science bulletit - Atlas Hymenoptera

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THE UNIVEI3SITY
OF'I(ANSAS
SCIENCE BULLETIT{
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Systematics and Phylogeny of the
Anthophorine Bees
(Hy*enoptera: Anthophoridae; Anthophorini)
By
Robert Iry. Brooks
V o l. 5 3 , N o . 9 , p p . 4 3 6 - 5 7 5
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September 30, 19BB
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To
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ProfessorCharles Duncan Michener
this study is dedicated
in anticipation of the
aa
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occasionof his seventiethbirthdav
22 September19BB
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and his retirement
1.,
from the departmentsof Entomology
;
F
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and Systematicsand Ecology
18 Mav 1989
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THE UNIVERSITY
OF I(ANSAS
SCIENCE BULLETIN
September 30, 19BB
Vol. 53, No. 9, pp. 436-575
Systematics and Phylogeny of the Anthophorine Bees
(Hymenoptera: Anthophoridae; Anthophorini)
Ronnnr W. Bnoors
CONTENTS
Introduction
Acknowledgem;;,; ::::
Historical Review .
Methods.
Characters Employed.
Tribe Anthophorini . .
Unplaced Species .
incertae sedis . .
Anthophorini
K.y to Genera of the AnthoPhorini
Genus Anthophora Latreille
Kty to Subgenera of AnthoPhora. .
Subgenus Anthophora Latreille s. str.
Pyganthophora new subgenus
Lophanthophora new subgenus
Mystacanthophora new subgenus
Caranthophora new subgenus
Subgenus Paramegilla Friese
Subgenus Melea Sandhouse.
Subgen us AnthophoroidesCockerell and Cockerell
Subgen us Anthomegilla Marikovskaya .
Rhinomegilla new subgenus
Petalosternonnew subgenus
Daslmegilla new subgenus .
Subgenus C/asodon Patton .
Subgenus HelioPhila Klug .
Anthophora incertae sedis . .
Genus Amegilla Friese
K"y to Subgenera of Amegilla
Zebramegilla new subgenus
Subgenus Amegilla Friese s. str.
Dizonamegilla new subgenus
Megamegilla new subgenus
Subgen us Aframegilla PoPov
Ackmonopsis new subgenus
Micramegilla new subgenus
Subgenus Zonamegilla PoPov . .
Notomegilla' new subgenus .
Glossamegilla new subgenus .
+39
+39
++0
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SysrpvrATrcs
AND PHyrocENy
oF THE ANtHopHoRrNB
BsBs
+39
Natuurlijke Historie, Leiden, who allowed me to stay in
his home and study his fine collection of anthophorine
This study provides a classification and
bees. I will remember him for his painstaking and
proposes a phylogeny for the genera and
beautifully illustrated works on bees and for the many
warm interesting discussions in his garden. I wish to
subgenera of anthophorine bees. The tribe
thank Yu. A. Pesenko of the Zoological Institute of
worldwide in distribution,
Anthophorini,
Leningrad for his help in establishing the status of
and considered here to include only two
Anthophorabactriana Popov and A. pubescenslabronigro
genera, Anthophora and Amegilla, is distinLebedev.
Mick C. Day of the British Museum (Natural Hisguished from other Anthophorinae by three
tory) and Astrid Loken of the Zoological Institute of
submarginal cells, the first recurrent vein
Blindern, Norway, brought to my attention many overmeeting the posterior margin of submarginal
looked details concerning the identity of Apis aceraorum
cell 2 near its midpoint, the forewing with
Linneaus.
simple aJar papillae, the second abcissa of
I am grateful to Lorraine Hammer of the Electron
Microscope Facility of the University of Kansas for the
vein M + Cu of the hind wing subequal in
opportunity to use the scanning electron microscope.
length to the cu-v crossvein, and the elonI am indebted to the following collectors and mugate, narrow gonostylus (if present) never
seum curators who have generously lent material: C.J.
flattened and paddle shaped. The Habrovan Achterberg*, Rijksmuseum van Natuurlijke Histopodini consisti.g of Deltoptila, Elaphropoda,
rie, Leiden, The Netherlands; A. V. Antropov, Zoological Museum of Moscow, U.S.S.R.; P. Arnaud and
Habropoda (: EmQhoropsis rr. syn.), HabroW. Pulawski, California Academy of Sciences, San
inphorula and Pachymelus, and formerly
Francisco; R. Ayala,Instituto de Biologîa, México; C'
distinct
as
are
cluded in the Anthophorini,
Baroni-urbani*, Naturhistorisches Museum, Basel,
from the Anthophorini as the Emphorini are
Switzerland; G. E. Bohart and F. D. Parker, Bee
Biology and Systematics Laboratory Utah State Unifrom the Eucerini.
.rersity, Logan; G. R. Brown, Biological and Chemical
Anthophorine bees are mainly xerophilic
ResearchInstitute, Rydalmere, Australia; C. Besuchet,
climates.
and abundant in Mediterranean
Muséum d'Histoire Naturelle, Geneva, Switzerland; J.
Among the North American species over
C. Cardale, Australian National Insect Collection,
90% occur in the dty southwest. In the Old
cslRo, canberra; H. V. Daly, IJniversity of california, Berkeley; C. Darling (presently at the Royal
World, where most anthophorine bees occur,
Ontario Museum, Toronto, Canada) and W' P'
the most speciose area is the Mediterranean
Stephen, Oregon State LJniversity, Corvallis; E. Diller,
Region. Of the L+ subgenera of Anthophora ZoàlogischeStaatssammlurrg,Munich, West Germany;
eight are found in the New World; only one
M. Dylewska, ZoologischesInstitut, Polska Akadernia
Nauk, Krakow; G. C. Eickwort, cornell university,
is restricted to that area. They are scarce in
Ithaca, New York; G. R. Else*, British Museum
tropical forests but are often found in trop(Natural History), London; M. Fischer*, Naturhisical montane environs. Amegilla is strictly
iorisches Museum, Vienna, Austria; S. Frommer and
Old World and flies in the summer, while
Riverside; Y'
J. C. Hall, University of California,
-Hirashima,
Kyushu University, Fukuoka, Japan; the
Anthophora is worldwide in distribution and
late P. D. Hurd, J.. and R. J. McGinley* (McGinley
commonly flies in the sPritg.
was formerly of the Museum of Comparative Zoology,
Cambridge, Massachusetts),National Museum of NatACKNOWLEDGMENTS
ural History Washington, D.C.; A. Jansson, Zoo'
logiska Museum, tlniversity of Helsinki, Finland; J.
I am sincerelygratefulto CharlesD. Michenerof the
University of Kansas,who providedencouragement, Càsevitz Weulersse and the late S. Kelner-Pillault*,
guidance,stimulatingdiscussionand aid throughout Muséum National d'Histoire Naturelle, Paris; F.
Koch*, Museum fur Naturkunde, Humboldt-univerand painstakingly read the manuscript. I am also
gratefulto GeorgeW. Byersof the university of Kansas sitât, Berlin; J. Kopelke, Forschungsinstitut Senckenberg, Frankfurt; w. E. LaBerge, Illinois Natural
for his aid and thoroughreadingof the manuscriptand
Hisiory Survey, Urbana; thç late M. A. Lieftinck*,
for his help in translatingRussian.
formerly of the Rijksmuseum Museum, Leiden, The
Special thanks are extended to George R. Else of the
Netherlands; O. Lomholdt* and B. Petersen, Zoologisk
British Museum (Natural History) for his help during
to
Museum, Copenhagen, Denmark; L. Masner and H'
his
home
for
opening
Museum,
my visit to the British
Goulet, BiosystematicsResearch Institute, Ottawa, Onme and providing enjoyable field excursions. Also my
tario, canada; s.J. Merritt, Texas A&M University,
sincere appreciation to Donald B. Baker for his work in
(Natural
College Station; C. D. Michener and G' W' Byers,
the bee collection of the British Museum
university of Kansas, Lawrence; R. M. Murillo, colespecies
difficult
many
out
sort
History) which helped me
gio de Postgraduados, Chapingo, México; G' M'
as well as discover the identity of some Walker and
Nitftiau, Bishop Museum, Flonolulu, Hawaii; J'
Nurse types. Thanks and appreciation also go to the late
Oehlke, Deutsches Entomologisches Institut, EbersMaurits A. Lieftinck, formerly of the Rijksmuseum van
INTRODUCTION
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SYsrBuATIcs AND PHvrocENY
most all major collections as well as determined and undetermined material in these
collections so that as many sPecies as possible could be placed in their appropriate
groups. I have attempted to make known the
unusual as well as common attributes of
anthophorine species so as to facilitate future
revisionary studies of the various groups at
the specific level.
All structures which could be studied flattened, such as sterna, were placed on slides
in glycerine for illustrative purposes. Larger
structures such as the genital capsule and
mouthparts were dissected and studied in
either alcohol or glycerine. Terminalia and
mouthparts were cleared overnight at room
temperature in about l0% KOH and then
transferred to one or two changes of water
for several hours (sometimes acidulated
water), finally to glycerine for study and
long-term preservation.
Preparation of specimens for scanning
electron microscopy entailed initial relaxing
in a humidifier for one to two days. Parts
were taken from the specimens and ultrasonicated in three warm baths (five minutes
each): the first, water with a detergent; then
70% alcohol; and finally, xylene. The structures, if relatively hairless, were allowed to
air dty; if hairy they were dried with air
blown through a piece of surgical tubing
attached to a micropipette. The specimens
were then mounted with glue (casein) on
stubs coated with Tubekote@ so as to give a
uniform black background. Specimens were
coated with 200 A of gold-palladium and
scanned with 15 Kv (200 spot size), using a
Phillips 501 scanning electron microscope.
Morphological terminology follows for the
most part Michener (1944) and for mouthparts, it certain cases,Winston (1979). New
terms for structures are explained in the
section on CharactersEmployed. The proboscis
is regarded as projecting downward from the
head. Thus the anterior surface of the mentum is the surface that would be called dorsal
if the proboscis were considered to project
forward. To save space the metasomal segments are abbreviated S for sternum and T
for tergum. Therefore sternum 6 is termed
36 and tergum 6 is T6, etc. Almost all
Anthophorini have bidentate mandibles and
OF THE ANTHOPHORINN
BBBS
++l
apically simple labra, thus only exceptions to
these are listed. All caPitalized colors are
from Ridgway (1912) excePt dark browns
and other nonspecific generalrzed terms
(..S., dark brown, ferruginous, amber.) All
characters of surface sculpture are described
using the system of Harris (1979) and were
examined under fiber-optic illumination diffused by frosted drafting film to eliminate
reflections.
Illustrations of terminalia and mouthparts
were drawn from cleared material using a
drawing tube. On shaded drawings the light
was positioned so as to strike the subject
from the upper left. For figures of the uPper
(anterior) views of maxillae, the whole maxilla was slightly flattened under a coverslip.
The orientation of the lacinia and penis are
not standardized and thus their shapes are
not meaningful.
Comments on distributions were prepared
from data taken from specimens and almost
always not from the literature' because there
are many misidentifications in the latter.
The generic and subgeneric descriptions
take into account all characters described in
the CharactersEmployed section and follow the
same lettering system to facilitate comparisons. That is, the characters of the genera
are directly comparable, and the characters
of the subgenera within a genus are comparable, but not the characters of subgenera
of different genera. Major as well as minor
trends of variation are noted in the descriptions. When one or a small group of species
differed from the majority in a character, the
differences and species involved are indicated in parentheses. Structures which are
described without indication of sex pertain
to both sexes.
Genital capsules were drawn with left and
right sides illustrating the dorsal and ventral
surfaces respectively. Dashed lines represent
hidden internal structures, while dotted lines
indicate ridges or denote areas of dark sclerotization. Dot matrices represent the penis
and other membranes of the genital capsule.
Typ. depositories of new species described
in Appendix 1 are indicated within brackets,
accorditg to the following list: [Ansfelden],
Austria, Collection of M. Schwarz; [Berkel"yl, flniversity of California; [Budapest],
ratu"qrorlpll tr'tt'"xorprtr{'Eb'eare
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rurroqdoqluy âr{}roJ sdnorttno er{J '(çgOt)
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zw
SysreprATrcs AND PHvrocENy oF THE ANrsopHoRINn Bnes
found in the male termin alia and mouthparts, therefore requiritg dissections. Anthophorine females have proven difficult to
separate and to place in groups; for the most
part, the subgeneric groups of Anthophorini
are based on characters of males. The sting
++3
apparatus was remarkably consistent among
the taxa and not found useful for placing
females in groups. The mouthparts and genitalia were dissected for almost all described
species as well as many undescribed ones.
The measurements reflect the variation
CUBITAL(1St r.M)
lst TRANVERSE
ALAR PAPILLAE
3rd SUBMARGINAL (2nd Rs)
MARGINAL (2Nd R1)
2nd SUBMARGINAL (lst Rs)
CUBTTAL(2nd r-m)
2nd TRANSVERSE
VEIN(2Nd M.CU)
2nd RECURRENT
lst RECURRENT
VEIN(1st m-cu)
(2nd MEDIAL)
2nd DISCOIDAL
(1st R1)
1st SUBMARGINAL
PRESTIGMA
BASAL VEIN (M)
1st DISCOIDAL (1st MEDIAQ \s
FOREWING
MEDIAL
COSTAL
cu-v
(1st
SUBMEDIAL CUBI
HINDWING
(2Nd CUBITAU
SUBDISCOIDAL
TEGI.JLA
HAMULI
PARAPSIDAL LINE
OCELLUS
SCUTUM
FLAGELLUM
PEDICEL
M"Cu (2nd ABCISSA)
VANNALLOBE
SCUTELLUM
SUBANTENNAL
SUTURE
ANTERIOR
TENTORIALP|Tq
JU94g--
EPISTOMAL
CLYPEUS
COMPOUND
EYE
LABRUM
MALAR
MANDIBLE
PREMENTUM
FOREFEMUR
HIND COXA
COXAL CORIUM
FORETIBIA
HIND TROCHANTER
Sugpt-Eunnt
SIGNUM
/
MESOCOXAL CARINA
PRECOXALSUTURE
HIND
TIBIAL SPURS
MIDTIBIAL SPUR
upinnsus
TOOTH
BASITARSAL
AROLIUM
cLAW-=-
AXILLARSUTURE
PRoPoDEUM
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tlrEual eql 'sewlprru rreql Euole /v\ar^ lerelel ruo{
parns€eru ere/v\ sererrrolateg eql (n) 'p?eH
'suorldrrssep pu? sde>1ur
Pesn
d8olourrurel eql urzldxa ot se^res tsll sFII
'/v\oleq
Pâlsll sralr€r?qr Jo sdnort Pâranel
eql uo poseq sr uorlJâs lxâu âql ur palueserd
DnqdlqruVJo uorlerglss€lr clreuetqns eqJ
'tu"lsuoJ elrnb are sJelJ"J"r{3
eseql fellf s/v\ol{s elqells^? eJe/v\ se snuatqns
qJee Jo serJads dueur ffi Jo pu" serJeds uoru
-rrror /v\al e Jo spnpl^Ipq Jo su"dqlnour Pu?
srl"lrue8 Jo dpnls eql 'ssalaqleuoN 'pelres
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do Àrrsus^rNo. sHJ
wn
I
I
I
?
I
I
SvsrnprATrcs AND PuvrocpNy
oF THE ANruopHoRINe Bres
SCLEROTIZATION
CIRCULAR
445
LATERALPROJECTION
PROJECTION
SUBMEDIAN
VENTRALLOBE
DORSALLOBE
S CLEROTIZATION
LONGITUDINAL
APODEME
SPICULUM
GONOSTYLUS-------------...
