitive 
condition. In Andrenidae as a whole the scrobal suture is absent in front of the scrobe. Therefore it is 
unlikely that the lower part of the curved groove noted above is derived from the scrobal suture, 
although this is the usual interpretation. 
41. Axilla of male with patch of velvety hairs (1). The presence of this feature is a unique apomorphy for 
Calliopsis s. str. ; it is unknown in other bees. Lack of such hairs is coded (0). 
42. Metanotum with lateral patch of short velvety hairs (1). This character, known only in Calliopsis s. str., 
C. (Perissander) and C. (Calliopsima), must be an apomorphy. Hairs in this area usually are not different 
than on the rest of the thoracic dorsum (0). In Calliopsis s. str. the patch is conspicuous and brown in 
males, small and white in females; the latter condition is also present in both sexes of C. (Perissander) 
and C (Calliopsima). 
44. Pterostigma with sides parallel or subparallel (1). Among Panurginae this character appears in the 
whole tribe Calliopsini and independently in a few other genera. Although this character is also present 
in some genera of the outgroups, it should be considered an apomorphic, since in Panurginae, the 
Calliopsini are the most derived bees. Pterostigma with margin basal to vein r divergent from costa so 
that the stigma is relatively broad is the plesiomorphy (0). 
45. Pterostigma with margin within marginal cell straight or nearly so (1). All the Calliopsini present this 
apomorphy, although it has also appeared independently in two unrelated genera. Most genera of the 
outgroups show the plesiomorphy, that is, pterostigma with margin within marginal cell clearly convex 
(0). 
46. Marginal cell shorter than distance between its apex and wing tip (1 ). Marginal cell much shorter than distance 
between its apex and wing tip (2). Most Panurginae and also the outgroups have a marginal cell about as 
Pantrginae: Calliopsini and Allies 247 
long as or usually longer than the distance between its apex and the wing tip (0); this is the 
plesiomorphic condition. 
49. Basal vein (fore wing) strongly curved toward wing base (1). A unique character for Protomeliturga; 
among Panurginae and the outgroups this vein is usually straight or slightly curved (0). 
52. Propodeal triangle smooth (1). Only a few genera of Panurginae (all of them Calliopsini) possess this 
derived character. These are: Acamptopoeum, C. (Verbenapis), C. (Hypomacrotera) and some C. (Microno- 
madopsis). Usually the propodeal triangle is striate or rugose, or at least minutely areolate (0). This 
plesiomorphy occurs in Andreninae and in many Colletinae. 
55. Tarsus 1 of female on inner surface mainly with hairs rigid, curved at apices (Fig. 14H) (1). A strong, 
unique character oi Calliopsis (Verbenapis). Hairs on this part of the tarsus 1 are rigid but straight (0) in 
other genera and in the outgroups. 
56. Femur 2 oj female with well-defined basal comb on ventral margin (Fig. lOD) (1 ). A uniform comb clearly 
differentiated from the hairs of the surrounding parts of the femur is present in all Calliopsini and, 
although appearing independently, in a few other genera of Panurginae. The comb of femur 2 of the 
female in the outgroups consists of stiff, short, dense, but irregularly organized hairs not very different 
from those of the surrounding areas (0); this must be the primitive condition. 
58. Basitarsus 2 of male longer than 5(1). This feature is present only in the Calliopsini [Acamptopoeum, 
Calliopsis s. str. , C. (Perissander), C. (Liopoeodes), C. (Ceroliopoeum), and several C. (Micronomadopsis) and 
in the Old World genus Plesiopanurgus]. In the rest of the Panurginae and in the outgroups basitarsus 2 
of the male is equal to or shorter than 3 (0). 
60. Tarsus 2 of male with tarsomeres 2-4 widened distally (1). This feature is found in a few genera of 
Panurginae and in Euherbstia and Orphana (slightly so in the latter) among Andreninae. Rather narrow 
tarsomeres (0) are plesiomorphic. 
62 . Tibia 3 of female on inner surface with keirotrichia forming a longitudinal dorsal or medial strip ( 1 ) . Tibia 3 of 
female with keirotrichia only at both ends of inner surface (Fig. 28D) (2). This sequence seems to represent an 
evolutionary trend. The lack of keirotrichia should be the most derived condition. Presence of 
keirotrichia on most of the inner surface of tibia 3 (0) as in the outgroups is the primitive condition. 
