Classification and phylogenetic relationships of the panurgine bees: the Calliopsini and allies (Hymenoptera: Andrenidae)

48 Pages·1991·24.9 MB·

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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 ofthe Tribe Calliopsini 218
Genus Acamptopoeum Cockerell 219
Genus Calliopsis Smith 222
Key to the Subgenera ofthe 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 Ashmead 240
Spinoliella
Key to the Subgenera ofthe Genus Spinoliella 241
Subgenera Spinoliella, 242; Peniella, 242
Genus Callonychium Brethes 242
Key to the Subgenera ofthe Genus Callonychium 244
Subgenera Callonychium, 244; Paranychium, 244
Cladistic Analysis 244
Appendix: A New Species ofCalliopsini from Argentina 253
Literature Cited 254
'Contribution Number3044fromtheDepartmentofEntomology, UniversityofKansas, 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 ofa taxonomic study of the genera and subgenera ofthe andrenid subfamily
Panurginae. Three tribes are recognized and described here: Calliopsini (with two new
subgeneraand one new species), Protomeliturgini(Protomeliturga)and Perditini(Perdita). The
Calliopsini includes Acamptopoeum, Calliopsis (Calliopsis s. str.), C. (Perissander), 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 ofthe 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 ofthe Calliopsini and closely related taxa. At least some clarification
oflines between genera has been attained. A broad interpretation ofthe genus Calliopsis is
proposed. Illustrations for each genus-group taxon are provided.
INTRODUCTION
The family Andrenidae contains two sub- strongly derived group, the Calliopsini,
families: Andreninae and Panurginae. A along with its closest relatives, Protomeliturga
study of the Panurginae at generic and (Protomeliturgini) and Perdita (Perditini).
subgeneric levels, with a classification and The three tribes are restricted to theWestern
phylogeny of the included taxa, was pre- Hemisphere.
sented by Ruz (1986). That study showsthat Robertson (1922) recognized Calliopsinae
the Panurginae is a monophyletic group and as one of the subfamilies of his Panurgidae
that several tribes can be recognized. but otherwise authors have not recognized a
The Panurginae contains numerous gen- higher taxon based on Calliopsis. The Calli-
era and, like the Andreninae, is found in all opsini, as here understood, contains five
continentsexcept Australia. Itsdiversity and genera: Acamptopoeum, Calliopsis (11 subgen-
abundance, however, are greatest in the era, two of them new), Arhysosage, Spinoliella
Western Hemisphere. (2 subgenera) and Callonychium (2 subgen-
The present paper is part of the sub- era). The genus Calliopsis is treated here in a
familial study mentioned above and treats a wider sense than in the past.
MATERIALS AND METHODS
About 20,000 panurgine bees have been the comparable structures being drawn at about the
studymaterialforthisandpreviousinvestigations same size regardless of differences in size of the
(Ruz, 1986). At least three species (3 individuals bees. Sternum 6 of females was drawn with the
each, ormoreifrequired)ofeachpolytypic genus latero-distal margin in its ordinary, curved posi-
and subgenus have been carefully examined and tion in order to avoid breaking it; therefore the
dissected. Structureslike male genitalia and asso- distal part of the sternum appears somewhat
ciated sterna, sting apparatus of female, mouth- narrower than in a flattened sternum. Figures of
parts, last tergum and sterKnaOH5 and 6, were certain structures are omitted ifsimilarstructures
cleared in a cold solution of (10%) for 12 are illustrated for related taxa or ifother sources
hoursorless, dependingon thedegree ofscleroti- with good drawings are available.
zation ofthe structures, then washed with water In the descriptions, each character (or groups
for several minutes and preserved in glycerol for ofmorphologically associated characters) is num-
later examination. bered. Forpurposesofcomparisonthesenumbers
The morphological analysis of the species se- have been maintained within taxa of the same
lected as representatives of each genus (type level. Certain characters(and numbers)are omit-
species plus at least two additional species when ted ifthey are similar in related taxa. Subfamilial
possible)wasmadeusingastereomicroscopeand, characters are omitted in the tribal, generic and
for details, a compound microscope. The names subgeneric descriptions, unless they represent an
ofthe species studied in detail are preceded with exception for the group; in this case those dis-
anasterisk(*)inlistsofspeciesundereachtaxon. tinctive characters are kept in the description.