PENISVALVEAPEX
PENISVALVE
PENIS
VALVEBRIDGE
PENISVALVEAPODEME
GONOCOXITE
GONOBASE
EYE WIDTH
MALARSPACE
EYE
CLYPEUS
MANDIBLE
PROTUBERANCE
CLYPEAL PROTUBERANCE
PYGIDIAL PLATE
GRADULAR PROCESS
G
MIDDISTITARSALBRUSH
esounf,€r-âsornsnr
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uor'e'I€rrero
pue(suunrdrn'::1Li::Tt'J-iiiliiiii''^ï,î:J'J;Tlt"1H:
'a1evs,o\o1ntoul1q
(o11t1{oyag)'v 'ç1'c'eruloeJeorf,nu sarnlcund uââM}aq puno:s dlerpatu alulnorlcund
orur Surpe:s d1pre1e1Surrnldlncs alelcund Surr*oqs (go ue{orq srrzq qlr,u) pr",1 jo *"1r.i ,ep-a;
n717saa
(o1tt1(ottrfl 'V 'g 'Surrn1d1nrs et€loâreorrrru qlrm snaddlc 'âFureJ nyatur (o.roq{ot1tuocws(y7)
otoq(oq\uy'y 'sn11aro
u"rpeu: o 'aurl leluo:; - g 'ztasorcrtu: rur 'zlesorceru= eru :C pue g ,rr ur* i.
'c prre ur ru.'' yg'
luasa"rde.râurl a["rs 'saaq aur:oqdor{lu' ur punoy dluoururoc surrnldpcg .€ .cld
v
:.,,-,,){*.,",,..'ù--...
',ft'',
#,,X
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'(tOg '3t.{) âtnc" or (t7
osl€ leur ssaco"r(
^ro1npo,rî
V
'8tg)
lunlq'aq uec qclqÀ\ 'ssacord r€lnper8 e olur
l,1pra1z1 pe8uolord sl IJ eletu eqt Jo snlnper8 aql
l.lpuorserro '(lgl 'tlg) areld prprtdd e sTrel pu€
qleâl uerporuqns o^w\lsreeq LL Jo xede aql uâuo
s€ lsnf 'ruasard ueuo sr (32 'tlg) alofl plp\E@
âleru er{J (d) '}uasqe }sotu1e ror{}oue ur pue
'a1nJe
,{1pszq
sr
a8prrq
aq} 'ranervroq 'n"toq{ot1îuV
'aleurEreurâ l.lpseq dlpnsn
Jo snuaEqns euo uI
sl (pz '8tg ur u/v\or{s luoruernsseru s4) anp\rq aalDa
s1ua( eqJ (o) '(tZ 'Etg) xade â^1e^ sruad er{} Jo
eseq âql ]e Eurrnseaur 'qlEuel or{t o1 al8ue aa.r8ap
06 v te ueTet sr qtple\ eqt pue urtreur râuur
eqt Euop pernsearu sr çEua1 eql 'llpsropocrde
xade eq] Eurrvrar4 'âJeJJns l€sJop sll uo ezlâs
â]nurru sJ€eq sder*p pue']sarc
pcrde ue qllm
re^âu 'pepunor Lpua8 s1 xa(o azlao s1ua{ aql (u)
'snldlsouo8 eril
Jo r{tpp\ pue qltual âq} ernsuoru
']sol sr ro ralslle
I rurorl salertsuourap pZ arnErg
e ol paf,nper sr tl serurleruos 'pazrloralrs dpqtrls
Âpo sl.ervtp lsorul€ pue âsolas sderv'p sr qclq^/'
snl(Tsouoï ercSuola dlpnsn '^norreu 'el8urs u l.po
sreeq alrxorouo8 r{reg (-) 'seuo âsounld l.11ensn
NIIs-rTnfl scNsrcs svsNvx .{o ^rrsus^rNn
sHJ
9tn
SysrnvATrcs
AND PHvrocENy
the first recurrent uein of the forewing joins submarginal cell 2 at or slightly distal to the midpoint
on the posterior margin of the cell. It is never
interstitial with the firqt transverse cubital vein
(Fig. 4u).(r) The sting is 2-3 times longer than the
gonostylus.
Bo$t length is measured in dorsal aspect from
oF THE ANrsopHoRrNe
4+7
BBes
the apex of the clypeus to the apex of the metasoma; it is 6-2+ mm.
TRIBE ANTHOPHORINI
PodaliriiLatreille,l902a,p. 377.
DAHLBOM
Anthophor i ni D ahl bom , 1835, p. 5.
Megillina Thomson, 1869, p.7.
A
B
c
D
F
G
H
F I G . 4 . F o r e ( A , C , E , G) and hind wings (8, D, F, H) of anthophorine and habropodine bees. A, B,
plumipes.C, D, Habropoda tarsata. E, F, H. tristissima. G, H, Pachltmelusconspicuus.
Anthophora
+48
THn IJNrvBRsrry or KaNSAS ScrnNcs BurrETrN
Diagnosis.Bees of small to large size, differing
from Habropodini as follows: first recurrent vein
joining posterior margin of submarginal cell 2
near midpoint, never interstitial with vein between submarginal cells 2 and 3 (Fig. 4u); anterior and posterior margins of submarginal cell 3
more nearly equal in length; marginal cell short,
shorter than combined lengths of submarginal
cells along their posterior margins (FiS. 4u);
spatha absent; gonostylus of male never paddle
shaped, one gonostylus per gonocoxite; apex of
57 of male never pointed but entire to emarginate; 56 of male not strongly convex medioapically; scopal hair simple except on posterior
margin of hind tibia usually plumose (Figs. 10ac), sometimessimple (FiS. 10d-e).
Description.a. Body length 6.0-2+.0 mm. b.
Flagellar segment 1 of female equal to combined
lengths of next 2.0-5.5 flagellomeres;of male, to
next 1.0-4.2 fl,agellomeres.
c. Face of male almost
always with white to yellow integumental markings, rarely all black; face of female with or
without white to yellow integumental marking;
mandible almost always with subapical tooth;
clypeus protuberant to relatively flat, profile projecting 0.3-1.2 width of eye; labrum usually
apically simple, margin thickened; maxillary palpus usuully with six segments, sometimes five or
four; labial palpus almost always with four segments, sometimes three or two; mentum usually
without a pair of teeth on anterior basal third,
length 0.5-0.9 length of prementum; paraglossal
length equal to 0.5-1.2 length of stipes; lorum
almost always Y-shaped. d. Malar space of male
and female 1.0- 16.0 times wider than long. e.
Midtibia of male sometimeswith various brushes
of hair (Figs. l2c, 13j, l8k). f. Hind basitarsusof
male sometimes with tooth, ridge, and/or swollen; hind tibia of female with simple scopalhairs,
though usually with a narrow posterior longitudinal zone of plumose hair. g. Arolia present except
in Amegilla.h. Basitibial plate of male present to
absent. i. 56 of male generally flat except for
lateral margin, medioapically emarginate to entire, rarely produced, disc variously modified
with depressions,protuberances, or ridges; 54
and 5 of male usually normal, rarely modified. j.
57 of male with disc apically emarginate to
bluntly produced, laterally simple or with one to
two lobes,length of apodeme0.2+-I.75length of
disc. k. SB of male apically narrow to broad, with
two small submedian lobes whose length is at
most one-fifth length of disc. l. Apex of gonocoxite of male variously modified, simple, bilobed or
trilobed. m. Gonostylus of male elongate and
narrow to greatly reduced to absent, never flattened and leaf-like. n. Penis valve apex is slender
and long to wide and short; sometimes apicully
hooked. o. Penis valve bridge present, spatha
absent (very weakly developed in a few rare
cases). p. T7 of male either with two subapical
teeth or pygidial plate; pygidial plate of female
with margin simple or carinate. q. First recurrent
vein of forewing joining submarginal cell 2 near
midpoint, never meeting frrst r-m. r. Hind wing
with cu-v subequalto secondabscissaof M * Cu;
cu-v slanted from M + Cu basad; vein M about
three times length of second abscissaof M * Cu.
s. Jugal lobe one-third or less length of vannal
lobe. t. Wings essentially bare with well developed alar papillae.
Distribution.The tribe Anthophorini with about
600 species is one of the largest tribes in the
family Anthophoridae, secondperhaps only to the
tribe Eucerini. Members of this tribe are found in
the tropical and temper ate zonesof the world with
particular abundance in xeric areas such as the
southwestern United States, Mediterranean region, northern and eastern Africa and Australia.
Biologlt.These bees are swift flyers. Almost all
of them nest in the ground, preferably in banks; a
few nest in wood [subgenus Clisodonand perhaps
Anthophora (Anthophoroides)signatal. Sometimes
more than one specieswill nest at the same site
and
(Schneider, 1982). I have seen 24. bomboides
abrupta share the same nest site at Lawrence,
Kansas [this was also noted at St. Louis, Missouri
(Rau, 1926) and Baltimore County, Maryland
(Norden and Scarbrough, 1982)l and A. bomboides
and califtrnica at Point Reyes National Seashore
in California. Anthophorine bees commonly reuse the same nest site year after year, unlike many
other beeq. When the nesting substrate is displaced from the original location, bees may be
attracted to it (Michener, 1960), suggesting that
there is some chemical nesting marker that seryes
as a lure. Perhaps this is one reason why more
than one speciescan be found occupying a single
nest site.
Earthen (or saw dust in the case of Chsodon)
cells are lined with wax and provisioned with a
semi-liquid mass of pollen, nectar and Dufour's
gland secretion (Norden et al., 1980). The wax
lining and provisions give off a strong cheesyodor
(Malyshev, 1925) from butyric acid, a product of
the triglycerides of the Dufour's gland and probably salivary or gut enzymes (Norden et al., l9B0)
and is soluble in many solvents (Batra, 1972).
The cell linings are thick and similar to those of
habropodine bees. Compared to those of Habropodini, anthophorine cell linings are not thicker
or do they differ in their separability from the cell
wall; furthermore, there is no difference in burrow orientation, contrary to Marikovskaya
(1e76a).
The mandibular gland secretion may function
as an alarm or aggregation pheromone in certain
speciesof Anthophora,since netted females give off
a mandibular gland secretion attracting other
females by the hundreds (Thorp, 1969; Brooks,
1983; Batra, 1984). Some species have males
which defend territories (Brooks, 1983) and females which vigorously defend their nests against
parasitic bees (Batra, 1978).
Polylecty is generally the rule, as exemplified
by Anthophoraplumipes(cited under ,4. aceraorum)
t
r
I
i
(,
AND PHvrocENY
Svsrer{ATlcs
I
')
I
I
I
I
I
F
oF THE ANrnoPHoRrNn Bnns
whereas speciesof Anthophoraare generally active
in the spring. Many species of Amegilla in the
Mediterranean region are diminutive and superficially resemble species of Anthophorasubgenus
Heliophila.
Taxonomy.Numerous keys have been made to
the adults of anthophorine genera; the most
important covering North America are by
Michener (19++), Mitchell (1962, only Eastern
U.S.), Stephenet al. (1969); India (Batra, 1977),
Europe (Friese, 1897; Schmiedeknecht, 1930),
Egypt (Priesner, 1957), and European lJ.S.S.R.
(Osychnyuk et al., 1978). Keys and descriptions
(Ruszkowski, 1966) and various species of Indian
Anthophorini (Batra, 1967). Only two species are
known to be narrowly oligolectic; they are Anthophora phaceliae n. sp. whose females visit almost
exclusively various species of Phacelia (Hydrophyllaceae) (Brooks, unpublished) and Amegilla
(Glossamegilla) f,orea which in Japan visits only
Impatiens textori (Miyamoto,
1962).
The nesting behavior of Amegilla is as in species
of Anthophora. Some species of the subgenus Asaropoda nesting in flat ground construct turrets over
their nest entrances (Rayment, 1951; Michener,
1965) . Amegilla flies generally in the summer,
GALEA
BLADE
B (
C Q
449
SUBGALEA LACINIA BASILACINIALSCLERITE
MAXILLARYP
STIPITALTHICK
STIPITALSCLERITE
I
l
I
SUSPENSORY
THICKENING
BASISTIPITALPROCESS
PROCESS
DISTISTIPITAL
GALEAL FLAP
GALEAL LUMEN
maxillaandcrosssectionof galea
FIG.5. Maxillae, outerviews. A,B,Anthophora(AnthophorQplumipes,
at midlength. C,D, Anugilla (ArnzgillQqudrifascinta, crosssection of galea at midlength and maxilla. E,
G, Anthophora(Heliophila) bimaculata,maxilla and cross section of galea at midlength. l, H, Anthophora
(Clisodon) terminalis, maxilla and cross section of galea at midlength.
F
I
I
i
,
s
r
I
I
F
I
;l
'eptu sxuronazlc (opfawo.to1) 'V
'O 'epru qofintgsn(oppawo"tn4) 'V 'J.el€ureJ
Dtru
'V'8. 'epureJ
-tofiqoc
(saprc^rotlfutpuV)
oc{tco{
(ororld
-otltuoqdoù 'V'v 'o.rorldotltuv aeuuetuv '
Jo
L'CI.{
'rarlaladaT
-tt 'F.IceA optsoq 'd1sr*o1zsop"U
onqounï
nyouûruta{'puerlg
.to?qatxa oypayy'pue"rlg
s?suatîorapp
'srnoq
pîsnqzaot 'uorarue3
s?npnae 're11e16 ota{tlnuuo
'a1rrpe11 oso1nïuo'1,>lsure8lf-^oztâuzn)tr moqoQo :(pa1zc
-pppppppppppppppppppppppppppppppppppppppppppppppppppppppppppppppppppp
esr/v\râqto sselun o.totldoqrv u\ peqrrrsep (leurtr"ro
erâf\ 11e ila parpnp ueeq lou e^Br{ tnq tsrxe sadft aqt
seseJ rr,al e ur) pedorlsep Jo lsol are saddr agl eJurs snuet
raqlle ur Ltureuao qtr/y\ paceld aq louuec Laqa 'Dnû
-?uv to ototldoqtuv rerltre er€ serJads EurrrrolloJ eql
srcss svJusDNr rNnroHdoHrNV
'srnoq s1ruo{aol1atnloapue srno q o1oî1tçsrun'1arqJrs,Dpqtt
-!un 'lalslC DîDtrunî 'srnoq olpo{s 'srnoq Dluls 'ra8r11
stlruzs oyfayng 'snrcrrqe i Dpnulues sldV 'srnoq oyouozo{n.t
'srnoq
Dîaldat 'srnoq
oyouozo"t@ 'srnoq
efi(dot(d
'srnoq oeoufit@ 'srnoq opasln{ 'srnoq onld 'sedsel zsol
'srnoq osoutïntaootïru 'srnoq nuottrcaw
-id 'lalsr
C oîouJo
'srnoq
otlu@ouopu'aueqlnoqeT
DîDuozotpera'lslJrlD
rpo{nstmu'srnoq
oîDtpnatpoafll'dorc.rno I Dsteut rydy'Aeg
oqoquo{'lalslC Dsl?trc?oppayg'srnoq
oqoùuçsoa.tltc'tezue4
Drytrylq sgv
:(asrm:aqto pelerrpur
sselun ototldotltuV
snua8 âql ur peqrrrsep ÂWut8uo ge) peÂortsâp ro tsol
rerllre are sadd.r ar{r âJurs rurroqdoqtuv
eqt o}ur dture}
-JeJ l{l}r\ paceld aq louuef, sercads turrvrogo; aql
ssrcsds qscv-IdNlf
.(lgO1) uaqdarg
puu onlrroJ pus (teOl) Laprgc141 '(gçOt)
reueqcrtrAldq uelr8 are rurroqdoqluv Is^rel Jo
ueryo eleur ;o talprur 'qlaal qllû\ pesrpour ueryo
ap1zurrr;o
ta1 pug :(nO)luasord Brlorv 'stsoufiotg
ellrarlsT o"aoq{otltuysnuag
' rrll€toru serul]etuosrler{
ol1fauV '
lluasqe ot parnpar dpeert ueryo snl
-dtsouot iprrqr lewq uo qloot rorratu"
elturs e qllÀ\ sderrrp lsorule runluaru
:(pç 'ttg) uorlcelord auo qtlÀ\ ssecord
ptrdrlsls€q :(cO '8tg) luesqs sIIorV
"llll€telll re^au
Drolldoqîuv'
rl"r{ lpadole^ep [e/u\ pue tuaserd sr(ervr
-[E lsorup sn1Â]souo8:(rtZZ'Etg) prlr{l
Fs"q uo qtoot rorreluB r{}l/v\ ro eld
-utrrlsruntueru :(l 'e 'eç 's8l.l) suorlraf
'slloruaplcco
(oa1ay4fl'V 't 'tnaratltoc(opt1do1ryeH)
'V 'd 'sa{twn1d (orotldoqruV)o,roqdotltuv'g 'stu
-n{tquoq (opo{otnsv) 'V 'g. 'oatqola (opûauosso4g)
'V'C'uaqwot
(o1lQawouog)'V'g 'ogotcsort.t
-ponb (opfawV) oppawy 'V 'ssacord IsteleTs
- q
-oxe : d'acsds leuralul : c'etprre [ete1e{soxe
'8uv*erp qoee
Jo lred re/v\ol uI per.IJBu€ eq
plno/v\srnds FIqIJ 'dlanrlcedser'uo11oq pu? lqtg
aql uo er? secsJrns rorrelus Pu? Jelno aq] ]3rll
qcns peluerro ere eelqll 'penluro srnds 1elqll pue
runlnqelerv 'acedslsura]ur e]BrIpuI seerepe]]oçl
'9 'cI{
's/vrer^pcrde 'aerqll
Pqq el€ureJSosecrdv
c
J
l
l
q
-ord
r{11À\
sserord
o/vu
[et1dr1slffiq:("0
'tuasa"rd erlorv
'tt.{) padola^ap
llalvr
rNnroHdoHrNVsHr do vusNsc oJ Àsx
'E.IX oppa opûay41
pu" zlrrrr"Jow sa(n.toat>lsrrolzssnc stsuatlryl'.{llurs xau,s
' srnoq o?ulurtrazas'dlsuret
s'asarrg
n-rtozleuzny zaoueup
o{aqqd'a1crpa11 ocna1ofo1d'ta>l1e6
,arrros'z1ra,rero141
qopdnod'z1rrrtero141suatsalo('[".IteA o\olpo{'rar1a1a
-dtT orntplrut'raTleM suaeelnl' e1crpa11Dttqsn
tnq'arro;
"[11"q ûuimlrutotnal 'srnoq oapopfiay 'elourdg n\ofinaoy
'Â:1snre8
l]-^ozlauzny lulmy'd>1srue8lf -^ozlevzn>{ ruos
-qoryt'a1crps11oeluDt!'1ara1coC,c!"ttory?a!'ta1le6DPld?t
v
g
Nrrs'r1nfl scNsrcs svsNvx do ^rlsus^rNn
sHJ
0ç?