63. Tibia 3 of male with keirotrichia forming a longitudinal strip (1). The keirotrichia in many bees are 
present on most of the inner surface of tibia 3 (0) and (1) should be considered an apomorphy. 
Reduction to a strip of keirotrichia occurs only in Spinolella, Callonychium, and one Old World genus. 
64. Tibia 3 of male with dorsal margin evenly carinate (Fig. 4F) or the carina fragmented, forming a series of teeth 
(1). This character appears in several members of the Panurginae but not in its outgroups. For this 
reason the presence of a tibial carina or of teeth may be apomorphic for the subfamily while its absence 
(0) is plesiomorphic. In this study, however, the ancestor probably had this feature; therefore the lack of 
teeth on the tibia 3 of the male in all Calliopsini should be interpreted as apomorphy. State (1) is 
present in Protomeliturga and exceptionally in Perdita. 
67 . Tibial scopa of moderately abundant hairs ( 1 ). Tibial scopa of extremely sparse hairs (2). The scopa is dense 
(0) in most Andreninae and Colletinae and for that reason at the subfamily level this feature is a 
plesiomorphy. Among the Calliopsini, however, condition (1) is the most widespread and may have 
been present in their ancestor; in this analysis, therefore, it is considered a plesiomorphy for 
Calliopsini. In this tribe only Spinoliella and Callonychium have the tibial scopa of sparse hairs (2), a 
derived state. A dense scopa (coded as in Table 2) appears in the cladogram as a reversal for Perdita, 
Calliopsima, and Verbenapis and should be considered as apomorphic for the tribes here studied. 
69. Basitibial plate of male represented only by a swollen area, laterally not delimited by a ridge or carina ( 1 ). This 
feature is known only in Spinoliella and some Perdita; in other panurgines and in most members of the 
outgroups the plate is delimited (0). 
70. Apex of basitarsus 3 of female with upper distal projection reduced so that apex is oblique ( 1 ). Apex of basitarsus 
of female transverse (2). A well-developed upper distal projection on basitarsus 3 (0) is widespread among 
bees. Both well-developed and reduced projections are also present in the outgroups of Panurginae. 
Thus it is difficult to determine the polarity. However, a truncated basitarsus occurs only in some of the 
otherwise most derived Panurginae, such as Spinoliella, Arhysosage, Callonychium and some Perdita. 
Therefore for Panurginae the direction of evolution is indicated. 
70a. Basitarsus 3 of male expanded laterally at apex on ventral margin (1), an autapomorphy for C. 
(Liopoeodes) (Fig. 22K). Lack of such expansion was coded (0). 
71. Tarsus 3 of male with tarsomeres 2-4 widened (1). This character has appeared in three different 
lineages among the Panurginae and is obviously apomorphic. Unmodified tarsomeres (0) are 
commonly found in many groups of bees including most genera of the outgroups. 
248 The University of Kansas Science Bulletin 
72. Tarsus 3 of male with tarsomeres 2-4 asymmetrical (1). Tarsomeres are usually symmetrical (0), a 
condition widespread in bees and wasps. Among panurgines, asymmetry, a unique and strong 
character, has been found only in some Calliopsis (Micronomadopsis), in C. (Liopoeodes) and slightly so in 
some C. (Liopoeum). 
72a. Distitarsus 3 of male widened medially, (1), a unique character known only for C. (Liopoeodes). In 
others the distitarsus is widest apically (0). 
73. Claws of female simple (1). Among Panurginae this character is present only in a few Old World 
genera and in Callonychium. Bifurcate claws (0) are widespread among bees and are obviously 
plesiomorphic. 
76. Tl-5 of male with posterior marginal areas (at least partially) pilose (1). This feature appears several 
times among Panurginae and other bees, and may be apomorphic. Since the metasomal terga in the 
outgroups present either condition, it is difficult to determine the direction of evolution of this 
character. However, the majority of the presumably more primitive Panurginae have terga with bare 
posterior marginal areas (0), a condition that may well be plesiomorphic. All genera with metasomal 
hair bands (character 5) have densely pilose posterior marginal areas. 