Illustrations were made using acamera lucida, The number of a given character used in the
Panurginae: Calliopsini and Allies 211
descriptions is not coincident with that used for ing that vein 1st r-m (2nd transverse cubital
Tables 1 and 2 and the cladogram (Fig. 30). vein) has disappeared. When three SM cells
In order to facilitate comparisons, the number are present, cells 2 and 3 are indicated as SM
ofeach apomorphy as it appears inTable 1 and in 2 and 3. M
Ruz (1986) has been maintained for the whole 50. Cell 2nd of forewings: 3rd discoidal cell.
revision of the subfamily. The numbers of the 69. Male terga: Postgradular depression width
characters not used in the Calliopsini and their measured along the longitudinal axis of the
close relatives are skipped in the present work. In body.
order to maintain the sequence of numbered 70. Pygidial plate: Glabrous plate on distal part
apomorphies, certain autapomorphies have been ofT7 ofmale and T6 offemale (the latternot
inserted with a numberalready used plus a letter. considered here), laterally delimited by car-
The morphological terms in the descriptions inae. The same area delimited only by hairs
are mainlv based on those used by Snodgrass or only a projection of last tergum has not
(1935, 1956), Michener (1944, 1965, 1981), been considered as a real plate in this study.
Rozen (1951), Winston (1979), and Michener 71. T7 offemale: HemitergiteofMichener, 1944.
parnedtaBtrioonosksm(a1y984b)e.dSifofmiceultchaarreacteexrpslawihnoesdebienltoewr.- hasInbdeeesncricboinnsgidmearregdinassoefxatelnegdesdeglmaetnerta,lltyh;etlheeg
20. Basal area and apex oflabrum: As indicated antenna has been considered extended forward.
in Figure IF The word vertex has been used to mean the
33 Tentorial pit: Small, rounded, the most dor- posterior margin of the vertex rather than the
sal part ofthe external depression associated whole top ofthe head.
39. Owribtihtst:heIannnteerrioocrutleanrtoorribailtsa,rmasss(Feiegn.i6nA,facDi)a.l abbIrnevoiradteerdtaos Tsavoer Ssppalcues, atrearbglae annudmbsetresrnoafatrhee
view (Fig. 6A-D). metasomal segments (i.e., not counting the pro-
46. Pterostigma and prestigma lengths (Fig. 2E) podeum). Thus Tl is the first metasomal (second
were measured on vein Rs; their breadths abdominal) tergum.
(maximum) were measured perpendicular to
the lengths. Prestigmal width was not meas- ACKNOWLEDGMENTS
ured to the wingmargin but is only the width
ofthe prestigma proper. Acknowledgments for the study material used
48. ppSrauerbsimesanortn),gihnSaasMlbecceeelnlllsca2l(lSe(Midf)S:oMnlFyocerltlwl2oe-nt-gS3thMascscouemlml--s ohinefrett.hhiesTwhsheeoryileems.aryeAvbibbserioeonvmiioatfttietohdnesinsuisnbufbpasameriqelunyetnhaetrsepeasgpievarersne
used for institutions or collections through the
rest ofthe work.
,penis valve This study would not have been possible with-
out the encouragement and assistance ofnumer-
ous people. I am particularly grateful to Charles
D. Michener of the Department of Entomology,
University of Kansas (KU), Lawrence, Kansas,
forhis generosity, advice, interest and help in the
preparation ofthe 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
basal their suggestions in editing the manuscript; Ed-
sclerotization basal ward O. Wiley (KU) for his advice and willing-
sspcilenreo-tiszhaatlpoend ness todiscuss aspects ofthis investigation; Frank
lfaitdegreal Koch ofthe Museum fiir Naturkunde der Hum-
duplication boldt-Universitat, Berlin, for lending Friese's
eWyiedth types and other valuable specimens, as well as
generously permitting study ofmaterial from the
collection under his care when I visited the
Museum; George R. Else ofthe British Museum
(Natural History), London, England; Till Osten
of Staatliches Museum fiir Naturkunde, Stutt-
clypeal protuberance gart, Germany; and Mario Zunino ofthe Museo
gCaelnFliiitogaplusiriaes,(d1Ao.,rsDaBil,aagDnr-daFvm)enattornadlshPveoirewdwistt;aerB(,mC)iS.n8oAlo,ofgmyMaalfleoe;r UseiindmtieavnIessdrti.istiTTutothyor,eidniloNa,tZeeowIoRtlaaoldBygcr,iluayanflsSfswioeisctBkke.i,mnadRtloNiybceealrwetdnsetJlleotarfyspRUeeunytisgvpaeeenrrcd-s-
C, D, S6 of females; E, Head, lateral view; F, John L. Neffof the Central Texas Melittological
Labrum. Institute, Austin (CTMI), lent and donated im-
212 The University of Kansas Science Bulletin
portant specimens. Professors Salvador Peris and Peter Schultze and Arturo Roig-Alsina (all from
Elvira Mingo of the Institute Espanol de Ento- KU) for their help in translating materials from
mologiaofMadrid, Spain, permitted my studyof German to English.
that collection. Yu. A. Pesenko ofthe Zoological The drawings were mostly inked by Donna
Institute, Academy of Sciences, Leningrad; Stevens (KU); some ofthem were also prepared
Klaus Warncke of Ruckteschellweg, Germany; by Carmen Tobar (UCV).
George C. Eickwort of the Department of Ento- The typing of the manuscript and editorial
mology, Cornell University, Ithaca, New York; assistance was by Joetta Weaver (KU). Many
the late Robert O. Schuster of the University of thanksalso toVirginiaAshlock(KU) forherhelp
California, Davis, California (UCD); Paul H. in typing and editorial work.