SvsrnuATICS
AND PrrvrocENY
with brushes; pubescence never blue or metallic,
rnetasoma generally with shaggy hair, sometimes
appressed; outer hind tibial spur not to o4y
pârtially isolated from inner tibial spur [(Figs.-6eg) see Commenlsunde r Amegillal; anterior surface
ôf mentum near basal third almost always without
"double," lumen
tooth; galea with a single, not
(Figs. 5b, g, h).
D e s c r i p t i o n .a . B o d y l e n g t h 6 . 0 - 2 4 . 0 m m . b .
Flagellar segment 1 of female equal to combined
lenfths of next 2.2-5.5 flagellomeres; of male to
.t.*t 1.0-+.2 flagellomeres. c. Face of male with
yellow to white integumental marking, rarely all
black, of female all black, sometimes with pale
integumental marking; clypeus protuberant at
leasi .7 width of eye in profile (except Mlstacantho-
OF T H E AN T H OPH O R IN N
BN N S
+5r
phora); labrum of male usually apically simple,
occasionally trilobed (as in Anthomegilla, Daqtmegilla, Petalosternln) weakly in Clisodon and some
Pltganthophoraand Rhinomegilla), labial palpus with
four segments [except /. (Paramegilla) fulaicauda
with three, A. (Heliophila) hololeuca with two];
mentum .5-.8 length of prementum, sometimes
with an anterior tooth on basal third fas in Melea,
Anthophoroides, and A. (Darymegilla) excisa]; paraglossal length equal to .5-.9 length of stipes; galea
with a single lumen (Figs. 5b, S, h). d. Midtarsal
brush of male present [.t in Anthophora s. str.,
Lophanthophora,and Pyganthophora(Figs . 2h, 12c)]
or absent. e. Midbasitarsal brush of male present
fas in Anthophrra s. str. , Lophanthophora,Pyganthophora, and Caranthophora) (Figs . 12c, 13j)l or
FIG. B. Flabella. Scale l i n e s r e p r e s e n t . 0 1 m m . A , B , Anthophora (Anthophora) plumipes, anterior and
p o s t e r i o r v i e w s . C , D , A. (Melea) abrupta apicoanterior and posterior views. E, F, Amegilla (Amegilla)
quadrifasciata, po sterior and anterior views.
.^^âIA
(qlfautV) nllpawV '3
ntoq{oqruV'C'^lrell l€srop 'nTntcso{t^rpnnb
lpsrop'nsua{mban (otoqdoqTunn4s(74)
's^ er^ epls pordu pus
'ngdn^rqn
(rrtrW) otoqdoqruV'9. ' V ' r u n r l o r € - e ' e l n J l q r o - o i g q u r t u I 0 '
Issrop
p u e O ' C ' V u l u r u r I ' l u â s â r d a " r( t p l t l l 8 r r u o ) s a u l la l e r s 'sal€uloJJo ellore pu€ s/v\€lrpulH '6 'CL{
'esso18 stl peqlrcsep (t901) s>1oorg pu€ reuer{f,ItrAI
,(puacar erour pu€ 'ntoqdoqîuv Jo spedqtnou âqt
pelerlsnilr (lOgf) srâpun€S pue (gZOt) €IerroC
eprâurrrlv '(gçO I ' t+Gl) reuer{rltrAl Lq pessnrslp
pu€ palerlsnllr l.1anr1sn€qxa s€^ n"toqdoqruVI€^
-rEI pu€ tlnpe yo l.Soloqdrour âr{J 'r6opq{toyy
'leudorloeu Lp-reurrd sI auo l.1uo pu€ uerer{€s
-qns erv eraue8qns eârql 'punoJ er€ €reua8qns
::H!l#
ii; :* $ï*,,iËJ;X,ï
i,îi{,T#Ji
-uopul
eql 'et1e;]sn11 ]deoxa seere lrqder8oa8
ur s€] tuaserd eletu Jo e1e1d lerprFÂ4 'ru 'eplû\
se 3uo1 se serul] I'g-l'Q çEua1 '(n11faut0uxl/Vur
alnf,e dllecrde) puno"r Lgecrde 'aleur8.reurâ dldaap
o] errlue dlpseq '(n11pautlqtuv u luasqe uorlcaf
-ord pseq) Surlcalo"rd l.yeseq aEplrq e^F^ slued
'ulq}
't '(pZ '8tg) âplm uer{t -ra8uol seurl}
ç'+-6'0
o] >ltlq] pu€ 3uo1 o] uor{s 'l.lpsropocrde pe^\el^
s€ 'xede e^F^ sluâd 'T 'sarceds rlteJ € uI luesqe
,(lqrssod 'uorlcas ssoJJ uI punor ol peuall€U 'eplm
ueql .ra8uol sâurlt 07,-Z '(o^rozldoqrunqdoTLuuur
'Vul sz]
ur su) 3uo1 oTfsadqnot7 (n11fawnn4)
lroqs
'[ 'aeut rvo aleilâur€l
]noqtl^\
peqollrt ro paqo[q 'aldurts
-orq'* uoo"î:f, ?i:!#:K,"::;,::o{i,ffi.,
al€ru yo sn1l.]souo3
:;,
ro r{}lm 'algord uI
uelperu qllm Je^ou 'aleutrec
aFpr-r pulpnlr8uol
l.lensn ur8reur qlTlnt aleureJ Jo a1e1d lerpr8dd
: [ ( r E I ' E ' t t 1 ' s E l . E )n t o q d o q t u n î { 4e r u o s p u e D M d
' '3'a]
-lllaH 'ntoqdotlTuoqdoT
]uasa-rd saurllatuos /I
'8tg) n116autr(soq
aleru Jo ssef,ord relnper8 :l(ltZ
ur os ,(pear8]
Pexag ut8reru lecrde
P€Irâ]u€orluâ^
'lçZ 'pG 's8tg) n11ûautnn4 euros
'
I
I€nlf,e [("lgZ
'n11ûautoqruVl a8prr
pue q1qdo1lag 'n11ûau.toulqy
'qlaal elnJe ol esnlqo
I€urpnlr8uol u€rperu e qll^\
o/vu r{}l^ l.gecrdeolperu peglpour urnrpr8Ld luasqe
e1e1duâr{lnt"ro [(ur->1 'r 'qg3 'tEt.E) ntoqdoqtuotldoT
pu€ o11Êawnn4 ârrros] qtoot l€ralel qtlm sâtul]
-euros pu€ [("-rt 'l 'qgz 'tçI 'rtl '37 't8t.E) Dlfî
-azuztzd lsour pue'ntoqdoqTun7r{4'ototldoqTunqdoT
se I{Jns suol}€f,Slpoul snoIJeA IIrI/v\ aleul Jo elIXoJ
-ouo8
ql8ue1 çL'I-0t'0 '3uo1
xady 'l 'rslp
3
o
3
o
'q '[(t 'pgl '3lg)
o] uor{s e1utuJo ZS Jo âruapodv
'uopos713 'Dll!î
o1y.qdoplagpu€ q1ûautoulqt 'napy1
-awnq?uv ur. srler{ eldturs qll^^ ut8reur rolrâJsod] (,
'n0I 't8t.{) esounld ur8ruur rolrâlsod ldacxa srl€r{
'8tg)
plqll
ledocs alduns qllm eleurâJ Jo 'a8pt.rpqq :(lOZ
pele8uole saurrlaruos snsr€l
leulpnlr8uol ro
qlool padolanep 11e/\.e o] ef,u€roqn]ord padolarrep
dpleart rorrelu€ u€ r{}lm peglpour ro pâpls-lel
-1ered lsoruls 'aldurts âleru Jo snsr€llseq pulH 'E
')Z,I 'qZ 's8tg) ntoqdoqtuorvCpue
']uesqe
ro
[(lg I
'n^rot1doq7un8(4'ntoqdotlTunq(o7''.lls's
o'toqdoqluvur
'J '}uesq€
su] luasard aleuJo r{snrq Iesre}ltslpplntr
Nrrs-r1nfl scNsrcs svsNvx .{o ^rISus^INI'I sHJ
7,çt
SysrpuATrcs
AND PHvlocENrr
The Dufour's gland was frrst illustrated by Trojan
(1930). Brooks (1983) illustrated the terminalia of
the New World species groups. Apomorphic characters for the genus are shown in the cladogram
(Fis.40).
Knv
1.
ro
SuscnNERA on' AivrlroplroRA
Seven exposed metasomal terga; thir-
oF THE ANrHopHoRrNe
2.
Bnes
+53
2
teen antennal segments; males.
Six exposed metasomal terga; twelve
17
antennal segments; females .
Midtarsus with middistitarsal brush
(FiS. 2h), usually with midbasitarsal
brush also (FiS. I2c); apex of S7 later'
ally with two lobes , a ventral and
FIG. 10. A-E, Outer views of hind tibiae of femalesto show vestiture. A-C show band of plumose hairs
along posterior margin oftibia, D and E lack such hairs. Scalelines (on lower right side) represent .1
C, Anthophora(Mysncanihomm. A, Anthophora(Anthophora)plumipes.B, Amegilla(Amegilla)quadrifasciata.
phora)capistrata.D, A. (Heliophila) bimaculata.E, A. (Melea)abrupta.F, Ventral view of T7, A. (Heliophila)
bimaculata.
nllfautoulqA
(qgZ '8tg) palurod
l.11eseq'Euo1 a8prrq e^l€^ srued :(lgZ
'Et.E)xadz Eurqrza"r a8pu uzlpaur
I€uIp
-nlr8uol qllm'paua11zg l.gzrlualosrop
'3tg)
'l gerpaurocrde urnrpr8l,d :(qg
Z
1a1z"red lsotule sut8"rzur l€relel pue
o'.TilJïtr
,T:1:i;î'
Ë;'ï,i:î'J,
'Etg)
nllpautoqruV '
GçZ
aleur8reure l.lpseq '1roqs a8pr-rq e^le^
sruad :(fCZ'Etg) suorlf,eford padolan
-ep dlTeem 'uerpaurqns lIeurs o/vu t{}lm
'xede Surqrea-r
]ou a8prr uerparu I€u
-pntÉuol qllm âlsruJo runrpr8ld :(pCZ
'8tg) pâ^aorreu sprnl]-om] pcrde qllm
alz8uola 'errlue l,gecrde urn11aqzld
'(E
II
'n1t's8tg)
'0r
alzur8-r€ruo l.lpcrdeorperu
sl.ertp ]sorule 9S lsaqol leerde uoqs
'snoreurnu qll1!\
]ou ulnIleqeu pu?
pe/v\orJ?u ]ou spJrrl]-o/v\] pcrde ueq]
'a;r1ue yr ro (rg 'Etg) suorlraford eill
-ra8ug ue] o] âerr{} q}lm l,lerrde 'aplm
puz uor{s urnfieq€U :(qt '8tg) epl^À
se 3uo1 ss pJrr{}-euo }soru le '"reaut1
pu€ uor{s sr(urvtp }sourls aceds r€lEtrAi
(qgZ 'Etg) seqol
0I
porde ilutus snoreurnu q4.tn. urnileqzg
ro (pCZ 'Etg) pe^^orreu sprrr{}-o/vu Ier
-lde qllm alz8uola 'attlua Lllzcrde Iunl
-lequg :QgZ 'qçZ's8t.E) aplm se Euol sz
prrr{}-euo uzr{} erour '3uo1 aoeds ælery
Dalary ' s8ur>1Jzurleluorun8alur .,vto1a(
qll^ âf,€J :QZZ 'Etg) off\l er{} uâe/v\}eq
uorlEJJ€ruep ou 'aseq ue{} Pezl}orelJs
Lpneaq erour ]ou â]Ixof,ouo8 Jo xade
:(IZZ'8tg) aurds prrde lnoqll^\ 'aldurrs
zlqll pug lruur tI uzr{} re}€or8 qr8ual
(nnuto{t1w pu€
saplotoq(oqluV '
ranrzllDa 'V u\ rrtogal,) s8ur>1rztu plueru
-nEalul â]1{llt.r{}lm ereJ :G+Z'Etg) eseq
pus xade ueamtâq uollef,Jzruep pezlu
-€letu lf,urlslp e, qll^À (alnsdzc lelrua8
Jo uollerzdard percelr uI Lpo uaas)
pazrlorâlf,s {pneaq e}Ixorouo8 5o xade
:$+Z's8lg) xadu s,aurdsr€eu peuesul
rnds lelql] e qtl^^ aurds ptrde Surnuq
elql] pulq lurur il ]sour le, r{}3ua1
' '(qZS 'e1t'u8tg) JIBq
5o ped Trlr{}
6
t{}lm gS Jolpu€ (--ç'g I -ç'ù geus
{poq ueq} 'rqol euo q}lm ro l.geratel
aldurrs xadu 3r :(pgZ 'eç2, 's8tg) pado
-lo^ep dplzam. l.1uo serurleruos 'saqo1
Ierâl€l omt rlll^A ZS Jo xadu lalqzrren
snsrelrseq pug l(a8pr"r ro r{}oo} rorr
-eluE lnoqlr/v\ pu? pepls-[elFrsd lsourp
'aldurrs snsr€lrssq pulq ueql
luas
-ard qloo] a18urs€ JI) prSll I€s€q r€eu
t{}oo} Jolrelu€ }noqlrrvt alduls runluanl
(pfZ 'aZZ,'r8tg) aqol l€rarzl
B
euo qllÀ\ ZSJo xadz :(l "1Z 'IZZ,'s8t.{)
a8prr Suorls ro rllool rorrelue r{llm
snsrelrszq pulq :(.:1.ZZ'Elg)prlr{} 1zseq
r33u r{}oo} rorre}u? u3 r{}l^A lun}uetrAl
uoposry
(p 't6z's8tg)
aldurrs l.11ecrdetunlleqzs ltuasqz sdeq
-rad 'alrxocouo8 3o xade r{}I/v\snoJoloJ
-uof, snll.lsouo8 laleluaplr] alqlpuetrAtr
' (tg 'Etg) seqot aTII
L
-;aEug qll^ l,lpcrdz urn11eqeg la]txoc
-ouo8 yo xads rrro{ pa}€f,rzurap l.1rza1o
snq] 'pazqorelrs ,(pqEr1snlr(lsouot !a1d
-urls seurrleuros 'alu]uaplq elqlpuutr I
.L
'9
(i':1
'6
.B
:\.).,,.oqdoqrun3(4
(r-t 'p 'qtI 't8lg) perlue^
pe>Toor{pu€ peue}teg }ou '"reln8uelrJ ro
peqo[q o]rxof,ouo8 ;o xadu :(ngt '3t.{)
sa8prr ]noqlrm 'aldurts gS Jo JI€r{ Iz}srçl
ntoq(oqlunqdoT
' (q 'niI 's8tg) p€rtua^ peToor{
'paua11eg 'aldurts alrxouo8
Jo xadz
:(pt t 'Etg) sa8prr anbrlqo Eur8ranrp
l.lpcrd€ Jo led u qllrvt 95 Jo JIer{ Ie}slCI
' '(ngt 'Etg) peqol
uoqdotltuDrDJ
-l.rt Lplear't ellxof,ouo8 Jo xadz :(pA I
'Etg) rslp
Jo r{t8ua1 sE 3uo1 sE sotrrl}
'1roqs
i'0 uuq] erour ou seuepode
's8tg) suotlcaf
'puorq dlzorde
/S :(pA
-ord a>lll-ra8tg qll^ l.lzcrd€ urnileq€l.t
ntaqdoqruV '
(q-;Zt 't8t.{) paqoilq lldaap o}Ixor
-ouo8 3o xade i(u7t 'Et.{) rslp Jo r{}Euel
su Euol se seurl] t' I ]s€el ]u sâurapode
'a1e8uo1a 'parvto-treul,lprrde
IS :(lZt
'q 'nB 'sFt.E)
urnilaqzl.t
l.lecrdu
eJllue
luasard
ç
a1z1dprpr8dd lluasard a1e1dplqlllsefl
'
]uesqu
+
a1u1dprpr8l.d lluasqe a1e1dIUIqIIIS€g
''@Z '8tg)saqol
9
'EgI 's8tg)
o / v ur { } l ^ Àr c ( a 7 7 ' p g z ' a O I
aldurrs Âgz-rate1/S Jo xade isaqsnrq
'aldurrs
snsJulpltr4l
,\uu
lnoqtlÀ\
'Et.f) auo
g
lesrop
@Z
NrJsrrng scNsrss svsNvx do ^JrsusAINn sHJ
'ç
.,
't
tçt
SysrnuATrcs
1 1 . Length
12.