79. Pygidial plate of male well defined (1). A pygidial plate clearly delimited by a ridge or carina is 
present in most Calliopsini and independently also in a few Old World Panurginae, all of them quite 
derived genera as shown by other characters. This structure in males is widespread among bees and 
wasps and is plesiomorphic for the whole group. In Panurginae, however, it seems to be apomorphic, 
the plate having been largely lost in ancestors of Panurginae. In most genera of the outgroups the plate 
is represented only by a bare, shiny, triangular area or by an expansion of tergum 7 (0) similar to that 
of the the most primitive Panurginae. This last condition is considered plesiomorphic for Panurginae. 
Reacquisition of the defined plate should be no great evolutionary problem since the plate is defined in 
females; appropriate genes only need to be turned on in males. 
80. T8 of male at apex rather densely pilose (1). Usually T8 has sparse hairs or is glabrous (0). However, 
in some Panurginae, like Perdita, T8 is clearly pilose, a condition not found in the outgroups. 
81. S4 of male with distal margin slightly produced medially (1), as in C. (Macronomadopsis) and C. 
(Verbenapis), although barely distinct in the latter. S4 of male with distal margin clearly produced medially 
(2). This character is evidently apomorphic, for it is known only in Calliopsis and the Old World genus 
Plesiopanurgus, in which it must have arisen independently. S4 in other groups, including the outgroups, 
usually has the distal margin concave or almost straight medially (0). 
84. S5 of male with distal margin slightly produced medially (1). S5 of male on distal margin with a rather small 
median projection (2). S5 of male distally with an elongate median projection (3). This feature is present only in 
the genus Calliopsis. The margin of S5 is concave or almost straight medially (0) in most bees and 
wasps; this condition is plesiomorphic. 
85. SJ-5 of female with hairs mostly appressed (\). Several Calliopsini and also Protomeliturga and and one 
Old World genus exhibit this character. Although this feature arose independently several times in 
bees, including the outgroups, it is likely to be an apomorphy in Panurginae since it is present only in 
otherwise derived genera. Sterna with hairs mostly not appressed (0) seems to be the plesiomorphy. 
87. S5 of female with distal margin convex medially (1). This is a unique and strong synapomorphy of 
Calliopsini. The rest of Panurginae and outgroups present the plesiomorphic condition, that is, S5 
with distal margin concave or almost straight medially (0). This condition is also widespread in other 
bees and wasps. 
88. S5 of female with median sclerotized area between gradulus and proximal margin of sternum (Fig. 12A) (1). 
This derived character is present in Calliopsini and it has also appeared independently in one Old 
World genus. (In Callonychium it is absent, presumably secondarily lost.) The lack of this sclerotization 
(0) is widespread in bees. 
89. S6 of male with distal, elongate, tapered projection on each side of deep emargination (Fig. 1 IH) (1). This 
structure has arisen only in some Calliopsini. The lack of such projections on S6 (0) is widespread in 
bees and plesiomorphic. 
90. S6 of male about three or more times broader than long (1). This strong, derived character is present only 
in Perdita. Usually S6 of the male is about as wide as or somewhat wider than long (0). The outgroups 
of Panurginae and many other bees possess this plesiomorphic condition. 
93. S6 of female with two proximal laminar lobes (Fig. 17H) (1). This character is found only in 
Calliopsini. Without doubt an excellent apomorphy, this structure is unknown in other bees or wasps 
which have the proximal margin of S6 in females straight (0). 
94. S6 of female without basal sclerotization (1). This is an autapomorphic character oi Callonychium. The 
Panlr(,inae: Cali.iopsini and Ai.i.iks 249 
presence of basal sclerotization (U) is cliaractcristic oi the rest of Panurginae and outgroups. 
95. S6 of female with basal spine-shaped longitudinal sclerotization (Fig. ID) (1). This feature is present in 
most Calliopsini except Callonychium. It has also arisen, no doubt independently, in a few other genera 
of the subfamily. .\ basal sclerotization of S6 (females) is not spine-shaped (0) in panurgine outgroups; 
therefore the acquisition of this shape is clearly deri\e(l. 