Arnaud of the California Academy of Science, This study was possible thanks to the financial
San Francisco, California; Wallace E. LaBerge, aid of the Direccion General de Investigaciones,
Illinois Natural History Survey, Champaign, Illi- Universidad Catolica de Valparaiso, Chile, and
nois; and William L. Overal ofthe Museu Para- especially the National Science Foundation
ense Emilio Goeldi, Belem, Para, Brazil, lent grants DEB82-12223 and BSR87-16817 (C. D.
important study material. RonaldJ. McGinleyof Michener, principal investigator).
the Smithsonian Institution, Washington, pro-
vided material from the Reed Collection then SUBFAMILY PANURGINAE
under his care at Harvard University, Cam-
borfitdhgee,SMnaoswsaEcnhtuosmeotltosg(icMaClZ)M.usReoubemr,tUWn.ivBerrosoiktsy metDailalgincosiisn.teSghuomretn-tt,on2g-u1e1dmbemeslounsgu;alfliyrstwistehgmneonnt-
ofKansas, Lawrence, Kansas(KU), helped in the of labial palp usually flattened; labrum (prox-
preparation ofdiagrams, made some preliminary imally) without lateral process to articulate with
photographs and provided certain specimens for clypeus as in Andreninae; paraglossa elongate
Astgurdiyc.ulTteurrre,y GArRisSw-oBledeofLtahbeorUa.tSo.ryD,epUatrathmenSttatoef asnusdpsenosmoerwihuamt otrapelroendgedrist(ailnly,Anusduraelnliynaaselownigdears
mUantievreirasli,tym,adLeogaavna,ilaUbtleahthe(pUaSnDuAr)g,inelecnotllescttuidoyn dsiusttuarlelyusaunadllsyhoarbtseernt;thmaanrgsiunsapelncseolrliduims)ta;llsyctrroubna-l
ofthe Bee Laboratory and kindly provided hous- cate (often obliquely); middle femur of female
oNifnegwtohenYomArmyke(rviAisciMtaNsnHto)M,Luoscgeoannut.mriJbeoufrtoeNdmaetvuaGlr.uaalRbloHeziesitndoeraJysr,., waminatidhnslvpyeanrotcnreatlpiblmiuaamraognsidenbhaaasiriritsdagrdesi,usstwa,iltlmyho;daesrccaooptmaeblpyrbeaassbeaunlntl-,y
lent many specimens for study, permitted the use dant to sparse; T2 with lateral fovea or at least a
okfintdhley pbereovicdoleldechtoiuonsionfgtahnedMhuelspeuwmh,enaIndvisvietreyd daa"rkgosnpootb;asgeo"notbhaastemaabysenntot(ebxecehpotmo3lgoegnoeursa wwiitthh
the Museum. Zine Ajmal de Toledo of the In- the real gonobase).
stituto de Zoologia Miguel Lillo, Tucuman, Ar- The three panurgine tribes considered in the
gentina (IZML), lent an important panurgine present paper, Perditini, Protomeliturgini and
collection under her care. Haroldo Toro of the Calliopsini, share the following principal charac-
Universidad Catolica de Valparaiso, Chile ters: 47. Submarginal cells two (exceptionally
(UCV), made valuable suggestions and provided three in Perditini). 66. Tibial scopa of female
numerous specimens. Dr. James S. Ashe (KU) usually (at least partially) of minutely branched
and Francisco Saiz (UCV) provided help and let hairs. 82. Gonocoxal apodeme not inflexed. 88.
meuse photographicequipment undertheircare; Sting short to extremely short, with first valvula
Carlos Gonzalez (UCV), Chile, and Stephen slightly sclerotized, without valve (except a pos-
ReyIeasm(KaUls)odiinddesbotmeedpthootRougdroalpfhiJcanwdoerrk,. Hans- sible rudiment in Calliopsis (Ceroliopoeum).
Key to Tribes Calliopsini, Protomeliturgini and Perditini
1. Marginal cell (nearly always) about halfas long as distance between its apex and wing tip
(Fig. 2B); iflonger, thenlowerpart efface, in profile, not orscarcelydivergent from eye. S6of
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 halfas longasdistancebetween its apex and wingtip. Lowerpart of
face, in profile, distinctly divergent from eye. S6 ofmale usually less than 3 times wider than
long; if wider, then distal margin with tapered projections and rather deeply emarginate
medially; or ifotherwise, ofdifferent shape than above. Almost never metallic 2
2. Tentorial pit at intersection of outer subantennal and epistomal sutures. Pre-episternal
grooveextendingbelowscrobal level. Middle femuroffemalewith hairsofventral margin not
well organized in a comb Protomeliturgini
Panurginae: Calliopsini and Allies 213
—
Tentorial pit in outer subantennal suture (Fig. 6A-D). Pre-episternal groove short, absent
below scrobal level. Middle femur of female with a distinct, well-detined comb basally on
ventral margin (Fig. lOD) Calliopsini
TRIBE P1:R1)1UNI ot outer subantennal and epistomal sutures. 45.