13.
t+.
AND PHvrocENy
usually 9-24rnrn; 56 simple
without pad of hair; gradular process
of T7 at most obtusely projecting, usually not developed (Figs . L3c, 8, 27 a);
apex and base of disc of 57 wider than
middle, apex laterally angulate cr with
two lobes as in Figures 13e, i, 169,
L2
20d,27d,k.
Length usually 7 .5-13.5mm and/or 56
with thick pad of hair (often hidden
until sternum is relaxed and pulled
apically); gradular process of T7 often
prolonged into tooth (Figs. 30f, g)
when body size is small; apodeme of
57 usually short, apex of disc laterally
simple, not with two lobes, lateral margins of disc parallel or apically converging, or if narrowed medially then
apex of disc laterally simple as in Fig. Heliophila
ures 30e, 30j, 32e, 32i .
Pygidial plate present (Figs. 13c, g),
sometimes lateral tooth below pygidial
plate also present (Figs. 20h, i, k);
gradular process of T7 sometimes
. 13
present .
Pygidial plate and lateral tooth absent,
apex of T7 with two submedian teeth
(Figs . 27 a, f); gradular process of T7
I+
absent
Basitibial plate present (FiS. 29a) and
hind basitarsus simple, nearly parallelsided; pygidial plate with margin carinate and not concave (Fig. 13c); gradular process of T7 sometimes present
Pltganthophora(in part)
(Fig. 13c). .
Basitibial plate absent or if present,
then hind basitarsus with anterior
process dilated, not parallel-sided; pygidial plate with margin rounded, almost never carinate (Figs. 20h, i, k-m)
if carin ate, then lateral margin concave
(FiS. 20n); gradular process of T7
. Paramegilla (in part)
absent
57 apically broad as in Figures 169,
27 d; SB with broad, deep apical emargination (weak in A. calcarata), lateral
margin on apical half concave (Figs.
15
16h , 27b)
57 apically narrow as in Figures 19e,
27k; SB with narrow shallow apical
emargination ,Iateral margin on apical
half angulate or concave (Figs. 19c,
oF THE ANItHopHoRINn
Bees
455
. 16
-.rgin
angulate,
spiculum projectitg somewhat basally
(FiS. 27b); 57 with apex shallowly
emarginate with two weakly developed
apicolateral lobes, apodeme with lateral process (Fig. 27d); apex of gonocoxite somewhat bilobed in pro{ile,
lower lobe weakly developed, not with
carinae, sometimes with a tooth on
lower inner margin of gonocoxite (FiS.
27e); face almost always with yellow
Petalosternon
markings .
SB with apicolateral margin prolonged
and rounded, spiculum not easily visible from ventral aspect but projecting
dorsally at a 90 degree angle (FiS.
16h); 57 with apex distinctly emarginate with two strong apicolateral lobes,
apodeme without lateral process (FiS.
16S); apex of gonocoxite complex in
profile with dorsoventral and lateral
carinae (FiS. 16k) ; face with white
. . Mlttacanthophora
markings
1 6 . T7 with two short submedian teeth,
their bases separated by convex lamellate (FiS. 19d) or angular apical margin; median longitudinal carina often
present; actual apical zorte of T7 as
seen from ventral view, narrow where
bent anteroventrad as in Figure 27 a;
hind basitarsus anteriorly with tooth,
ridge
blunt process or longitudinal
(Figs. 19g, 20g, j, o); apodemes of 57
almost always short (see character j of
Paramegilla), .34- .7 B as long as length
of disc, lateral process absent (Figs. "
19e, 20d); apex of gonocoxite in profile
simple to weakly bilobed (Figs. 19b,
. Paramegilla (in part)
20b). .
T7 with two long submedian teeth,
their bases separated by rounded apical margin; actual apical zone of T7 as
seen from ventral view wide where
bent ventroanteriad (FiS. 27f); hind
basitarsus almost paralled-sided, anteriorly simple; apodemes of 57 long,
1 .00- L .7 5 as long as length of disc,
lateral process present (Fig. 27k); apex
of gonocoxite in profile deeply bilobed
. Dasltmegilla
(Figs.27il..
t 7 . Flabellum narrowly elongate, apically
27h)
1 5 . SB with upi-i.r.t.f
'elrxoJouo8 yo xede pu€ elnsdec
1e1rua3Jo r*âI^ Ierlue^
' ounq.rn(ntoq oqTuo
cn\s(7,,y)otoqdotltuv' g. 'O 'oTonsn{tqonb (n11fawy) o11fau.{V pue nsua{tnban (n^rotldoqtun
d
-nTsdyy)uoq(oqruVJo al€ural 3o srp8rrls 'fl 'V 'ssaco-rdIeurref, Ierelzl - dl 'sn1l.1souo8- 3 'snlzur_ rrr
'I 'CId
'urnlan - n lurru }uâserdar(rplr rq8rr uo) saurl el€)S 'alnsduc
I
1e1rue3pu€ reueelf, l€uueluv
I'
âraruoga8eg il(nnxa
ql8ual ur pnba
I
'V) esournld edors
Jo auoz Ieurpnlr8uol
rorrelsod qlrm pue plro6 plo eq] uro{
uaql qloo] qll^A JI] qloo] rolreluz lno
-qll^ 'aldurrs prlr{} Ieszq uo run}uatrAtr
IZ
' rat{}â8ot ueTe} serouroga8ug g'g-i'Z
JXeu o] {}3ua1 ur lenba I erâuroga8eg
:QZZ 'Etg) qlool rolrelue padola^ap
IIa1!{'e qll^ prlr{l psEq rEeu unluetr l
^to?r,i,L'J'
il:l,iïff
?:':::##"
u,,
' (pCZ
ollpautoq?uv '
'Etg) pe^\orreu sprn{}-o^u
lecrde q4llt
eletuola 'a;rlue Llleerde unileqelg
'Jls 's ntoqdoqruV' ' (pZt '8tg) asournld
srrzr{ pdors qtlm zlql} pulr{ Jo ur8reur
ârrl
pu1pnlr8uol rorralsod :(fZt'8tg)
-ua xade 'sprrq]-olvu prrdz uI pemoJreu
'papls
lalzred ssel ro erolu urnl
'02
'6I
-laq"H :GZl 'Etg) eplm se Euol sz prlr{}
-euo lsour le 'padola^ap llarlt dlare-r
'1-roqs sLz^,ntp lsotule acuds ruletrAi
' â01 a"rn8r.{ul
6I
sz aldurrs srrer{ ledocs q}lm zlql} pulq
;o ur8r€tu leulpnlr8uol rotratsod :(qgZ
'Etg) seqol
lzcrdz llzurs snorerunu
'Etg) pe^\orreu
qllm urnileqzg ro (pçZ
sprlq]-o1ûl lecrdz qll^ alz8uola urnl
- l a q z g : ( r g Z ' q ç e ' s E I . { ), p l ^ s z E u o l s €
prlr{}-euo ueq} erotu 'Euo1 acuds r€letrAl
' râr{}a8ot ueT"l seraruol
0Z
-1a8egç'ç-Z'Z ryau ol {r8ua1 ur pnba 1
ereurolla8eg :(rg '8tg) seqol pcrdz e{ll
-ra8ug 3uo1 lnteJqtlm 'aplm urnlleqzl,{
.BI
BI
' roqle8or ue{"} sereruollaFeg
0'+-Z'I
]xeu o] r{}Eua1 uI lznba I eJOuolla8
-uH :(qgZ '8tg) seqol 11uurs'1roqs sno
-rerunu qlrm ro (pCZ 'eB 's8tg) orllua
lou
Nrrs-rTnfl scNsrcs svsNVN .{o ^Jrsus^rNfL sHJ
9çt
SvsrnuATrcs AND PHvrocENY oF THE ANrHoPHoRrNn
2r.
to next 2.2-5 .5 flagellomeres taken
. 22
together
Hind basitarsus with posterior distal
process acute (Figs. 22h, 23c); 56 with
subapical tooth; posterior margin of
hind tibia with simple hair (Figs. 10e,
22h); body longer than 13 mm
t trtro
Hind basitarsus with posterior distal
process blunt as in Figure L2d; S6
posterior
subapical tooth;
without
margin of hind tibia with plumose hair
(FiS. 10a); body no longer than 13 mm
22.
23.
24.
25.
26.
port.ri.;'- urgir';i n;A*!#rT";{:;
simple hair (Figs. 10d, e, 22h); length
. 23
6 - 1 5m m .
Posterior margin of hind tibia with
plumose hair (Figs. 10a, c, l2d);
. 2+
length rc-2+ mm
Face black, body length l2-I4 mm;
Mediterranean region .
Paramegilla (in part)
Face almost always with yellow to
and
markings
integumental
white
body length 6-15 mm; if face black
then length from 1 1- 13 mm and southern African in distribution .
Heliophila
Clypeus almost always black, rarely
if
with pale integumental marking,
markings are present then not in shape
( (T"
or if in shape of
of inverted
"T"
inverted
, then flagellomere 1
longer than combined lengths of next
25
three flagellomeres
Clypeus with pale integumental mark( (T' )
(FiS.
irg in shape of inverted
1BS); flagellomere 1 equal in length to
next 2.2-3 .0 flagellomeres taken toCaranthophora(in part)
gether .
Metasoma with apical bands of hair
complete and/or surface covered with
26
semi-erect to erect hair. .
Metasoma with apical bands of hair
interrupted medially or surface covered evenly with appressed hair (Figs"
Paramegilla (in part)
2 1c-e) . .
apically
without
Clypeus normal,
hooked or wavy hairs or basal pecten;
flagellomere 1 equal in length to next
2.2-5.5 flagellomeres taken together
457
BeBs
Dasym;g'i uo,t ;eî:#;:l'f,::,'oI;:,:,:;?o:
-
ophora in part) , Paramegilla (ir part),
Petalosternln and Pltganthophora
Clypeus flattened, with hairs apically
hooked or wavy, with a basal pecten
(Figs . 17 a-e); flagellomere 1 equal in
length to next 2.2-3 .4 flagellomeres
:*:i :'::':
"rr,,'
",à,i,eio,o1
i,, p*iy
Subgenws Anthophora
Latreille
s. str.
PodaliriusLatreille , l}02a, p. 430. Name suppressedby'
ICZN Opinion 151 (Hemming, l9++).
AnthophoraLatreille, 1803, p. 167.
Typ. species: Apis pilipes Fabricius, 1775 (-Ap;t
plumipesPallas, 1772, see Comments)
designated type
and Anthophoraput on Official List of Generic Names
by ICZN, Opinion 151 (Hemming, l9++).
Lasius Panzer, 1804, H. 86, T. 16. New Synonym.
Typ. species:LasiussaluiaePanzer, 1804 (: AnthoQhora
crinipesSmith, 1854, p. 324). Monobasic. (See Nomenclatorial H istory).
MegillaFabricius, [1805], p. 328. See Richards (1935)
for comment on the date.
Typ. species:Apis pilipes Fabricius, 1775, designated
by Michener, 1984, p. 139.
Diagnosis. Male without pygidial plate, Ylt]r
two sùbmedian teeth on apex of T7 (FiS. 12k);
midtarsal, midbasitarsal, middistitarsal brushes
of male almost always present (Fig. I2c); 57 on
lateral margin of disc with two well developed
lobes, a dorial and a ventral one, apodeme long,
never with lateral tooth (FiS. 12n); SB wide
spiculum from ventral aspect short and
apically,
'(Fig.
l2î); apex of gonocoxite deeply
biunt
bilobed'(F-igs . l2f , g); flabellum apically entire,
14); basitibial plate of male
not lobeà lfig.
weakly delimited to absent.
Deicription. a. Flagellar segment 1 of female
equal to length of next 3.25-+.04 segments tog.ih.t; of mâle to next 2.00-3.75 segments. P
Fu.. of male with yellow markings, of female
black (except labrum of A. melanognathahas lightyellow'spot); mandible of male with palg spot. c.
Malar spuce of female about five times wider than
long, of male about six times wider than Jo-"Si
flabëilum apically simple, without deeply lobed
anteriot *atgin (Fig. 12j); mentum simple without anterior iooth basally, mentum about .6-.7 as
long as prementum; paraglossa about .6-.8 as
lonf as itipes.
-(Fig d. Midtarsal brush of male well
. l2c) (except absent in lanata,
devéloped
weakly devèloped in fuluitarses and subterranga)..e.
Midbasitarsal brush of male well developed (Fig.
I2c) (except absent in lanata), f . Middistitarsal
brush' of male well developed (Fig. 12c) (except
weakly developed in lanata). g. Hind basitarsus of
. l2a, b). h. BasitiEi4 plate of
male simple Gtigs
'defined
to absent. i. 56 of male
male podrly
apicomediaily emarginate [emargination slight to
',\ âr^
'lS p,re ,t,râr^oprs 'rslp '/S '1q '141'rnar,r
'e1eur;o
'1 'itlara ps.rop '(pallruo
lertuo^
["Jlue^
99
leq) apu Jo l,J Jo xady 'y '^el^ Jor.râlrre 'urnleqelg 'f 'srr,reraâprs pu" Ierluâ^osrop 'alnsdeo
'g 'g 'aprue; go ^4,âr^eprs 'peeq pue ,v\el^ Jotno 'ta1 pur11 'g 'q '(rvrer,rrelno) epur;o Ealpnu
I"rIu3C
pue purH 'p 'g '(aler,r"re1no)8a1pulH 'V 'gggo xede;o sserord uerpeuens:dus 'qsnrq lusrelprur
'BS;o xede yo ssacord
=qtur 3qsruq lesr"lrlsrppru:qpur 'qsnrq lusrelrs"qprrll-qq*
lerer€l=dl
:sa(ryn1((y) 'ts 'N-I 'g-g iapur 'm1îouâouo1rw
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7,9+
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AND PHvrocENy
oF THE ANrHopHoRINs
BnBs
+63
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,91
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AND PHvrocENY
t
!t
I
I
Etymologlt. Lopho.s(Gr.), ridge or crest, indicating the apically diverging ridges on the distal half
of 56 of the male, plus Anthophora.