96. S6 of female with basal sclerotization free at least at apex (1). A basal sclerotization on S6 is usually 
fused to the sternum (0). Therefore, when the sclerotization has a free apex, it seems clear that this 
condition is apomorphic. 
97. S6 of female with strong, almost straight ridge on lateral margin (Fig. ID) (1). The Calliopsini (except 
Arhysosage), Perdita. and some Old World genera possess this character, which is lacking in the 
outgroups. A lateral ridge on S6, if present, is usually curved (0), a condition which is here considered 
plesiomorphic. 
99. S6 of female with premargmat hairs in well-organized rows forming a continuous or medially interrupted patch 
or band (Figs. lOH, 12C) (1). This unique, strong character appears only in all Calliopsini. Premarginal 
hairs on S6 are usually organized in a dense patch (0). This primitive condition is present in 
Andreninae and Coiletinae. 
102. S 7 of male reduced to a transverse bar with two small latero-distal projections (Fig. 29D) (1). This unique 
apomorphy is known only in Callonychium. Usually S7 is more conspicuous and of a different shape, 
commonly v-shaped (0). 
103. S7 of male with small brush of hairs near middle of each lateral projection (Fig. 15D) (1). This sternal 
brush is only found in Calliopsis (Liopoeum); its absence is coded (0). 
105. S8 of male with a small lateral projection between body and distal projection (Fig. 1 IF) (1). This strong 
character appears only in Calliopsis except for C. (Verbenapis) in which it exists only as two weak lateral 
bumps. It is absent (0) in other bees. 
108. S8 of male robust, strongly concave laterally (Fig. 4G, H) (1). This is a unique, strong feature, 
present only in Protomeliturga. A thin structure with unmodified lateral margins is the plesiomorphy (0). 
111. Gonocoxites {ventral view) connected by membrane (Fig. 1 8A) ( 1 ). Gonocoxites partially or fully fused to each 
other (Fig. 25A) (2). The gonocoxites are completely separated (except at the very bases) (0) in the 
presumably most primitive Panurginae and in the outgroups as well. Therefore this feature must be a 
plesiomorphy. The evolution of this character seems to be toward a full fusion of the gonocoxites. 
113. Gonocoxite (dorsal view) proximally with well-developed, mesal, usually lobe-like process (Fig. 15A) (1). 
This strong character is present in Calliopsis (Liopoeum) and in one species of C. (Hypomacrotera). This 
structure is always lacking (0) in the outgroups. 
115. Gonostylus about 1/2 to 1/3 length of gonocoxite (ventral view) (1). Gonostylus vestigial, about 1/8 length of 
gonocoxite (2). Gonostylus rudimentary (recognizable only by the presence of few, short hairs at apex of gonocoxite) (3). 
Gonostylus absent (4). Reduction in gonostylar length occurs in a few Old World Panurginae as well as in 
Protomeliturga and Calliopsini (almost lost in Callonychium). This strong character is rather rare in other 
groups of bees and lacking in the outgroups. Gonostylus longer than gonocoxite (0) is the ancestral 
condition. 
1 18. Volsella rudimentary or absent (?) (1). Most Panurginae, all Andreninae and Coiletinae have well- 
developed volsellae (0). Arhysosage, Spinoliella, and Callonychium are the only Panurginae that have 
apparently lost them. 
119. Volsellae connected to one another by basal, narrow and weak, membranous bridge (\) . Volsellae connected to 
one another by widened membranous area or by slightly sclerotized cuticle (2). Volsellae partially fused to one another 
(3). Volsellae fully fused to one another but line of fusion visible (4). Volsellae absent or completely fused forming a 
highly sclerotized plate, coded as missing (9) for Arhysosage, Spinoliella and Callonychium. In the outgroups 
and many other bees the volsellae are free from one another (0). They may be attached to the 
gonocoxites or penis valves but not to each other. Volsellae mesally free is certainly a plesiomorphy. 
120. Volsellae without denticles (1). Denticles are usually present (0) on the volsellae. In Protomeliturga 
and some Calliopsini, however, the teeth are lost. 
122. Penis valve (lateral view) with longitudinal row of hairs (1). This autapomorphy for Acamptopoeum is 
absent in the outgroups, which lack such hairs (0). 