Pre-episternal groovecurved, meetingscrobe. 46.
This tribe contains a single enormous genus, Pterostigma large, with sides divergent; margin
restricted to North America. It has not been within marginal cell clearly convex. 47. Marginal
mreocnloapsshiyflieetdiicn.tIhnetphreesceunrtrenwtorckl;assitifiiscadtiivoenrsitecbount- cwaepiletl.hxudsaounradslallwyibnhoagrldfteirpasuosrluoalnelsglsy.aus6n1t.doisoHttiahnnedcde.tib6b9ei.tawToe2fe-mn5aliotefs
tains only the genus Perdita, whith is suljciivitled male with posterior marginal areas glabrous or
into 21 subgenera. nearly so. 72. T8 ofmale usually tapered distally
2.TIhnetefgoulmleonwtingusfueaaltluyremsetcahlalriacctaenrdizewitthhisyterlilboe:w barnodadhlayircyo.nc7a6v.e.S577.ofS6feomfamlealweiatbhoudtisttahlremeatrigmiens
markings. 33. Tentorial pit usually at intersection as broad as long (length measured on midline).
82. Gonocoxal apodeme not inflexed. 83. Gono-
mca
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)
PerditaSmith, 1853: 128. Typespecies: Perditahalictoides
Smith (monobasic).
The list of subgenera and synonymy for each
can be found in Hurd (1979), with another
subgenus in Timberlake (1954).
I havenot attemptedtostudythesubgenera(or
genera?) encompassed underthe name Perdita. In
the following description, however, I have at-
tempted to indicate the variationwithin 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 ofthat subgenus
have the specified characters: Perdita s. str. {halic-
toides Smith); Pentaperdita {albovittata Cockerell);
Perditella {larreae Cockerell); Hexaperdita {ignota
Cres-
crawfordi Cockerell); Pygoperdita {interrupta
son); Heteroperdita {trifasciata Timberlake, female);
Glossoperdita [pelargoides(Cockerell)]; Hesperoperdita
{trisignata Cockerell); Alloperdita (novaeangliae
pr Pt Viereck, female); Macrotera [texana (Cresson), bi-
color Smith]; Macroterella (mortuaria Timberlake,
male); Macroteropsis{latiorCockerell); Pseudomacro-
tera {turgiceps Timberlake); Allomacrotera (steph-
anomeriae Timberlake, male); Epimacrotera (ainsliei
Crawford); Callomacrotera {maritima Timberlake,
Figure 2, A-E. Forewings of males. A, Proto- acapulcona Timberlake); Cockerellia {albipennis
meliturga turnerae (Ducke); B, Perdita halictoides Cresson); Cockerellula {opuntiae Cockerell); Pro-
Smith; C, Acamptopoeum submetallicum (Spinola); cockerellia {albonotata Timberlake); Xerophasma (be-
D, Calliopsis (CeroUopoeum) laetum (Vachal); E, quaertiana Cockerell). Additional information on
Callwpsis (HypomacTotera) callops (Cockerell and charactersand theirvariabilityamongandwithin
Pmocrate=r)m.arginpatl=cpeltleraopsetxi;gmbav;=basparl=vepirne.stigma; Dsuabngfeonretrha(1o9f91Pe)r.dita is given by Ruz (1986) and
214 The University of Kansas Science Bulletin
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, veryshort, usuallynearlyhalfaslongas
distance from apex to wing tip (Fig. 2B) (not as
short inXerophasma andMacrotera). 48. Submargi-
nal cells two except in Xerophasma and Alloperdita,
which have three, but seconMd small and petiolate
anteriorly. 50. Cell second sometimes absent
or weak. 51. First recurrent vein usually close to
first transverse cubital, sometimes meetingit. 55.
Propodeal triangle with no visible hairs (some-
times with sparse, minute hairs [e.g., P. (Al-
lomacrotera)stephanomenae, Epimacroteraainsliei, 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,
Figure 3. Perdita halictoides Smith. Male: A, B, slightly curved (or almost straight), somewhat
Genitalia, dorsal, ventralandlateralviews; C, D, shorterthan basitarsus 2; stronglycurved at apex
dSo8r,sadlorasnadl,vevnetnrtarlalviaewnsd; lFa,teTr8al; Gvi,ewSs6;, vE,entSr7a,l irtnohtaMrnaiccrhtoiwtaiecreoa.na6sm1.olsTotingboifaasi3nbonafesrfietsmaurarslfueascse,o3,meswwoimhteahttikmleeeiss-s
view. Female: H, Sting; I, Sting, ventral view; J, very sparse toward the middle, only at ends, or
S5, ventral view; K, T7; L, S6, dorsal and apparently absent; male with dorsal margin of
ventral views. tibia 3 usually untoothed, with keirotrichia usu-
ally on most of surface, sometimes sparse. 62.