Dispar Species Group
b. Labrum of male apically with three projections, median projection emarginate, suggesting
that of many Heliophila rnales (Fig. 301). e. Midbasitarsal brush of male well developed (FiS.
15a), occupying one-third to one-half length of
brush of male well
basitarsus. f. Middistitarsal
developed (Fig. 15a). g. Hind basitarsus of male
somewhat round in cross section and dilated. i.
56 of male apically emarginate medially with a
large, deep depression on each side of disc (Fig.
14f); 56 of female normal without median subj. Disc of 57 of male bry_allyapical projection.
-of
tooth-like protuberances. k. SB of
with pàir
male itrongly angulatç apicolaterally. m. Gonostylus of male round in cross section, about 3.5
times as long as wide. n. Apex of penis valve
about 1 .6 times as long as wide viewed apicodorsally. o. Penis valve bridge basally emarginate,
!.7'times as long as wide. p. Pygidial plate of
male not distinct, represented as a median shelflike projection, apically wide, about .4 times the
distance between the apices of gradular processes.
q. First recurrent vein joining submarginal cell 2
distad of midpoint. r. Sting in repose attaining
apex of gonostylus, about twice as long as gonostylus.
'
Included Speciesand Biologlt Included here is only
species , A. (Lophanthophora)
one Meditérranean
dispar Lepeletier, the biology of which in Egypt
*as desciibed by Ibrahim (1976, as A. speciosa
Friese).
oF THE ANrHoPHoRrNn
Bnns
+65
Porterae Species Group
with a
of male medioapically
b. Labrum
apical margin
projection,
weakly emarginate
sometimes flexed outwardly, greatly produced. e.
Midbasitarsal brush of male absent (weakly present in A. atricilla), though ursina and dammersi
derived, dense
different,
have an apparently
group of flattened, orange hairs in the same
position as a basitarsal brush. f. Middistitarsal
brush of male weakly to well developed. g. Hind
basitarsus of male flattened. i. 56 of male apically
entire (sometimes emarginate as in ffinis, agama,
biciliata,, disparilis, neglecta, robusta, turcomanica, and
ursina), with a shallow depression on each side
(FiS. 14d) (depression absent in disparilis); 56 of
female normal, without median subapical projection. j. Disc of 57 of male with a basal pair of dark
scleroti zed areas and/or tooth-like protuberances
(Fig. 14e). k. SB of male apicolaterally rounded
(FiS. 1aS). m. Gonostylus of male flattened to
round in cross section, short to long, 2.5-8.5
times as long as wide (Figs . l4a, b) (long in
atricilla, and turcomanica, 20 times longer than
wide). n. Penis valve apex about as long as wide
as viewed apicodorsally. o. Penis valve bridge
basally emarginate, | .6-2.3 times as long as wide,
short in disparilis, 1.0). p. Pygidial plate of male
distinct, apically wide, about 0.3 the distance
between apices of gradular processes (Fig. 15c).
q. First recurrent vein joining submarginal cell 2
basal to or distal to midpoint. r. Sting in repose
attaining or slightly surpassing apex of gonostylus, 2-3 times as long as gonostylus.
Comments. A. disparilis Friese is a strange species in that the male has an all black labrum,
greatly flattened hind basitarsus with a tooth and
process, mbb:midbasitarsal
FIG. 15. Anthophora subgenus Lophanthophora,males. gp:gradular
brush, mdb:middistitarsal brush, mtb=midtarsal brush, pp=pygidial plate. A, Midleg of A. (L.)
dispar, outer view. B, C, A. (L.) porterae,midleg, outer view and apex of T7 (hair is omitted), dorsal
view. D, A (L.) hispanica,midleg, outer view.
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osuâp e qlyr\ 'errlua ut8renr pcrde 'rleq {cBIq
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B9r
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AND PHvI-ocENY O F T H E A N T H O P H O R I N N
unmodified face and the male has an unmodified
55, 56 and genitalia. A. urbanais at least bivoltine
(Mayer andJohansen, 1976), flying from the end
of March to mid November.
BNNS
+69
Included Species. A. (Mystacanthophora) urbana
"primitive"
species,
Cresson. Even though it is a
it has the largest range of any North American
Mltstacanthophora- Texas and Nebraska (exclud-
Scale lines (on right side) represent .1 mm;
FIG. 17. Anthophorasubgenus Mystacanthophora.
fs=flattenedorroundstouiseta.hs=apicallyhookedsetaofpecten.A,B,Anthophoracapistratafemale,
and
face and closeup of base of clypeus. C, A. montana,baseof clypeusof female. D,E, A. arequipensis
A. hitaris, uppei part of clypeus of females. F, A. walshii, ventral view of male metasoma showing
pattern of pubescenceof 55-56.
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opfauqog
'?Bg 'd 'g+61 'esnoqpu€S
]ou âl€ru Jo rlsnrq lesre]rs€q pu5{ râuul : (tg I
'Elg) e^ecuoc dlpeorq âceJrns pr€ûuno 'peue11eg
Âq" prtrutrsap 'tLL1 'se11e6soa.nstdv :sercedsadda
'dd '
'asarrg o11ûauolo4 dlleer8 snsrelrseq lqtoot IBsBq ]noqlr^\ ro r{}lin\
'gZ pu€
BI
L6BI
aletu Jo snsrelrseq pulH 't '(lSt 'ttg) pedole,rap
aserrf, oq4gfiau,rrrnd snueEqng
IIâÂ\ alsru Jo seqsnrq l€srelrlslpplul pu? I€srslls€q
'?ulqc
-plni 'J-e ']uâsq€ olstu Jo r{snrq lesre}PltrAl 'p
'sedrls se Euol se
L' lnoqe essoltered iurnluauard
se Euol se 9' ]noqe runluotu 'pr5{l l€seq uo
qloot rolJslu€ lnoqltrvr 'aldurÏs runluetu lseqol OTII
-ra8ug lecrde lere^âs qllÀ\ run[âqeg l8uol se epl/v\
se serurl L'0I-0'g âlelrr Jo 'Euo1 sB epl/v\ s€ serull
'f, 'Qulprrt 'V ul
L'TI-ç'ç oletueJ go eceds r€lenl
luasqe) Treru eled qllrnt. urnrqel :Qu1paquI ]uesqe
'ap! ur umorq) (Egl 'Elg) >[reru ellq/v\ pedeqs-a
peue^q qllû\ aleureJ 3o sneddlc i(all,{^ elqlpu€rrr
qllm lulpaq ù Icelq âlqrpueur 's8urlreur a]lqrvr
er€d 'q 's]ueur8as
r{}lÀ\ el€ur
]xeu o]
;V1I
Jo
eleurrr5o 'sluautas 0'g-7,'f, ]xau 3o qfual ol lenbe
'uot4{ttcsaq
eleureJ Jo y luaur8as re11a8e1g 'e
'peleredas dleprr*
'(fgf 'Elg)
qloâl
lecrde
'luasqe el?ur
'uerpaurqns o/vu r{llm
Jo e1e1d
LL
prpr8dd :(rgt 'Etg) aqol papunor Ierelel pu€ eqol
preleloctde
u€rpatuqns qlyv\ urtreur
alep8ue
'Etd
'poqs pue psorq aleru
uor{s
Jo BS :(pgl
euepode 'Euo1 s3 epr/v\ s€ ]noq? JSIP qllÀ\ elsru
'8tg) pel€llp eptu
Jo rnrueJ Puu elql]
Jo IS :(tgt
esoles
l,]tnecuoc
pulq :(qgt'ttg)
Ieurpnlrtuol
(1es-reds 'durqs qllm snsr"tls€q pull{ Jo ereJrns
'tlg)
pedole,r
râuur Jo ur8reu rotralsod :(lgt
-ap ilâ/v\ sâqsnrq Issr€]IlslPplur pue l€sr?]Iseqplul
'V
'luasqu qsnrq
lesrslplur qllû\ olsru :Qu1pat1
eseurr{O aqr rdacxe) (Egl 'tlg) Treru aled pedeqs
-I parre^ur qll^À eleurâJ yo snaddt3 'stsouflotq
urols"e pu€ elpq
uJâr{tJou o} seeJ€ uuadorng
urelsee puB ueeusrrelrpetr tr aql ur punoJ ere
'lcunlsrp ssel ro ârour ar€ sarcads xrs ar{J
dtql
'ds 'u oxocout{s
'(snrcrrqe.{)
'uaTJ
pue
suacsaqn(
'rer1e1a
-Iy lraytnauy 'ureqtuq
aîo! 'r.{JIV
lulpazl
-dtT umo{np (ntoqdoqTun"rop) 'V 'saleadgpaprytul
.stsouîotq eql
ur ue^r8 sralceJer{J âleru Jâr{lo dq pue 'snsrelprur
pu€ erqrlpru eq] uo rleq Jo ,(sogsnrq,, dq 'qloo]
lecrde ue pue Fseq € eJeJrns rorJâ]ue slr uo sr€eq
lsrl] snsr€lrs?q peue]leu pu€ elqlt pue rnuâJ
puFI pelelp e Âq,peleredas dppeer er€ seletu âr{I
'srrerl esorunld
Jo pueq € qlrû\ elql] pqq eqr Jo
ur8reur rorrolsod aqt turzreq dq poteredas oq u€c
selsrual slr qcrq/v\ ruoJJ o71.qdo11ag
o} pe}?la"r dlasolc
otoq(oqruorv7 'sruaaal7
]soru sr dpueredde
'o"toq{otltuV
snld 'aleura; erll Jo ereJ erll uo sTreru epd eq]
o] EuureJer xgard e 'etel'('rC)
orDJ '(lo1ow(1E
'turu
's 'snldlsouo8
B'Ll-ç'B qlEual
;o qfual serurl
'QIo! ur Eurssedrns dlrqErls) snldrsouo8
ç'Z-0'Z
3o
xade tururelte elrnb 1ou dlensn asodar ur turlg 'r
'lurodplu
o1
lelslp Z IIeo leur8reuqns Eururolurâ^
'b 'luasqe
luerrnror lsrrd
LLJo ssâcord relnper8
:(fg t 'Etg) sas€q relno rrâr{} uee^{.}oq acue}srp
sB Euol se t'-Z' qr8ual rreql 'qlaal uerpauqns
paleredas dlapunr Z r{tyit dgerpaurocrde aleur Jo
'd 'aprrvtse Euol
4; lluesqe âleurgo areld lerpr8d4
se ç'I-6'9 'dgewq eprû\ pue uorls 'aleur8reruâ ]ou
'eldurrs se^p^ sruodyo a8p1rfl 'o '(eBt 'Etd dles
InB' "::::;
;::- ;y" ::i#;-;;"
eddl
'(eruro;
-lleC elug ol erqrunloC qsrlrrg urorllnos) rseo3
er{} o} tsâr* (euroqelTo pue sesuey Erl
rgrred
Nrrs'rTnfl scNsrcs svsNvx do ÀJrsusArNI) sHJ
0L+
SysrnuATrcs
AND PuvrocENy
oF THE ANrHopHoRINn
Bnns
+71
dufouii, all male except G. A, B, Genital capsule, dorsoventral and
FIG. 18. Anthophora(Caranthophora)
D,
E, 57, ventral and side views of disc. F, 56, ventral view. G, Head'
ventral
view
sideviews. C, SB,
H, Hind basitarsus, inner surface. I, Hind leg, outer surface. J, Apex of T7, dorsal view. K, Midleg,
outer view
e-re(p-y 'I 'H) r,J;o sacrdy.,,^âr^
relno,nw.rofu,uac
(ù .v to3elpurg .3tflff"ini'ii#:;;
'tnpra (d 'v 'olq"p
'y 'o71oa(ù 'v
'N-x 't*er^ lolno ta{qmt7 (
.y
Q) V pue
fu)
l" 41 ;o xady
4)'
'f 'r*arapsrop'o{rwwos(4)'vJoLrpxedy'1
'ââr^l€srop'1a3oxedy,11 .meïa.re1nà-,8e1
;oEe1pur11
'c ' ^âr^ JorJelus 'urn11aqu1g'g 'srr,rerrr
'gs pue
'gs 'g-c 's^ âr^ eprs pu?
PulH
["rlue^
/s
leJluâ^osrop
'elnsder
'g, 'V 'sllosvq (ù
.V ,11-y .sapur ,opûawo.to4 snuatqns ototldotltuy
.02 .CIù
l€trueC
,g ,V .e["u ,soan(optâauoto4)ototlilotltuV.6I .CId
'BS 'C 'si\Àer^âprs pue
l"rtuâ^osrop epsdzo pfpeC
âpl^À ss gg'-çt' r{loo1 pedolarrep IIe/û B ot drunq
JorJolue ue qll^À sdurvrlelsotu1e e1eru Jo snsJelrs€q
pulH 'E 'luasqe e1etu Jo seqsnrq qt1,v\ Er1plnt
'sadrls se tuol se
J-p
B' tnoq€ essol8ered lurnl
-ueruard se Euol s€ 9'-g' lnoqe runlueru 'qlool
JorJelue lnoqluvr 'aldurrs pJrql l€seq Jeeu urnlueru
lsaqol a>p1-ra8ug pcrde lerâ^os rlllÀ\ urnfieq€H
l8uol se apy/t sB serurl 0't I - L' ç eleur Jo !Euo1
se epuv\ s€ serurl O'tI-ç'ç elsureJ ;o aceds rel?tr tr
'r 'selqrpu€ru pus runrq€l snadÂ1c uo sTr€ur
alud
ur8reur pcrde '(suo,ra(ns 'V ur ârrlue ur8reur)
dgerpatu eleur8rerue dyvrorreu ol dpeorq âFtu
'r.'(oaupflntta{'Vvl padolarap Ll4eervr
Jo gSyo xady
'suota(ns 'V 'oau.tnqa 'V 'untoau1oq 'V u\
luasard)
luesqe dlpnsn eletu Jo a1u1d 1erqrlrseg 'q '(lOZ
'Etg) snoqpq 'pare11p ueryo apru
Jo rnrueJ pue
Blqll pqq :(fOZ 'Etd pale8uole seurleruos a1etuJo
snsrel pu5{ i(oautïnua{ 'V pue 'suacsaqnn 'v 'u,rntoau
-lnq 'V ur aldurrs) eEprr pqpnlrtuol
rorrelue
us rllr/v\ ro (o 'f 'tgz '86I 'sttg) Euol se Âqeseq
'urn11eqe1s
'g 'arera
,r, ," . ',,!"rîË1Ëî.T:'i:-iliËt::.ï',tfîT;?"iiî
rorrârrr€
rerluâ^
Nrrs-r1nfl ssNsrcs svsNvx do ^rISusAINn sHI
zL+
AND PHyrocENy
SysrBuATrcs
:
N
A
oF THE ANrHopHoRINn
BeBs
+73
s}l 'Euo1qo,,
Eurl're,r sâqol pcrduqns
ruo{
slsoutnlp
se runurels slt{} sprocar aqs o176aIaDI0SJo
rorl ur q8noçle 66olrurels rIlB Jo xode pue alruJels
r{t4 Jo seqol lecrdeqns I€rete1 pare8uole dl8uor}s,,
eql sr relrereqc turqsrnturlsrp reqloue ler{} pe}e}s
osp (OgOI ) edelsnoTlretrN 'exel o/v\l req uââ/v\]
-oq delre,ro dleprm r{rlr{ù\ sqrtuel 3o runnurluoo
e sr aral{} esnereq (edelsnoTlretrAI nsues) on!8
-araorzd ruor; n1lfauolos e}eredes 1ou seop ereruol
-1a8eg lsrg er{} Jo qfuel eqJ '(sa{g4cotâ')sluau8es
t'g or (soa4) sluoutas g'Z ro (n370) T;Z eleureJ
âql w '(onp1a) 6'2, ot (soa4) stuaur8as ç' I sI el€ru
aql uI atuer eql (edelsrtoTrr€tr\I nsues) o117îatu
-o.tzd uI 'Qmouotqv prJe lanrnzasouo)stuatutes 0'?