123. Penis valve complex (1). The presence of complex folds in the penis valve is found only in some 
subgenera of Calliopsis. Most primitive Panurginae and the outgroups possess simple penis valve (0). 
125. Penis completely separated from penis valve (1). Most Panurginae and outgroups possess a penis 
partially or fully fused to the penis valve (0). In Acamptopoeum and Calliopsis, however, these structures 
are completely separated. 
250 
The University- of Kansas Science Bulletin 
126. Penis with internal sclerotizatwn (1). An internal sclerotization ot the penis is a rare feature in other 
bees and is totally absent (0) in the outgroups. It is found only in Spinoliella and Callonychium. 
127. Sting short to very short, not reaching (or at least not surpassing) stylus (1). Sting rudimentary (2). Females 
of many bees and wasps have a long sting which is commonly exposed. This is true in colletids and 
some Andreninae. In Panurginae, only the most primitive ones have an elongate sting (0), which may 
be seen from outside but less frequently than that of CoUetinae. A short sting is present in most 
Calliopsini (except Acamptopoeum), Protorneliturga, Perdita, and in a few Old World genera. It is 
rudimentary, no doubt the most derived condition, in Callonychium. 
128. Sting truncate at apex (1). This strong feature is present only in Acamptopoeum and some Perdita 
(e.g., P. halictoides). Commonly the sting is acute at the apex (0). An apically truncate sting must mean 
that, even if it is elongate, it cannot be used for introducing venom. 
129. First valvifer elongate (1). The first valvifer (triangular plate) is usually more or less triangular (0). 
In Perdita and independently in Callonychium, however, the first valvifer is elongate, a modification 
probably related to reduction of the sting. This clearly derived condition is not found in CoUetinae and 
is rare in Andreninae. 
130. Valve of first valvula rudimentary (Fig. 24J) ( 1 ). Valve absent (Fig. 16B) (2). A well-developed valve of 
the first valvula (0) is present when there is a long sting, as in CoUetinae and some Andreninae. The 
tendency in evolution is toward the loss of the valve, which is correlated with the weak sclerotization of 
the first valvula. In Protomeliturga, Perdita, a few Old World genera, and most Calliopsini there is no 
valve. A tiny projection that may be a valve rudiment is present in C. (Ceroliopoeum) and some Old 
World genera. In this study the absence of a valve in the three tribes involved should be interpreted as a 
plesiomorphic character and the presence of a valve-like structure in Ceroliopoeum, as an autapomorphy. 
Seventeen taxa (15 taxa used by Ruz, 1986, 
and 2 additional taxa, Liopoeodes and Ceroliopoeum) 
and a total of 52 characters have been used in this 
study (Tables 1 and 2). Of the 130 derived 
characters previously considered (Ruz, 1986) for 
the whole subfamily, 53 were eliminated for being 
invariant among, or not applying to, the taxa 
here involved. Besides, 17 autapomorphies plus 
characters 70a and 72a added later, five con- 
vergences (see Table 1), and three characters with 
consistency indices (C.I.) 0.250 or below, were 
not used in this cladistic analysis. The three 
characters eliminated were the following: 26, 
Flagellomere 1 of male about twice as long as 2 or 
less. Even though this feature is diagnostic for 
some genera, it appears independently several 
times among Panurginae. 100, S6 of female with 
curved band of hairs on premarginal area. This 
character is known only in Calliopsini and seems 
to be derived, but it appears to arise independ- 
ently and to reverse several times within the tribe. 
116, Gonostylus partially or fully fused to gono- 
coxite. The gonostylus is lacking in several genera 
and subgenera of Calliopsini. Therefore this 
character was not informative in this study. 
The presence of hair bands on the metasomal 
terga (character 5) was only considered as a 
character for males. In several taxa among the 
Calliopsini these hair bands are present in both 
sexes. In Calliopsis (Liopoeodes), however, they are 
present only in females. The C.I. of this character 
of females was too low to be considered in the 
analysis. 
The polarity of variables (sequence of character 
states within each variable) has been determined 
using outgroup comparison. The outgroups con- 
sidered for the entire subfamily are: Andreninae 
(the other subfamily of Andrenidae) and Col- 
letinae (the subfamily considered the most primi- 
tive among Colletidae). 