Description. 1. Length 2-11 mm. 2. Integument Tibial scopa usually ofrather short hairs (long in
metallic (at least partially), except in species of some species); in most cases apparently simple
the subgenera Macrolera, Aiacroterella, Xerophasma, but with minute branches, with ratherlong alter-
Perditella, Cockerellula. 5. Pubescence in general nate branches in P. (Callomacrotera) maritima Tim-
short, usually minute on most of metasoma, berlake; scopal hairs usually sparseormoderately
especially on terga. 8. Head narrower to wider abundant, dense in Macrotera and Callomacrotera.
than thorax. 11. Glossa much shorter to longer 63. Hind tibial spurs usually slightly curved,
than prementum; usually with flabellum. 17. strongly curved at apices as in Macrotera; outer
Galea! comb of0-15 setae. 20. Labrum less than spur usually shorter than inner. 64. Basitibial
twiceasbroad aslongto morethan twiceaslong; plate of male well defined. 66. Claws bifurcate.
basal area usually glabrous, sometimes almost 67. Metasoma in male usually wider than to
fully pilose; distal margin of basal area a ridge; sometimes narrower than thorax. 70. Pygidial
labrum somewhat protuberant, with apex in- plate ofmale absent, though T7 at apex usually
flexed or not. 25. Mandible of male simple to with median projection (projectingarea truncate,
bidentate, or with prebasal tooth-like projection. bifurcate, or tapered). 71. T7 of female as in
30. Inner subantennal suture usually curved, Figure 3K. 72. T8 of male generally somewhat
sometimes almost straight. 33. Tentorial pit usu- tapered at apex, distinctly hairy (Fig. 3F). 74. S4
ally at intersection of outer subantennal and of male with distal margin broadly and gently
epistomal sutures (sometimes in outer suture concave or almost straight. 75. S5 of male with
though close to the epistomal suture). 34. Anten- distal margin widely and gently concave (some-
na! sockets below middle of face [e.g., P. (P.) times almost straight). 76. S5 of female with no
halictoides Smith] to slightly above as in Xe- median sclerotized area proximally and no gra-
rophasma. 36. Antennal flagellum of male un- dulus; distal margin broadly and shallowly con-
modified, shorterto longer than head. 37. Lower cave(Fig. 3J). 77. S6 ofmale threeormoretimes
mesal paraocular area rather flattened or slightly wider than long (length measured medially), dis-
convex as in P. (P.) halictoides (sometimes some- tally with a very wide V-shaped emargination or
what protuberant). 39. Orbits usually sub- margin almost straight (Fig. 3G). 78. S6 of
parallel, sometimes convergent below as in female with no proximal laminar lobes, basal
Panurginae: Cai.i.iopsini and Allies 215
sclerotization fully fused to sternum and not basal area hairy laterally. 33. Tentorial pit at
spine-shaped (similar to that of Protoineliturga), intersection of outer subantennal and epislomal
lateral margin with ridge almost straight or sutures. 45. Pre-episternal groove extending be-
curved, longitudinal basalcarinaabsent, duplica- low scrobal level. 46. Pterostigma with sides
tion membranous (sclerotized area below it), diverging; margin within marginal cell convex.
distal margin distinctly concave or narrowly 61. Hind tibia of male with dorsal border a
emarginate medially, S6 with small sparse hairs carina. 69. T2-5 ofmale with posterior marginal
proximallv, forming dense patch at both sides of areas pilose laterally forming a hairband. 70. T7
midline distally (Figs. IC, 3L). 79. S7 of male ofmale with a stronglateral projection. 76. S5 of
with two distal lateral lobes and with shallow to female with distal margin almost straight. 82.
deep V-shaped or concave apical emargination Gonocoxal apodeme not inflexed. 83. Gonocox-
(Fig. 3E). 80. S8 of male cross-like with distal ites short, fused ventrally. 85. Volsellae well sepa-
part wide to narrow, truncate, rounded or ta- rate from each other, without denticles. 87. Penis
pered; basal part wider to narrower than distal extremely wide and completely fused to penis
projection (Fig. 3C, D), bilobed, or bifurcate in valves. 88. Sting very short, first valvula little
some species. 82. Gonocoxal apodeme well or sclerotized, without valve.
usually strongly developed (laterally or medially
in ventral view), not infle.xed (Fig. 3A, B). 83. Genus Protomeliturga Ducke
Grootniozceodxciutteiscleeltoongfautlel,y cfuosnendecvteentdrablyly.sli8g4h.tlGyosncloe-- (Figs. 2A, 4, 5, 30)
setlvolnugsatef.us8e5d. Votloselgloaneocaotxtiatceh,edwteolleacdhevoetlhoeprebd,y ProotposmieslittuurrngearaDeuDcukcek,e,191192:0763(,mo9n0o.baTsyipc)e.species: Calli-
small membranousarea; withdenticles. 86. Penis Diagnosis. Segments 3 and 4 of labial palp at
valves elongate, simple, usually tapered apically, right angle to segment 2. Basal vein (forewing)
fused to each other generally rather extensively strongly curved. Terga with distal hair bands
(sometimes free only at apices). 87. Penis mem- laterally. Male. Antennal flagellum much shorter
branous, usually almost aslongasand widerthan than length ofhead. Tibia 3 with dorsal margin a
valve, fused to valves in great part. 88. Sting sharp carina. T7 at apex strongly bent down and
short(sometimestruncate, e.g., P. halictoides), not forward, with a conspicuous projection laterally.