o1 (n1onçtasap)sluaurEas L'7, lxau aq] o1 ql8uel ul
pnba sr ereruogaEeg âl€ureJ ]srg arl] '(wn.towsouo)
ol
(o1ocry"rasaù stuaurtas
stuatutas
ç' I
0't
lxeu âq] ol r{}tue1 ur lenba sl arauolla8eg aptu
'o1713auo70s
q paceld edelsnoTlretr\tr r€ql
tsrg egl
sarcads eql q 'exet eql eleredes o1 (araurollaEeg
]srg: ) ,,luourtâs r"letng puores,, âïl 5o çtue1
aql pâsn ede>1s^oTrretrAI'ade âql Jo acuuraqntord
eq] 9' sI ler{l 'algord uI aJuereqntord leeddlr arues
'ù qtQ
ar{} e^Bq(Wstomaqa() ollfauolos pue (soa4
-azaDrDd
Jo serreds ad& eql l.lapvvr serre^ snadÂ1c
eq] Jo uorlcalord eqr 'dlrepurrg 'ureq] pâuolslÀua
eqs sB exv,t t{}oq Jo satcads eqr tuoure dpear8
serre^ aeue8 erll Jo qlpyv\ eql !o716auo.tzd tuor;
o1lfauolog turleredes uI ânle^ ou Jo ere sTeeI{J
âpyv\ oqJ 'saletu Jo s1e1d lerprtdd eq] Jo sepls
eql Euole qteâl Jo acuesard eq] puz BS pue lS Jo
saqol lere]Blocrde paleEuola dltuor]s eql '1 arenr
-ogateg pale8uole'snaddlc
Eurlceford l,1Euor1s
'sleaqc oplÀ\ er{r uodn peseq opûawolo4 luor;
(OgOt) ede>lsnoTIrBIAI
paleredas
oypauopg
prlq] eq] ol tll8ual ur pnbe lnoqe sI
sndled drugrxeur a{} Jo }uetutas puocas eq} r{rlqrnr
ur pue elql] pulq aql Jo ur8retu rorralsod aqr
Euop rlsr{ esounld >1ce1se1utuoJ ern qrrqa ul DII!8
-nanrLd u/v\ou{ dpo eql sePnlcw (o\oan\ 'o\otoua{
'oso4tquo 'V ' 't'a) dnorE sercads ,(qtrorrrelou
'(p 'cIZ's8tg) etral eql uo spu€q rleq pcrde
V
l,gerpaur a^BrI sarcads Luu141 'GtZ
pardnrretq
'Stg) sel€ru
Jo ?S uo rl€t{ Jo 6(u/v\orc,, retlnced
'sllosaq
uI dnort rer{}ouv
e s€q €lsv
IBrlueC
ur s€ (qf Z 'ttg) /v\oleq pue eloqe tur8renlp sllqro
'ttg)
reuur eql wq dnort u€f,r{V euo }nq (B I Z
lalpred ssel ro ârotu sqqro reuur eq] e^Bg selelrr
o11ûauo"rn4 due141 '(OgOl) e(els,roTlrel4tr dq se/v\
sernlJnJls Jo uorleJlsn11r Pue uols
optfiaun.tD1
Jo
-snf,srp ]saq aqJ 'orel{ pessnrsrp }ou o11ûawo.to4
'o11paw
urqlr/vr sdnor8 satceds duenr ere erer{I
-DrDd ueql ellsurtuJel olelu luorarurP pue runlueru
eI{} Jo prlq} Isseq eI{} reau q}oo} rolreluv vu ar\sq
CIalarypue sapzlrollfuq\uV 'rleq ]cere-Itues ol lrere
qly/' dllensn pue ('**
ZI-0I) 11eursst saptotoqd
-owuv seerorl/v\ rlsrl passardde qtrr* Pu? ('**
?I
'S 'If urelso/v\r{}nos eqt ur rncJo
lnoqe) atrel are
'uerdorqlg
t€r{} q1ûauon4 Jo serceds o/v\} ar{J
pue Jlloreeled dlalalduroc lsorul€ sr n11ûawn"to4
pue rrlrreeN sr s?pt0.t0q{oq7uVecurs uorlnqrr}slp
dq Lpsea lsoru saprcnq(oqruV tuor; pale.redas sr
olltflawn.tzd 'poqolrrl tou 'lurod eldurs e ol turuoc
'aqol pepunor a8rul B rllr/v\
tou
BS Jo eseq eql pue
'a1e1nEue ro papunor
lS Jo urEreur 1erât€locrde
eqt Eurneq ,(q paleredas ere se1etu aql '('rtt
'n1oan1 'V
rdecxe) rl€q esotunld Jo pueq r{tl/u\
ur8;uur Jorrelsod aql pu€ ssacord lelslp rorrelsod
tunlq e qtl^À elql] pug aql Eurneq dq epurâJ
eql ur Dalary uro{ sreglp }I 'sapto^t0q{oqruVpu€
Dalaw ot pe]€1ar /(1esol3 sI n11ûauon4 'sruawwTc
'(loyasaa sef orygïsaa pue
e(els,roTlr€tr\tr
fiotluaqcstpa{ s€] rcryaqcpa{'016I)
pue (utntoauloq 'V ' e06 I ) uoued Lq pepodar
ueeq e^eq o11ûauotvd Jo sarceds u€rserng eerr{}
yo ser8ololq Eurlsau eqJ 'S '{l ura}se/v\r{}nos eq}
u1 dery dlrea ol qf,r€tr\tr 'pFo114 pto eql w lsn8ny
ol qrJ?tr trruoJJ t(g olpaaza2rDdJo sercadg 'Gop\g
'ruru
's 'snlLlsouo8
+Z-ç'g {}Euo1
3o çtuel saurrt L'Z-B' 1 Eurls l(çtua1 reldlsouot
t+' se reJ se Eurssedrns) sn1(lsouo8 ;o xede Eur
-ssedrns o] Eurure]]e esoder ur Eurlg '-r 'lurodpgr
Eururol ure^ luârrnoer
o] I€lsrp Z IIâr purtreuqns
'b '(p 'rIZ 'sEt.E) dlerpeur peldnrrelq rlerl
]srld
Jo spueq lecrde ro eJuef,soqnd lJero-rruos esuâp
ro (ttZ 'Elg) posserdde qll,v\ uego etuos"leru
:(pOl
lluasqe âFru Jo LL Jo ssecord repper8
'tlg) €urrer
leurpnlr8uol uerporu qt1,v\ sarurteruos
'(tOZ 'Etg) qlâel u€rpatuqns EulluroJ ateut8retue
dlapr*r ol dyvtorreu xade qll/vr dpo sorrrrleruos
'[(rOZ 'Etg) n1ntoua{ 'V pue ranrlaulnq 'y
rdacxe]
surtreur eleurrec lnoqlrrvr a1e1d '(qOe 'tlg) ses€q
ralno rrâr{} ueo/vqeq aJu€}slp se Euol se 0Z'-90'
qlool Iera]BI '1eseq B r{}r/v\ sdervrp }sotul€ a1e1d1er
-prtdd padolanep 11eff\q]lrv\ (11ensn el€tr\tr 'd '(nOZ
'e6l 'sEt.E) aplÀ\ se Euol sB serult g'Z-V'l
çEue1
'(suo.ra{ns 'V pve 'oaut3n.r.tr{'V 'o\otoura{'y rdacxa)
aleurErsrue dlpseq âAle^ sruad yo a8pufl 'o (1ps
-JopoJrde parvter^ s€ epl^/r se Euol sB setull +'7,-0'l
e^le^ sruad ;o xady 'u '(sa{qtcnÊ 'V uI durnq
02,'q'n6T
I l e u r s o ] p e r n p e r s n l d l s o u o t ) ( q ' n'pauelleg
's8lg) epp\ se tuol ss serull g'B-0'7
dlln
-nsn e1etu 5o snldlsouoC 'ru '(qgt 'ttg) peqollr]
o] peqo[q d11eem algord ur 'elduts dlanr]e1ar
s^(ervr
eleru Jo âlrxoJouo8 Jo xady'l'luasard
-le ]soutup rrortezrlorâlrs l€ulpntr8uol :(tOZ 'c6I
's8tg)
lqtrerls ssal ro erotu ur8reur leretpl !e1e1
-ntuu pue dlerpaur aleur8rerue l.ldaep ol dpqErls
el€ru Jo BS 3o xady '>1 '(saun1 L'1-Z'I 'onïtlncou
'sa(qtcn.tE'V 'suoutaql 'V ur tuol) (pOZ 'a6I
'V
pue
'goqs eruep
'stt.{) rslp se tuol se sâtult
BL'-+g'
-ode lluasa-rd sâtulleruos eeJB pâzlloJelJs dplearvr
qtnoqt eruerâqnlord lerel€l lnogll/v\ cslp Jo oseq
isaqol t-Z qllÀ\ dgecrde 'pscnpord ro arllue 'eleu
-rEreura dllerpetuoctde eleur Jo IS 'l 'GlZ 'Etg)
JI€II esuop Jo , (u/v\oJJ,, uelpoul e qllÀ\ selulleluos
aletuJo ?S :(tOZ 'q6I 'ttt.E) uolsserdtp asre^suerl
o1 >l€o1v\ r{tyv\ pexâU dgerluen
o^€ruoc tuorts
Nrrs-rTng scNsrcs svsNvx do ^rrslrs^rNo. sHI
+Lt
SysrpvrATrcs AND PHylocENy
rounded, merging with apex, to strongly laterally
expanded and tapered in the following series of
species.
Subapical lobes of Bth abdominal
sternite not developed, its apex varying from
medium width to narrow and strongly elongated
The
in the following sequence of species.
oF THE ANrHopHoRINn
BnBs
+75
species of Paramegilla (sensu Marikovskaya) show
the same variation in the apices of 57 and SB. The
last character of the male, namely the presence of
teeth on the pygidium, is common in both her
Paramegilla and Solamegilla, thus not of value in
separating them. For these reasons, I have syn-
FIG. 21. Anthophora subgenus Paramegilla. A, A. gracilipes, male, frontal view of head. B, A. semirufa,
male, frontal view of head. C, A. ireos, male, dorsal view of metasoma. D, A. olgae, female, dorsal view
of me tasoma. E, A. martensi, female, dorsal view of metasoma. F, A. deserticola,male , lateroventral view
"crown"
of hair on 54, and numbers sterna'
of metasoma. arrow indicates midventral
o]€r€dâs ol alelu eq] Jo snsrel elpplul or{} uo
rleq tuol uâUo 'asuep yo e8ur rJ e pue snsrzlrseq
pulq eql uo qtoot e Jo ecuesqe ro acuaserd
eql se qJns srelf,€J€r{c pa}lurll-xes pu€ 'eceds
r€letu er{} Jo }ueurdole^ep Jo ]unoru€ er{} 'eJeur
-ogeteg lsrg or1]
Jo rptuel âql uo l'pretu pârler
eq 'ssalanlel ss/v\ g IIâJ purtreuqns
Jo r{}peerq
âql teql Euuntouy 'oapw ol osle lnq (llerelco3)
ranrollna otoqdotltuv ol dpo ]ou elqzcrldde sernt
sapzlrll!{otltuV }eqr pe^erleq ( I ç6I ) e)plreqtull
'(gtO1) asnoqpu€S dq pasodord
s?/v\ Dqaw errr€u ]uaruâcelder eqt eJuer{ 'snua8
ueretdopldeT
e roJ uorlspuerue u€ s€ (f tg r )
zrssety dq dlsnornard pesn uoeq p€r{ Dssauaaî!ruV
arusu eql 'alBru ar{} roJ snsr"trssq pqq peq}oo}
aql pue 'epura; eql roJ re]f,€reqe l'a>1pdrcurrd aqt
se g IIoc leur8reurqns Jo r{lp€erq releart aql Sursn
'dns n4(ntqo ntoqdoqruV ro1 Dssaozaaqluveurcu rrreu
'[rogstg
-aE
aqt
pasodord
uosueqog
ç06I uI
'rutu
LI-?I
's 'sn1d}souot se tuol s€ ocr/v\1'snldlsouo8
{}Eua1
go xade Eururelle lou asodor ur Eurlg 'r 'lurodplur
ol lelsrp Z 11âcpurtreuqns
tururol ure^ ]uerrncer
'b 'qloot
lsrld
lecrdeqns qlr/v\ elsrueJ Jo 9S :$ZZ
'ttg)
]uâsqe eleruJo /IJo ssof,ord relnper8 lseseq
rolno rrâr{l ueâ^r.}oq oJue}srp se tuol se ç I '-0I '
qleel uBlpeluqns lunlg orur qlyvt 4a iluasqB e[?ur
go areld lerprtd4 'd 'aprrurse tuol s€ seurr] g'I-B'0
'aleur8r€rua
'eldurrs dgeseq e^l€^ sluod
lou
yo a8prrg 'o '(c 'qZZ'stt.f) dgesropocrde pe^\ârl
app\ se Euol se sorurl L'l-Z'I o^le^ sruad 3o xady
'u 'ap1,v\se Euol se serurt
'paua11eg epur
B-ç
Jo
snldlsouoC 'ur '(c 'qZZ'stt$
seruereqnlord tur
-lcalord dplearvr '1ere1e1o/vu ro auo pue uorlceford
'tuo1 euo qU^À 'eldurrs d1e^tr1e1ereletu
lecrde
Jo
elrxorouo8 3o xedy '1 '(pZZ'ttg)
luesqe rrrortezrl
-orelrs
lewpnlr8uol lpeqoprl dgeseq'suortcalord
'1, '@zz
pl?relsl olncs o/vu qlllv\ dlecrde eFru Jo BS
'Etg) rslp
'
3o çEuâl se Euol sB seurrl I I - L'0 âtuep
-ode
leate
Ieretel pezrlorelcs Tr€p ro seruereqn]
-ord lnoqlr^r rsrp
Jo eseq laqol prelel pepunor
atrel qllÀ\ 'aleur8reure lou pepunor dlpcrde epru
'l 'GZZ'ttg)
luasard uorsserdrp I€relBI leete
Jo IS
pcrde pezrlorâlf,s dpqtrl qll^À eleur8rerue dl8uorls
o] dplea.rvr dgerpaurocrde el€ru Jo gS 'l ']uesqe
eleutrrgo aleld lerqrlrseg 'q '@gZ'qZZ'sEl.D elnre
ssacord letslp qllÀ\ snsrelrs€q pqq '(qZZ, 'ttg)
JIBTI esorunld Jo urtreur JorJe]sod lnoqrr/v\ olerueJ
'ttd
atprr ro r{}oo} rorre}u€
Jo elqll pulq :(lZZ
qlyv\ peuelleg dgeretel eprrr Jo snsr€lrseq pulH
'E ']uesqe sor{snrq r{}l^Àalsrrrgo EaplI
'J-p 'sedr}s
I
se Euol se L'-g' essol8ered lurnlueurerd se tuol
s€ B'-1,' lnoqe urnlueru '(ttZ,Z'Etd qloo] rorrelue
qtyv' Prlr{t I€sBq reeu run}uâru lsaqol o:[1-ra8ug
Iere^es qll/u\ dgecrde rrrn11eqeg l8uol sB aplrnr
se serurl +'B-0'Z al€tu Jo 'Euo1 se oprlv\ s€ serurl
'o 'Treru eled qllÀ\
B'L-Z'g eletueJ Jo eceds reptr I
eleru Jo alqrpueur l>1ce1qeleureJ Jo 'moged qlp\
ap1eutrrJo
ac€d 'q'slueurtos L'I-g'I lxau ol eleruJo
'.raqla8ol stueru8es g't-1-'f,
lxau 3o qfual ol pnba
eleureJ Jo 1 lueu8es rege8elg 'e 'uot\{ulsaq
'r{çZ
ern8rg
ur
u/v\oqs
se aceds rel€ru padolanap dlalerapou
o] [orv\
qlyv\ sarceds crlcrealed :(rtZZ 'tlg) {}oo1 rorro}ue
qly prlr{} I€s€q r€eu urn}uaru :(pZZ'Elg) peqolrr}
dgeseq pu€ pe/v\orreu llpcrde BS pue (rZZ 'ttg)
eqol pâpunor a8rel € r{tr1v\dlpralelocrde IS e1eru
'QZZ 'ttg) aleln8ue ro peqloot urtreur rorrelue
r{}yv\ eptu Jo snsr€}rs"q pu1q iqtoot pcrdeqns q}r/v\
'qZZ 'stt.t) elnf,e ssacord
eleutrrâJ
Jo gS :@gZ
lelslp
rorratsod qll/n\ eletueJ
snsretrssq puFI :(t&Z
Jo
'r0I 's8l.f) asournld
'eldrurs urtreur rorralsod
lou
'stsouantq
Jo srrer{ q}l/u\ âl"ureJ Jo elql} pulH
'f,rs"qolnv '/gBI 'des o4{ntqnn,roqdoqruy
:sarcadsadd;
'(uosgeqolf zs.çrowafi
-uv roretu?u
r'au)6eq
.i";,tJ"'J,rjiliriËi:o7'^
rrs"qouory' LtBI 'ÂeSo4{nrqnn,rot1{otltuy
:sarcadsadd;
'zrssetv
"3"ifi*';:,u;'
'(t+st
DS
s?otuaqtuv
ïi:'rii
'selsls pâtlun urâlsam aql w 'opnonaln{ pue
slwofitluac 'sarcods o/v\l r{}y/t 'ecgr;y pue ers€Jng
uI peerdsepyit sr snuatqns slql'(E"U)
Dnpta
pue'sloorg
ona?ttsaa'(E.tX) nî1oa'z1rrvrero141
Dgualuuqezrl'(1sm,o>1zsop€ll stsuauaqspal'olueqcs
-lpâd suaptsrnl 'asarrg ocnu(s 're1p14 suon{ns
'(eserrg)
'oluaqcstpad
'(aserrg) r{"t
sa{tut{s
4îrys1os
'z1r*re,toIN
'(uos
-xtaas 'zlruretoIN
llqaua7ps
lappnr
-rlrlrg) rlllcî"tponb.' zlrnerotr\tr txilslonzaqsrd'sloorg
aaaaturaouo('rar1a1ada1 oâopo{ 'zete 4 ocuold 'zlr.nre
-rotr\l pata{'z1uvrero141 ranrDzasuto'o>lueqcsrpod
'essrrg
'o>luaqcslpe.il
ao71o
sttlu?atJoenla
szlouotpuara 'oluaqcslped
où@ouolaut 'o>lueqcs
-]pâ.{ tsuaLta.a 'reusarr4 ota7qncou 'asarrg ofi@oc
-na1 '11ere>lcoCslunal 'pnerrg DparDI 'o>lueqcslpad
ruuow{noq 'stnoç1 slrolnâa.r.n'(sege4) soan 're1pp1
n4{1cuz 'aserrg
szquaazqlq 'z1rrvrero141 azlnranq
'z1rrvrero141sa{qnnÊ'eserrg
sldocnolî'z1u*ero111
mounsolï'.M
s.zlnlqqocm{'uu€tusre^g saQaln{
'e>lelraqrulJ
'z1nturotr41
opnntta1n{
stu"tonaod
'o>lueqcslpâ.{ suatsaangf'rar]a1ada1 naut7n.t"tr{'(t
l
-^llo) o4o"roua{'d1smo>lzsope}I rctluatlcpa{'puerlg
ruslailca 'opoqrrg
slwofitoqc4da 'l.1sr*o>1zsop€U
oaunqa 'uueursJelg
Dlqnp 'z1rrvrerotr41 Dlnttln
-sap 'e4JIpaH slurneîatne 'rcTJIV .t0lo2u0t 'o>lueqcs
-lpo.{ szuroeraCIlt 'z1rr*ero141 .t{otsttrlc 'uossar3
stwo{t"tquac '(e.(elsnoTrretrAI)
nsuagnfioq 'zata4
Dpuqq 'qlprs sllDsDq'oluaqcslpâd sa(lqnq 'rar1a1a
-dtT ranrnauf)q 'z1rtr-totr41 tloflotTso 'asarrg oïnaro
'reTJIV osltpquD'(E.tX)
sunuranD'o>luaqcslped
'sapa{g papnleul
'V
(o71fawo"to4)
'(pZ 'Stg) sîuronanle
xmnuzrqo
Jo eletu eqt Jo aeuualue peqqnlc aql pue (t t
'ttd
qnflo.rTso'V Ja e1etu er{} Jo eeuua}ue âlerres
leq/v\eruos eql s3 qJns n1ûauo.tD{ u\ dqlrornralou
ete sJaIJBJBTIJ xes drepuocas anbrun eruos
' opûauolzd repun npûaunlos pezrurduo
NrrsT-rng scNsrDS svsNvx do Àrrsus^rN^[I sHJ
9Lt
SysrpuATrcs
AND PHvroGENy
Anthophlra s. str. (in his sense) from Anthophoroides.