The data (Table 2) were analyzed using a 
computer program (Swofford, 1985, PAUP ver- 
sion 2.4) based on parsimony. The options se- 
lected were the following: 1. MULPARS. 2. 
Rooting. 3. Global branch-swapping. 4. Delayed 
transformation optimization. Only three most 
parsimonious trees were found, which showed 
almost no topological diflferences. Therefore only 
one tree (Fig. 30) is illustrated. Statistics of the 
tree: length=115; C.I. = 0.6. 
The cladistic analysis of the data in Table 2 
indicated that Calliopsini is a strong mono- 
phyletic group, some of its synapomorphies being 
unique among all bees. It also shows that this 
tribe is subdivided into two clearly dilferentiated 
lineages, and that the sister group of 
Acamptopoeum is also well supported. 
The relationships among the nine taxa at the 
top of the tree, however, do not seem to be very 
stable due to homoplasies. Moreover, some taxa 
or groups of taxa are differentiated by reversals 
only. Calliopsis s. str. and Perissander, the sister 
group of the remaining seven taxa, may appear in 
a trichotomy depending on the distribution of 
character 42. This character may arise earlier in 
the cladogram (as a synapomorphy for all the 
nine taxa plus Calliopsima) and be reversed later 
Panurginae: Calliopsini and Allies 
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252 
The University of Kansas Science Bulletin 
Figure 30. Cladogram showing the phylogenetie relationships of the genera and subgenera of the 
tribes Perditini, ProtomeHturgini and Calliopsini. Autapomorphies and convergences (see Table 1) are 
also included here but only data in Table 2 were used in cladogram construction. 
Pani'rginak: Calliopsini and Allies 
253 
tor tlie scNcii taxa nicniioned abo\c. On the other 
hand, if the characters ehminated (see above: 26, 
100, 116) are maintained, some of the trees show 
Acamptopoeum as the sister group of the whole 
tribe, ahhough there are only two characters that 
would support this arrangement. 
The subgenera of Spinoliella and Callonychium 
agree in all characters shown in Table 2 and 
therefore were not segregated in the cladistic 
analysis. 
The two outgroups of the Calliopsini, Protomeli- 
turga and Perdita, were considered here to show 
how they relate to the tribe Calliopsini. A prelimi- 
nary study that did not include Protomeliturga 
because it was not then available, indicated that 
Perdita was the sister group of the taxa now 
included in Calliopsini. 
APPENDIX. A NEW SPECIES OF CALLIOPSINI FROM ARGENTINA 
Calliopsis (Liopoeodes) xenopous, new species 
Diagnosis. This is the only species of Liopoeodes 
and can be recognized by the subgeneric charac- 
ters. Its general appearance is like Liopoeum but it 
is mostly black and not so hairy. Male. With 
apices of forewings brown as in C. (Hypomacrotera) 
and some species of C. (Perissander); tarsus 3 
modified. Female. With hair bands on metasomal 
terga. 
Male. Length 6-8 mm. Forewing length 5.5 
mm. Coloration. Head and thorax black; face very 
slightly blue metallic and the following parts 
yellow: clypeus (lateral area), lower paraocular 
area beside inner orbit (usually L-shaped spot), 
supraclypeal area, underside of antennal flagel- 
lum (upper side brownish black), foreleg (on 
tarsus, outer surface of tibia and apex of femur), 
sometimes also with tiny spot at apex of femur 2 
and at base of tibia 2. Tegula brown. Wings 
slightly yellowish; veins, pterostigma and fore- 
wing at apex brown. Terga brown-black, slightly 
metallic blue; sterna brown. Punctation. Labrum 
with basal area smooth, shining, impunctate. 
Clypeus shining with punctures fine and rather 
abundant. Rest of face with integument slightly 
microareolate, punctures mostly smaller than on 
clypeus. Gena with punctures even smaller. 