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 ofall the
diversitythatitpresents, hasseveralapomorphies
that make it adistinctive group. Accordingto the
cladogram (Fig. 30) this genus is the sister group
of the Calliopsini and Protomeliturga together.
Rozen (1966) has found that Perdita has distinct
larvalcharactersdifferent from those ofanyother
Panurginae.
The pollen balls in thecells ofseveral species of
Perdita are coated with a secreted, cellophane-like
layer, a synapomorphy shared with the Calliop-
sini(Rozen, 1967, andpersonalcommunication).
liProtomeliturgadoes notcoverthepollenball, this
fact may mean that Perdita, hot Protomeliturga, is
thesistergroupofCalliopsini. Thiswasthe result
that I obtained in a preliminary cladistic analysis
using a smaller set ofcharacters. w
Distribution. This genus occurs in southern G \-y H
Cmaanlaad(aH,urtdh,e U1n97i9t)e.d States, Mexico and Guate- A,FBi,guGreeni4t.alPirao,todmoerlsiatlu,rgvaentturrnaelraaend(Dluactkeera)l.vMiaelwes:;
TRIBE PROTOMELITURGINI C, S7, dorsal and ventral views; D, S8, lateral
view; E, T7; F, Leg 3, outer view; G, S8, dorsal
The main characters of this monotypic tribe and ventral views; H, S8, ventrolateral view,
are the following: 20. Labrum of female with tc=tibial carina.
216 The University of Kansas Science Bulletii
Female. Middle tibial spur finely and densely cu-v 1/4 to 1/5 as long as second abscissa of
toothed. Tibia 3 more than twice as long as M-I-Cu. 54. Dorsal surface ofpropodeum about
basitarsus 3, with scopa mostly oflong, minutely as long as metanotum. 55. Propodeal triangle
branched hairs. Inner hind tibial spur conspic- densely pilose, especially on lateral areas, with no
uously curved. striae. 57, Basitarsus 1 about six times longer
uyfPrealaulcatbDlerheeosews(cro.ccrfileapyn4btpf.ciueeoenumnMsd.aeialont1enf)ta.gm,Lseaeaonlninemengrda,catolhsawnmris5saetp-hlahoi8lsrctmusn,pooomofu.mtsyoteis3hinl.tnolllrLoymoawooxwwaesmeprtaaprrnprokdpaefsarsr.asdltoeoecdrg5o-,.-sf tFlct59ohheo.naammgnnuMbeirhrbandrilo2donftlaoafedfwseemftlliaeilnoblmniedaagm;lelaevatslesapoelrubnosarapsvoenioemddtfneatrbarrmeosnasotuldrhs2men-2sao4ertwtxgiuhecitnaslhnmnesoalfbdrtiialihngsyferha,teidlleeelmydfyit.imnwinouiem5rtdte8e..ehs
sum ofthorax. 6. Integument microareolate, dull and dense teeth. 60. Basitarsus 2 of both sexes
on most ofhead and thorax, shiny on metasoma. about as long as 1 and shorterthan 3. 61. Tibia 3
t7a.shabPnruonlacodtnugarsaesntdhfoirnnaaex,r)r.moow1se1t.rlyGtlhdoaesnnssateh.soor8ma.xewH(hesaaotdmeltwoiinmdgeeesrr o(sFfuirfgf.eamcae5lHee)x,mcoepwrtiethctlhoakseneirttowotircdieocrhasisaallooannngdmasovsebtnatsrioatflarismnuansre-r3
ltohnagnerprtehamnen2t-u4m.toge13t.herS.eg1m4.enSteg1meonftla2bioafllapbailapl gsihnasr;pmcaarlienati(baisai3n Pwsietuhdodpoarnsuarlgusm)ar(gFiign. a4F)s,trwointg,h
mpms2aeeo(lgnrapmteseainbtn3tohlau2oontfnsogttl-htawtrhbioeaictaneelg3utaeipasmdaneldlspboe4nelasgpor)rn.togaijesce1ru6cl.tpatrtahMeetapadnaaxlniapgl3ta.llleaar1tn5ypo.galbr1etSlaeangdtod-eof okdwtiiefhsbiatilraaotollntlgryss,,ihpcouabhrrirtfsaaeenrwawcsihottienhhndadnmfohieramnisiaunarltnlseee.m(,ras6orfa2imgn.neideTnwitsibseltieiitahmghlph;bltelsryo)ca.uontcpc6euah3rre.mvssoeHosodimtnnellad-yyt
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
slightlymorethan twice aswideaslongin male, a
little less than three times in female; moderately
protuberant (about 1/3 width of eye in lateral
view); distal marginwith roundedprojectionnear
lateral margin of labrum (Fig. 51, K, L). 30.