Timberlake's idea of Anthophora s. str. did not
include Heliophila ( -- Micranthophora), Amegilla., or
Clisodon which he considered genera. Timberlake
was the first to reco gnize the similarities between
Melea and Anthophoroides.
Comments.Melea is most closely related to Antho'
phoroides. Even though the two share derived
characters such as the short first flagellomere, the
mentum with an anterior tooth near the basal
third, the hind basitarsus of the male with an
anterior tooth, and similar male terminalia as
well as plesiomorphous male characters such as
oF THE ANTHoPHoRINn
477
Bnns
lack of midtarsal, midbasitarsal, and middistitarsal brushes, they are distinct groups . Melea can be
separated from Anthophoroides in the females by
the hind tibia with the scopal hairs all simple, 56
with a subapical tooth and the posterior distal
process of the hind tibia acute. The males can be
separated by their larger size (L3-I7 mm.), the
widely diverging rather than almost parallel apodemes of 57, the trilobed rather than rounded
base of SB, and the blunt rather long and acute
tooth of the hind basitarsus.
Biolog. Melea fly from March to August or
September. Several derived biological characters
adp
FIG' 22. Anthophorasubgenus Melea. adp:vcute distal Process of female hind tibia, l:lorum,
m=mentum,tm=toothofmentum. A,A.(M.)pkgiata,apexofgonocoxite,ventralviewofrightside.
B-K, A. (M.) abrupn. B, C, Genital capsule, side and dorsoventral views. D-F, SB, S7 and 56 of male,
ventral ,rie*t. G, Fhbellum, anterior view. H, I, Hind leg of female and male, outer views. J, Apex of
male T7 (hair omitted), dorsal view. K, Lorum and mentum, side view.
e^le^ srued yo a8prrg 'o (lesropocrde pe/v\er^
ss apl^À se Euol sB serurl 9'I-6'0 'geurs a^[e^
sruadyo xedy 'u 'ap1,v\se Euol s€ serull /'9-0.g
eputuJo snldlsouog 'ru '(J 't+Z 'stl.l) pezrlor
-elrs Trep pu€ eleln8ue dgerelel 'peqogq dgearvr
turreadde elrxoJouo8 serurloruos 'pedola^op
dpleem eqol pralnl lecrdeqns qlp\ rôot ftcrdL
q8noql 'eldnrrs dpueredde eptu Jo elrxorouot
'l '(E+Z 'Etg) luosq€ rrorlezrlorelrs
Jo xedy
IBu
-lpnrÉuo1:qt7 ern8rg ur se
algord 'papunor
BSJo
pue pedola,rep dpleam reuroJ lural€locrde
'(E+Z '8tg)
'turrceford
sâqol
Jo rred uerpotuqns
'pozrlorelcs
d11er1ua.rt
rlllÀ\ dgecrde âletu Jo
'T '(p+Z 'Etg) oslp Trep
BS
3o çEuel w Euol se g'I-0'I
eurapode iseare pozrlorâlJsTrep ro sorueraqnlord
]noqlr/v\ dleseq rslp ia+Z ern8rg ur s€ algord
'rqol
peorq
qllÀ\
punor
dgecrde
el€ruJo IS
lerelel
'l '("+Z
'ttd luasard uorssardrp
1erelel lpezrlor
-elrs (pqEU 'dgerparu eleur8reura (prrorreu xade
rlll/u\e[?tu Jo 9s 'l 'tuesqs al?ru 3o ereld Isrqrlrssg
'r{ '(l+Z 'Elg) snoqpq 'pare11pdllensn aptu
Jo
rnruâJpulq :(l+Z'EtO xede r€eu awds uo pauesur
rnds lelql] ouo 'eurds pcrde 'louelue rllrm pe]Blrp
el€utrrJo
erqrt puFI :$.'l+Z'stl.{) pe^rnc o1lq8rerls
rlloo] Jo ur8reur JorJelu€ raddn 'qlool rorJalue
qlyit epur Jo snsrelrseq pulH 'E 'luesqe eleru
'y-p 'sedlls se tuol s€
Jo saqsnrqqlrm trlplry
B'-ç'
lnoqe essol8ered lurnluaurerd se Euol se L'-ç'
rnoq€ runlueru '(lZZ'Etg) prlql IBsBqrsou r{loo1
rorralus r{}yv\ runlueru iseqol a>p1-re8uglere^es
qly/' dgecrde urn11eqegltuol se âpp\ se serur]
'(sanq] gI zanrolpa)Euol se epr/v\
t'I I -0'L el€ru3o
'c '(o1oufus
se seurrl L'B-t'9 eleuralJo acedsrel€trAtr
uo sTrer.ualrr{/v\ldacxa) Trelq elBuroJ1o '(onutortpt
pu? unr7llno 'vul rvrolel')ellq/v\qllÂ\ el€tuJo ec€d
'q 'slueur8as
0ç' I -gg' I ]xeu ol epur Jo lroqtet
-ol sluaur8asgg'g-0ç'Z ]xau Jo {fua1 o} pnba
aleural Jo 1 luaurtas releEelg 'e 'uoz7{t^tcsag
sr d8olorq âsoq/v\ serreds crlcreepd râqlo dpo âr{I
'@gOI 'lTsû\ol[,om) Dsllfa DlelA se
qrns seqc]e^
elqee8euerrr e s€ lerlualod e^eq
Jo rotsulllod
,{eu o4ofo1d 'V '(OOgl) punl-Ernquaseg
pue
(nZOey)gaoqran '(OZO1)opell '(trcù rrunrx
puu cr^opslf '(ZOO1)ueslelN '(q 'nI96I) rc2co1111
'(OeOl) ede>lsnoTTretr^tr'Qtpunuo"t
ss pegodar
'9t6t)
'GtOù
sorC
>lezseuefl
se qons (ouqaurn(
'tr lnoqe ere Dalaryrrlrree
se pepoder) o7o13o7d
-Fd tururecuor sreded luepodurl '6o7o19
'(nzz
'Elg) uortcelord pcrd€ ot uor1lppe ul
q]eât FrelBI
o/vu rlll/!\ elsul Jo âlrxorouo8 Jo xedy 'l 'Euo1
se l.lpseq op1,v\ su Lt'-tt'
qlool 'JIBI{ pcrde uo
qloo] IIBurs r{}lû\ ol€ru Jo snsrs}rs€q pulH 't '(qçe
'ttg o] repms) sarurl
'tuo1 sB
B't-g'Z eleur Jo
aplm se serur] t lnoqe olerueJ;o aceds æletr I 'r
dnorg sarcodg e1erte14
'otoq{otltuV
'sl1oatoq
(otoqdotltuogot
learoq dpo eql ere deqt
-s(W) 'V pue ulplsîlC r{tl6 '?ueqlv
Jo }sotu pue
BrqtunloC qsrtrrfl 'sa1e1g pâ]1un âr{} }noqEnorqr
punoJ ere pu€ rrlrrv
er{} o}q reJ aEuer daql
pue uerrreruV quo51 ere sarreds reqlo eq] luurq3
uJâlseaqlnos ur sJnJJo suotrtÊru 'V 'uosseJC
'1era1co3
'dqrly
slpruapîceo pue
suo{tÊru
saptnq'utoq'iev,Soy{n"rqo(oqrW) 'V 'saltadg papnlcu7
'(çgO1 'erleg
pue uoproN) sçed lsuoture] rrern tururl]no s]ca[
-go ol 1r dldde pue 'uorlarces puel8 ælnqrpu€ru
qlyv\ ll turxrnr dlqenrnserd 'qsnrq 1erqel eq]
ur ecrnl eq] tcelloc 'ansslt ('-I DauzsDtautlsz;r) drus
-red /v\eqJ o4{n"tqn sepur aql 'srrer.I enrldrospe
Jo
Peuelleu eug Jo r{snrq pcrde ve seq ulnrqel eqr
sercads reqlo ge Suorue
leqt ur rurroqdoqluv
Jo
'Stg) Dalaç
oïTI r{}oo}rorre}ue qruv\Jm
Joserceds
enbrun ere suo{tÊru pu€ oy{n,tqo 'V Jo seletu
leseqr€âu runtuaru :(l+Z'Et$ dlecrde pezrlorâlrs aqJ 'o1(ntqo 'v
Jo dtolorq eql uo pesoder (ZAOI)
dplrep eleur Jo elrxocouo8 :(E+Z 'Elg) cslp ruor;
'dgeuort1ppv
'dnort
q8norqrers
pu" ueproN
alEue tq8rr le turlceficrd 'Euo1runlnrrds 'pepunor
dgeseq 'aqol tr€ralelpepunor pue âqol uerporuqns sarcads seproqurog egl se DalaWJo serceds crlcreeu
er{} Jo dtolorq pue uo ernteretrl oqt peû\orler pue
aleptu€ rllr/v\dgerarelocrdeBS :(p +Z 'Etd rerllo
srrt€ruelsds eql pessnrsrp s€r{ (g86I) sToorg 'serr
r{ree or Ieilered }sourle seurepode 'eqol pepunor
'llz,
-eds esewqC euo qllÀ\ uerrreruV
eplÀ\ t{}1,v\dlleralelocrde e1eru
/S :(l
çro51 dpreru
'stt.l) qlool rolrolue qll/v\ eleur Jo
'(îop1g puo sluarawoC
sr dnort sarcads sF{I
Jo snsr€]rs€qpu5{
'@zz
:(f 'l+Z 's8l.f) aurds 3o prlr{} pcrde uo peuesur
'ttg) uorlcaford pcrde o] uorrlpp€ ul q]oo] pre]€l
rnds FIqp e qllÀ\ 'dgecrdsorâlue slqll puFI uo
'l '(lzz'tlg)
aurds qllÀ\ e1uurlurur gI-0I tp8ual 'stsou7mq
e
elBru
qlr/v\
Jo alrxocouo8 5o xedy
'JIeq
'uorleu8rsap
Fur8rropue crddtouotrAtr Euol se dlewq apr/v\se ?ç'-Zt' r{}oo1
les€q uo
'9681 'geraTroDunnlpo sntrqopod:sarcads
qlool etrel qlp\ eleTuJo snsrelrs"q pulH 'E 'saurrl
addl
'(rzl 'El.l or rspurs) tuol s€
'B+ 'd '1061 'geralcoD pu€
IIârâ>[roCsaprctotldoqruV+'B-L'ç alsur Jo
ePy/' se seurrl B'1-ç'ç ol?rrleJJo aceds r?l?tr I 'r
IIereTroD pue
dnorg sarcadg seproqurofl
Ilare{c oJ saplo.roq{oqtuv srrrta8qng
'seeq
ruo{*sra'uereqr (,Jrî,T?
î,:rii;"r""5',ti,ï
elqutrrnq Jo scnurur dpuaredde ere oapry 3o sarceds
oql Jo IIV '(ggO1 'qoorg)
ror^eqeq tur8urts Jo
out(nsn1qo'nM stsuauoyïuow'gera1co3 nyoa"t"ta{ssol luaredde eql pu€ soJueJluâ lseu eql JâAo s1âr
'nM
'otoqdoqruV rrr.
-rnt
szznrlq1zrautt
(rrtrW)'V'sa4ca(g
papryguJ
Jo uorlJnrlsuor epnlcw âseql
'(çgOI 'leld) opatta{ 'V sr.u/v\ouT exel râr{}o urorJ Eur}eredes oapry o} anbrun eJe
lr
NrrsT-rnfl sDNsrss svsNvx do Àrrsus^rNn sHJ
BL+
SysrnuATlcs
AND PHvlocENY
basally simple, 0.75-1.10 times as long asrvjde
(FiS. 2+f). p. pygidial plate of male absent; T7 of
*ale with iwo short submedian teeth .I+-.26 as
long as distance between their outer bases (FiS.