Rather sparse and fine on thorax, denser and 
very fine on upper half of metapleura. Propodeal 
triangle basally with fine and weak striae; shin- 
ing, smooth posteriorly. Metasomal terga shiny 
with punctures small, but abundant, intermixed 
with sparser and coarser punctures; punctation 
coarser and deeper on T7. Metasomal sterna with 
punctures sparser and somewhat larger than on 
terga, mostly present distally toward middle line, 
except very fine distally on S5 but absent on 
middle line. Pubescence. Relatively sparse, fine, 
mostly brownish or black (whitish on venter of 
thorax and metasoma); hairs medium-sized 
(nearly as long as or longer than antennal scape), 
similar to those oi Liopoeum on head and thorax, 
somewhat abundant on clypeus and between 
antennal sockets; short on labrum, forming a 
median tuft below labral ridge. Metasomal terga 
with hairs mostly very short, appressed, rather 
dense (similar to those of Hypomacrotera); not 
appressed, somewhat longer, sparse, and dark on 
latero-distal parts of terga, denser on T6 and T7; 
short, not appressed, and sparse (mostly absent 
on middle line) on sterna. Structure. Head about 
twice as broad as long (8.3:4.2). Inner orbits 
sinuose, convergent below. Clypeus more than 
twice as broad as long (4.5:1.9); distal projection 
(beside labrum) slightly rounded at apex. Inter- 
alveolar distance almost as broad as alveolo- 
orbital distance 1.1:1.3). Inner subantennal 
suture angulate, less than half as long as outer 
and slightly shorter than width of subantennal 
area. Frontal line with upper part in a shallow 
groove, lower part (little more than half of total 
length) a well-marked carina reaching inter- 
alveolar area. Facial fovea five times longer than 
broad and 1/4 length of eye (1.0:0.2:4.0). Al- 
veolo-ocellar distance about four times longer 
than distance between alveolus and epistomal 
suture and about as long as antennal scape 
(2.1:0.5:2.2). Ocellorbital distance about four 
times greater than that from lateral ocellus to 
vertex (1.9:0.5). Alveolus diameter similar to that 
of median ocellus (0.6:0.6). Antennal flagellum 
slightly shorter than head, flagellomeres longer 
than broad except second about as long as broad; 
two first flagellomeres somewhat shorter than 
remaining ones. Gena (lateral view) slightly 
broader than half of eye width (laterally) 
(1 .3:2.1). Pterostigma slightly more than twice as 
long as broad (1.2:0.5). Prestigma about 1/2 of 
pterostigmal length and 1/2 pterostigma width 
(0.7:1.2:0.25:0.5). Marginal cell more than four 
times longer than broad and longer than distance 
from its apex to wing tip (4.5:1.0:3.1). Thorax 
narrower than head (7.0:8.3). Mesoscutum more 
than twice as long as scutellum and almost five 
times longer than metanotum (3.7:1.6:0.7). Pro- 
podeum with basal part almost horizontal. Leg 1 
without modifications. Leg 2 with tarsal segments 
much more elongate than those of forelegs; basi- 
tarsus more than ten times longer than broad. 
Leg 3 with tarsus clearly modified; basitarsus 
much shorter than middle one, and wider and 
with strong outer projection at apex; tarsomeres 
2-4 much shorter and much wider than those of 
middle legs, with projection similar to that of 
basitarsus, 3 and 4 widened toward inner side; 
distitarsus longer than those of fore and middle 
legs and widest medially instead of at distal end 
(Fig. 22K). Metasoma (maximum width) narrower 
than head and slightly broader than thorax 
254 
The University of Kansas Science Bulletin 
(8.3:7.0:7.3). Fovea of T2 almost invisible and in 
ventral position. T8 as in Figure 22G. S5 with 
rather short median distal projection; apical hairs 
dense and somewhat longer than projection (Fig. 
22C). Genitalia and associated sterna as in Figure 
22A, B, E, F. 
Female. Length 8.5 mm. Forewing length 6.2 
mm. Coloration. Black, except mandible with 
small basal spot yellow, apically mostly ma- 
hogany; antennal flagellum (extended forward) 
with upper side brown-yellowish, underside dark 
brown; tegula, legs, pterostigma, and wing vena- 
tion brown. Wings slightly brownish, infuscated 
on tip of forewing. Punctation. Mostly fine as in 
male but much denser on metasomal sterna. 