Inner subantennal suture almost straight. 31.
Subantennal area wider than halflength ofinner
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 paraoculararea slightly convex. 38.
Facial fovea narrow and shallow in male, wider
andwell-markedin female. 39. Orbitsconvergent
below in male, subparallel in female. 40. Ocelli
mostly above orbital tangent. 41. Vertex convex.
42. Gena of male (lateral view) narrower than
eye, ofrather 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 Figure 5. Protomeliturga turnerae (Ducke). Male:
within marginal cell convex. 47. Marginal cell A, 86, ventral view. Female: B, Labrum; C,
rather pointed at apex (obliquely truncate. Fig. Labrum, lateral view; D, S6, dorsal and ventral
2A); lengthlittle shorterto somewhat longerthan views; E, T7; F, Sting; G, S5, ventral view; H,
dncFaiiuasrblsscitetctaleanlllrcl.e2sec-t1uf5-wrr23oro...emn54t0F9ita.osv.reSaeiBupwnabeismxdnaaigltrsotgvawewiniinitntanhlgfsrtctorceiolumpl.n-gvf14li8ryas.lbtcooSuntuurgrtvbeaermnadsas.vrtleghor5ian1s-nge. LalHnaeedbgardlu3,,ambfoorrunoutlnmetyraolimvniiateKntw)de;.d)ltI.ap,t=eHMraeatllaenedtv:,oireJifw,arsloAnpn(tihattal;einrvcnsidaepo;wm=iK(tdh,itaseitdLra,,sl
second abscissa of M-fCu. 53. Hind wing with clypeal projection.
Panurginae: Calliopsini and Allies 217
apex; inner more ccmspiciiously cui\ed, espe- from the two museums are certainly conspecific.
ciallv in female. 64. Basitibial plate of male well Discussion. The cladistic analysis shows this
defined, margins carinate. 65. Tarsus 3 with no species to be the sistergroup ofthe Calliopsini. It
sirmnmoaadmliiflfeimscaauaplbtieeic.qaoulnas6l7p.reoixjnceMecmeptattiloaen,sb.aosim6in6atn.aerrsiCunlsshaomworasftleerdfeetemshpaoallmnyeeowcwulheitaftetthr; pfpiraresntsuersngitigsnhtea.tshePerrioegsbeanobuflsydesrbuiegvcgeeadsutsfseeaatoufrreaistt,shealralttpehrroialumigthhiaviaetr
broader than thorax. 69. T2-5 of male with bands on the metasomal terga, Ducke (1907)
gradulus posterolaterally absent and with erroneously placed it in Calliopsis.
postgradular depression rather shallow; posterior Distribution. Prolomeliturga is known only from
marginal areas of Tl-5 in male and 'ri-4 in Sao Luiz de Maranhao, Brazil.
female pilose, forming bands laterally (band less
distinct on T5 of male and T4 of female); lateral TRIBE CALLIOPSINI
fovea of T2 (both sexes) rather narrow, gently
dcoatefopnarsfpepeesimxscaeusldtoe.ruos7an0sgp.lryoiPjnycegucirtFdviiioegandulrldeapotleaw5rtnaEel.laoynf7d(m2F.aifglo.erTw48aaErb)ds.oe,fn7wt1i;.mtahTTl77ea usrpghTeihrnieas,e.tirsIibtelh,isetfmbooeuslntodwdoenarliysveeridinesgtrhoeofuwpitessotfcehratnrhaechtePemarins--,
trapezoidal. 73. Metasomal sternaofmale mostly using the same series ofcharacter numbers as in
withhairsstraight, short, appressed, exceptonS6 the generic descriptions, in order to avoid repeti-
obliquely directed toward midline. 73a. SI-5 of tion in the descriptions ofthe genera as well as to
female with hairs as in male but longer and emphasize the unity ofthe tribe.
branched on premarginal areas. 74. S4 of male 30. Inner subantennal suture about as long as
with distal margin slightly and broadly convex. antennal socket diameterorshorter. 33. Tentorial
75. S5 ofmale with margin straight medially. 76. pit in outer subantennal suture. 45. Pre-epister-
S5 offemalewith neithermedian proximal sclero- nal groove rather short, absent below scrobal
tized area nor gradulus; distal margin broadly level. 46. Pterostigma basally nearly as wide as
and gendy concave (Fig. 5G). 77. S6 of male distally or little narrower; side within marginal
broadly and gently concave medially (Fig. 5A). cell usually straight. 48. Submarginal cells two.