Pubescence. Similar to that of male but not as dark; 
mostly shorter than antennal scape on face. White 
hair bands distally on T2-5 (interrupted medially 
on T2). Structure. Headless than twice as broad as 
long (8.4:5.6). Inner orbits slightly sinuose, 
slightly convergent below. Clypeus about three 
times broader than long (5.5:1.8); marginal pro- 
jection as in male. Interalveolar distance nar- 
rower than alveolo-orbital distance (1.1:1.6). 
Inner subantennal suture curved toward middle 
line, little less than half length of outer subanten- 
nal suture and slightly shorter than width of 
subantennal area (0.5:1.1:0.8). Frontal line with 
distal carina shorter than in male. Facial fovea 
about four times longer than broad and about half 
as long as eye (2.2:0.5:4.2). Alveolo-ocellar dis- 
tance about five times longer than distance be- 
tween alveolus and epistomal suture and slightly 
shorter than antennal scape (2.1:0.4:2.5). Ocello- 
orbital distance about nine times greater than 
that from lateral ocellus to vertex (2.9:0.2). Al- 
veolar diameter similar to that of median ocellus 
(0.5:0.5). Antennal flagellum shorter than head. 
Flagellomeres of similar length about as long as 
broad except first and last two somehwat longer. 
Gena (lateral view) narrower than eye laterally 
(1.3:1.8). Pterostigma slightly more than twice as 
long as broad (1.4:0.6). Prestigma somewhat 
longer than half of pterostigmal length and 1/2 of 
pterostigma width (0.9:1.4:0.3:0.6). Marginal 
cell five times longer than broad and longer than 
distance from its apex to wing tip. Thorax nar- 
rower than head (7.3:8.4). Mesoscutum slightly 
more than twice as long as scutellum and about 
four times longer than metanotum (3.5:1.6:0.8). 
Propodeum similar to that of male. Legs with no 
modifications. Middle femur, on ventral margin, 
with well-defined brush on basal half. Middle 
tibial spur somewhat shorter than basitarsus 2, 
almost straight, with small teeth. Metasoma about 
as broad as head and broader than thorax 
(8.5:8.4:7.3). T2 with lateral fovea weakly 
marked, narrowed at ends. S6 and sting appa- 
ratus as in Figure 23C, E. 
Type material (all from Argentina). PROV. JU- 
JUY: Holotype male and allotype female, 
Huacalera, 17 km N Tilcara, 2800 m, 1-6-1972 
(D. J. Brothers) (KU). Paratypes. 35 males and 83 
females, same data as holotype; 3 males, 
Huacalera, XI-3-5-1968 (C. C. Porter); 2 males, 
same locality, X-10-1975 (G. M. Bohart); 1 fe- 
male, Posta Lozano, XII-15-17-1967 (C. C. Por- 
ter); 10 females, end of Garganta del Diablo, 
Infiernillo, Tilcara (no date indicated) (Solbrig); 
3 males and 1 female, 10 km S Humahuaca, 2900 
m, 1-6-1972 (D. J. Brothers); 24 males and 16 
females, 9 km S Humahuaca, XII-10-1975 (G. 
M. Bohart). Additional paratypes. PROV. 
TUCUMAN: 2 males, San Pedro de Colalao, 
Dpto. Las Trancas, XI-1951 (no collector indi- 
cated); 1 male, Horco Molle, XII-10-23-1967 (C. 
C. Porter); 1 male and 1 female. Las Lenguas, 
XII-27-1972 (J. L. Nefil; 1 female, same locality 
and collector, XII-5-1973. PROV. SALTA: 1 
female, Payogasta, 1-23-1966 (C. C. Porter); 1 
female, Rosario de Lerma, XI- 17- 18- 1983, Ma- 
laise trap (M. Wasbauer). Paratypes have been 
deposited in the following collections: KU; MCZ; 
USDA; UCD; CTMI; AMNH; IZML and 
UCV. 
Comments. A gynandromorph from Jujuy, Ar- 
gentina, was found among the specimens from 
the MCZ. It has the head and legs 1 and 2 (right 
side) like the female; the rest of the body like a 
male. 
Etymology. The specific name is from Greek xeno 
meaning "strange" and pous, foot, referring to 
the modified hind tarsus. It is a noun in apposi- 
tion and does not change with the gender of the 
genus name. 
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