78. S6 of female similar to that ofPseudopanurgus 55. Propodeal triangle glabrous. 58. Middle
except longitudinal basal carina absent; duplica- femur of female, on ventral margin, with a
tion well sclerotized; distal margin slightly con- distinct, well-definedcomb. 61. Tibia3 offemale,
cave medially; premarginal area fully hairy but on innersurface, with keirotrichia present toward
sparsely so medially; rest of sternum with hairs dorsal border and at base and apex, sometimes
minute (Fig. 5D). 79. S7 of male wide, distally only at the two ends; male tibia 3 with dorsal
withaV-shapedemarginationandalateral, short border untoothed. 62. Tibial scopa, on outer
projection; proximal arms forminga U (Fig. 4C). surface, of sparse to extremely sparse, mostly
80. S8 of male wide, large, strongly carinate minutely branched and simple hairs. 69. Tl-5 of
laterally and much exposed distally (surpassing male and T1-4 offemale with posterior marginal
T7) (Fig. 4D, G, H). 82. Gonocoxal apodeme area usually completely or partially hairy (short
well developed (dorsal view), rather wide, not hairs). 70. Pygidial plate ofmale usually present.
inflexed(suturealmost invisible)(Fig. 4A, B). 83. 72. Tergum (T)8 ofmale usuallyabout aswideas
Gonocoxitesmuch shorterthan penisvalve, fused long and roughly hexagonal but with lateral
medially. 84. Gonostylus fused to gonocoxite, angles rather rounded. 73. Sl-5 of male with
slightlyshorterthan halflengthofgonocoxite. 85. hairs mostlydirected straight backward. 76. S5 of
Volsella strongly lateral in position, free mesally, female with a median sclerotized area between
fusedto gonocoxiteproximally, withoutdenticles. proximal margin and gradulus except in Callo-
86. Penis valves wide, completely fused to each nychium (Fig. lOE); gradulus usually short except
other and to penis. 87. Penis long (slightly sur- in Callonychium long and recurved; distal margin
passing penis valve), extremely wide, with two broadly convex medially. 78. S6 of female with
msheosratl, nlootngrietaucdhinianlg sstcylleurso,tifizresdt vaarlevausl.a8l8it.tleSstcilneg- spcrloexriomtiazlatiloanmienxacreptloboefs;difwfietrehntbasshaalpespiinnes-loimkee
rotized, valve absent (Fig. 5F). Calliopsis(Nomadopsis) and absent in Callonychium;
sttpyoeapCTcleoihlmeoesmot,efymnpaPtePrts.oem.lraoiltmTaueehrll)iniestafruvrargaeogiemaln(aultlbushelrecentieMsorftuaokyersnp(eeotDhuwuifnscPekasmeortan)aull.edeynysfaewrnaoEsdmm:iolltenihceoe- bdlsagilaoioslsnnmytagaeliwlatiltuCytsadhlcli(ellneaeaisaxrotlcopsesttbsipirasztoosamnsatf.geliu,ow1scn.he;audasrmftuioroanenslaseltcaAylrbfeohsarrfyeolosnstmmotuios;zrsaefdgtddaieucs)speec;tlxriwclaocieaiftatpgeththrsitaotmtlnehiriimrnundanusgrutuei-me-;n
Goeldi (Belem, Para, Brazil) (not dissected) and sparse hairs except apically with well-defined
two specimens (one male and one female) from rows ofdense hairs forming a lateral patch or a
the Berlin Museum which, although labeled continuous, usually curved, band ofhairs. 80. S8
"Typus," are not the real types. The specimens of male with distal projection well developed;
218 The University of Kansas Science Bulletin
basal body with a lateral acute projection except sclerotized, valve absent [except a rudiment of
in Acamptopoeum and Calliopsis (Verbenapis), which valve in Calliopsis (Ceroliopoeum)].
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 cdp
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
acsloumbmxas.iletlne,t8?6)no.raarsPrreuiondnwiisAmrbehrvnyiastdloagvserea.ysge(8,d8co.oSrpmsipSanltloiellintyegelllfuyaussfuaeuandlsdletydChatrsloholouogrngotyhncooh-a-r Any,cFhSip=igiunumorleiel7(l,aCaA(l,PleonBni.eylcHlhaei)uammd)ascuolfamtamanadl(ieSbspui,lnaovrleean=)t;raBBl,rveCitaehlwelsso..-
greatly reduced (except surpassing stylus in cdp dista=l clypeal projection; es= epistomal
Acamptopoeum); second valvifer (triangular plate) suture; c clypeus, lateral area; p paraocular
usually unmodified; first valvula only slightly 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 ofmales, frontal views. A, Acamptopoeumsubmetallicum (Spinola); B, Calliopsis
(Calliopsis) andreniformis Smith; C, Arhysosage ochracea Brethes; D, Callonychium (Paranychium) chilense
(Friese). tp=tentorial pit; ff=facial fovea.