24b); gradular process of T7 of male absent. qFirst recurrent vein joining submarginal cell 2
distal to midpoint. r. Sting in repose not attaining
t o s u r p a s s i n g a p e x o f g o n o s t y l u s, 2 . 2 - 3 . 0 l e n g t h
of gonostylus. s. Length 10-13 mm.
Commenls.T. D. A. and W. P. Cockerell named
Anthophoroidesbecause of the five-segmented maxillarypalpi. This character is only of value at the
rpe.ific lével, being found also în A (!gro1"eg.illa)
Timberlake
futaicauda and in many Heliophila.
"1tOS1)
included species of Meha with Anthoph.0r0r.des (see Commenls unde r Melea). Anthophoroides.is
-ori closely related to Melea and its separation
from that subgenus is discussed in the Comments
on Melea.
Biotog. Species of Anthophoroides, a New World
subgenus, are xerophilic. Linsley (IP+2) gate ?
detaliled account of the biology of A- linslefi.
Giltiam et al. (1984) analyzed the microbial fauna
oF THE ANTHoPHoRINn
BBBs
+79
of an undescribed species of Anthophoroides.The
territorial behavior of the males of Anthophoroides
is the same as that of Melea in that territories with
distinct boundaries, usually enclosing host plants,
are defended against conspecific males. The females of Anthophoroides nest aggregatively in
earthern banks excep t signata which nests in dead
pe.rs.
Joshua tree stumps (Yucca) (R. Snellingand possibly also in the ground (Brooks,
.o*-.)
are generally po_lylectic
They
unpublished).
though phaceliae is oligolectic on Phacelia (Hydrophyllaceae).
The species which inhabit the regions of the
Sonoran and Colorado desert are probably univoltine but possibly bivoltine. There is a normal
spring brood(s?) from February (two records in
Jànuàry) to mid Jtly. In those dty areas which
Lu,r. -id to late summer rains such as southeastern Arizon a, califtrnica has a second brood. .4.
marginata, the only truly neotropical species, .is
r.^ronul in southwestern Arizona and probably in
the seasonally dty tropical areas of México but
rnay be multivoltine in other Mexican and Cen-
females.Scalelines(onrightside)represent
FIG.23. AnthophorasubgeneraMeleaandMystacanthophora,
of Anthophora(Melea) abrupta and'.4.
plates
Basitibial
B,
A,
proc"ss.
.l mm; dp=iosteriorii.tal
abrupta and A'
(Mystaiantiophàre capistrata.C, D, Orrt"t view of apex of hind basitarsus, A. (Melea)
trata.
c
aPis
(My stacantho
Phora)
'/ûer^
'oclu,rofitoc
'V p Ee1 pur11
relno
('ù
'| 'rtern relno 'trl pulH 'l 's^{.or^
'H 'C'/v\âr^
'Jslp
ePIs pue
ler}ue^'BS
Ier}ue^osrop'alnsdec
1e}rueC [
'l,S'g'O'1v\or^
o
p
l
s
'
e
l
n
s
d
e
c
p
l
r
u
a
g
'
3
'
r
v
r
e
r
^
Jo s/ûer^ eprs puB leJlue^
Iesrop'(prrrl*o leq) LI Jo
xedy 'g 'rvrarnlerluan '9g 'y 'tun.rllpa ('ù 'v'l..-v 'septu 'sapto.ror1fuqruv
snua8qns otoqdotltuv '+Z'cl.f
ffi3
Nlrsa'rng scNsrcs svsNvx do Àrrsus^rNf) sHJ
OB?
SvstnuATICS
AND PHvrocENY
tral American areas.
Included Species. A. (Anthophoroides) californica
marginata Smith,
Cresson , linsleyi Timberlake,
phaceliaen. sp ., signata n. sp ., aallorum (Cockerell),
and six undescribed species. They range from
Colorado and Texas west to southern Oregon and
California, south through México (including Baja
California) to Honduras. A. californica and marginata are the only two species of the twelve which
range into the neotropics.
Subgen us Anthornegilla
Marikovskaya
AnthomcgillaMarikovskaya, 1976a, p. 55.
Typ" species:AnthophoraarcticaMorawitz, lBB3. Original designation.
oF THE AnrHoPHoRrNn
Bnns
48 1
apically trilobed labrum (FiS. 25i); apodemes. of
S7 of male short, widely spaced and diverging
(FiS. 25e); SB of male with long-apodemes and
dpièulum projecting anterodorsally in the same
plane as disc (Figs . 25b, c); aPex of penis valvewithout fine, small hairs; female with hairs of
posterior margin of hind tibia simple (similar_ to
FiS. 10e); flabèllum apically entire, apical half to
twô-thirds narrow and elongate (Fig. 25d).
Description. a. Flagellar segment 1 of female
equal to length of next three to four segments
together, of male to next 2.0-2.5 segments. b.
face of
Faie of male with light yellow-white;
of male labrum
female black; apical margin
trilobed, mediar lobe emarginate (FiS. 25i). c.
Malar space of female about 1.7 times as wide as
long, of male 1.6-2.0 times as wide as long (FiS.
25h); mentum about .7 as long as prementum,
,*3:r::'
î,oY'li',:Iï
i ffiliffii"**.'TiH
B
FIG. 25. Anthophora(AnthomegiltQarctica, rna)e.A, 56, venfial view. B, C, S8, side and ventral views'
vie"rr.-E, 57, ventral view. F, G, Genital capsule, dorsoventral and side views.
D, Flabellum, Lt".io.
H, Head, side view I, Labrum. J, Apex of T7 (hair omitted), dorsal view'
s€ ecr/vç ]noq€ '1urod dreqs B s€ pa8uolord dgeseq
se^F^ sruad 3o eEpug 'o (gesropocrde peÀrer^ se
eplû\ se Euol sB serurt L'C ]noqe '(EgZ 'Elg) pe/v\or
-r€u dlpcrde pus eletuola e^lel srued
;o xody
'u 'eplù\
se
Euol
se
serur]
B al€ru yo snldtsouoC
'ru '(q 'tgz, 'tttg) alrxorouo8
Jo eqol lerluâ^ Jo
p"retel eete snouerqrueru € pue pssetu uorlcalord
e qly/. luorlceford lerlue^ >lee/v\ r{}lÂ\'aldurs
e1etu Jo elrxocouo8 3o xady 'l '(rgZ 'Eld tuasqe
uorlezrlorelJs leurpnlrtuol ilq8rerls ]sorul€'el€l
-n8ue dpleam ur8reru leralel 'peqo[q 'pervrorreu
'T '(pgZ 'Etg) cslp
(lecrde el"tu
Jo BS
3o qfual
ue{} retuol dpqtrls ltuol enrepode iaqol l€srop
e ll ol luseq dpq8rls pu€ oqol lerlue^ e 'saqol
Tee/v\ o/v\] rlll/v\ ur8reur lBra]elocrde leil]ua dlecrde
aleru Jo IS '1. '@gZ 'ttg) p€rluol paxeg urtreur
'errlua dlecrde eluru
'l 'luesqe apru
lecrde
Jo gS
'q 'o 'pgI sarntrg ur sB rrer{
Jo e1e1d plqllrsefl
aldurrs Eurneq ur8ruur purpnlr8uol rorralsod qlu*
al€rrrâJ Jo elqlt pqq ieldrrrrs epur Jo srsrelrseq
pulH 'E 'luasqe e1eru Jo seqsnrq rllyv\ EtplIN 'J
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-ard se Euol se 6' ]noqe 'prID les€q reou q]ool
rorrelus lnoqlr/v\ runlueur :(qgz 'Etg) seqol uoqs
:@gZ, 'ttg)
snorerunu qllÀ\ dllecrde un[eqel]
epp\ uer{t ra8uol ep}ll e aceds æl€tr4tr 'r 't{r8ua1
ur ruru gI-1 1 essolt '€xo) pqq turssedrns asoder
8uo1 eap8 lada
r{tpl M
Erl
ur '(-ZT-I1)
Jo
L'l
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elgord
ur
eleruso
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'req]eEo]
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lxeu o1 pnba ]noqu âletu 3o
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olsureJ Jo 1 luour8es rege8elg 'e 'uot1{t.tcsaq
'e 'p01 sern8rd ur se
rler{ eldurrs turneq ur8reur leurpn}rtuo1 rorralsod
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/v\oJJ€u se petuolord 'e1nJe dleseq sa^[€^ srued
'(tgz 'Eld /v\orreu pu€ ele8uola e^[B^
3o aEprrq
e}rxocouot ;o eqol ler}ue^ Jo
sruad '(,{gZ 'ttd
peJe]el eete snoueJqruaru e pus peseru uotlcelord
e qll/u\ epsdec plrua8 :(qgZ '8lg) seqol uotls
runlleq?u
snoreurnu qlr/v\ dlecrde'pedÉqs-uoorq
iurur çI-+1 essol8;o çEua1 'exoc pug Eutssedrns
asoder ur eelet 'Euo1 dleurarlxe suedqlnour lepun
uerll re8uol aplll e aceds relstrAI 'stsouflotq
oI61'lare>1co3
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Isrluer ruor; srnrro snue8qns sIqI
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oua81q7'zlvvrero141 snaaucos(tc'z1u*ero141 Dplnu!
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n s u a ï m f t a q ' z 1 u u e r o 1 4 1s a ( u ' n n ' z 1 r t t r e
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sr s€ 'saeq
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lsoru dllua
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ur PunoJ sr uorlezrloralJsep l€s€q r{}lû\ parued
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âtrxorouot
eql Jo eqol lertue,r - ] eelleslo^
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(q 'eg LGI) edels,roTlr€tr\tr
l.geseq pelrerrp
runlnords pue seruepode Euol qll^À BS pu€ qleal
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eql Jo srâ]c€rer{r leuol}lppe sql dq peteredes ere
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pat€1er
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ot
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ol Ielsrp Z 11â]leurtreuqns Eururof ure^ ]uorrnf,er
'b '(fCZ 'tlg)
luesq€ eleru Jo LI Jo sse
]srld
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eleru yo areld lerpr8d4 'd '("lEZ 'Elg) eplÀ\ se Euol
s€ seur] t' uor]eur8reura '(atprrq uo seJuereqn]
-ord Jo rred e qlyv\ oua31ru1) uol]eul8reua
daep qll^À â^[€^ sruad 3o a8prrg 'o '(EçZ 'Etg)
rleq lnoqlr/v\ '(gesropocrde pe/v\er^ s€ oplÀ\ se
Euol sB serurl g o^l€r srued;o xady 'u '("1çZ 'Elg)
elrxocouot Jo eceJrns Jeuur uo sJrerl qll^À Jelsrlq
e sB turureurer es€q aqr dpo '(oua7gol 'Z w lsol
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'l '(c 'qç2, 'sttg) rslp agl sB aceld arues
3o xady
aql ur pue l.llesroporalus Eurtcalord 'a41-aurds
qloq 'tuo1 tunlnf,Ids pue saurepode 'luasqe uol]
-uzrlorelos Isurpn]rtuo1 'a1e1n8ue dlleralelocrde
epru Jo BS '{ '(tçZ 'Etd seere pozl}oralrs TJep
ro âcueraqnlord lerolel lnoqll/v\ JsIp Jo es€q lcsrp
'padola^ap dlleervr utEruur
;o qfuel 6' anrepode
'pepunor al€tu
xady 'f '(nçe
lS
Jo
5o
I€relelocrde
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Iere}Blocrde /v\al " }dacxe alduns rletl
'l 'tuesqe
'arrluo dgzcrde ol€ur
Jo gS
IIE lsorule
oleru ;o areld lelqllrsefl 'rI '(rOl 'Etg o] rupurrs)
eldrurs elql] pug uo euoz rolrelsod purpnlr8uol
Jo srreq qll^À eluurâJ leldrurs â1eur Jo snsrelrwq
seqsnrq qll/u\ Et1plf.t
pulH 'E ']uasqu eleur
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zB+
SvsrnuATrcs
AND PHyrocENy
oF THE ANrHopHoRrNn
Bees
+83
FIG. 26. Anthophora (Rhinomegilla) megarrhina, male. A, 56, ventral view. B, Flabellum, anterior view.
C, Head, side view. D, E, 57 and SB, ventral views. 4 Apex of T7 (hair omitted), dorsal view. G, H,
Genital capsule, side and dorsoventral views.
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0'I I -L'ç
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paqo[q L14eam algord ur e1etu Jo e]rxorouo8
'l '@tZ 'Etg)
luosq€ dlensn uortez
Jo xady
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ol
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'urur
6[-8 rp8ual eluru
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se Euol
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luesq€ âletu Jo LLJo sserordrepperE
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ruor; Burrec uerpatu e dq pepl^1p dpulpnlr8uol
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e qlp\ dlprpeur
'd '(qgZ 'ttd opp\ se Euol
ep1eur
Jo LL p xady
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sHJ
+Bn
SvsrnuATrcs
AND PsvrocENy
oF THE ANrsopHoRINn
Bnes
B
K
485
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pue
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ù v ?*errr1er1ue,r ;o xedy
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4;;o xedy
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s n u e 8 q n so t o t l { o t l t u V . g Z , C t t
d,g,C.y.sepur
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runlueru :(p 'c6Z's8lg) arrlua dgecrde urnfiaqeg
rlll/n\ cslp Jo elpprur 'rqol Issrop pazrtoralcs tq8lt
!Euo1 se epl^À se sârurt g'ç lnoqe epur Jo 'tuo1
pu€ eqol lerlue^ pezrlorelrs TrBp e 'seqol orvr]q]lm
se âpr/v\ se serurl 4 lnoqe eletuoJ 3o aceds rel?trAl .r
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Jo
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'(ugZ 'Etg)
'q 'aldurrs eletuJo snsretrwq pulH
dgerluarr
a1e1d
l€rqrtrseg
'E 'tuosqe el€ru
'J-p 'sedrls se
rlerl Jo qcled'punor'r{rl.{r
q}l^À â^le^ srued
Jo seqsnrq trlplry
'agl arn8rg ur sB aldurs elqll pqq uo ur8reur
tuol se JIer{ }noqe essoltered lun}ueruerd se tuol
rorrelsod Jo srrerl r{}lÀ\ eleureJ :(p 'c6Z'stlg)
se 9' lnoqe runtueru '(oslwa ldacxe) qloot rorrelue .
'eldurs
arrlua dlpcrde runilaqell :(qOZ 'ttg)
palurod
lnoqlr/v\
PrI{l leseq reeu runlueru iseqol
puu ele8uole a1e1d1elqllrseq qlyv\ oletueJ l(u ralce
a>p1-ratug lere^as qtp\ dgecrde urn11eqzg iEuol
-Jer1f, 'uotg{u,csaq ess) elrxoJouo8 qlp\ snoroloJ
$s3apr/v\ se sourrl L-ç 3[Bru5o 'tuol s3 epr^{.s€ sarurl
'r 'aqol uerpetu aleurt
-uoJ'pazrlorelJs l.1a1a1duroc'lueserd
ç-t e1eureJJo aceds re1etrAtr
JI'a1eur
-r€tue qlyvt 'Quo\lom pu€ pslexa 'Dx.totsnutur eldrurs
ralelueprrl'lTH,iH
Jo snldlsouo8
'suot{tagco1'V ul os Lpleârv\) (8
'Elg) uoura?stlnpd
,îîïK3
n
'6981'uossar3 sqDununy
ororldoqîuv
:sarcadgedda
se paqolrr} dgecrde â1eru Jo rrrnrq€l !>1ce1q
ur
'6+I 'd '6181'uotlu4 uoposllC âFural
'r*o11ad ot elFI^À durearc el€ru
Jo
Jo er€d
'q 'slueru8as
ç'Z-0'Z ]xeu ol epru Jo lraqla8ot
olted uopos!1, snuetq.s
r{}tua1 o} lenba }noqe
sluatutes eerqt lxeu
Jo
al€urrreJJo 1 tuaur8as rege8elg 'e 'uoq{uosaq
'GtZ'Elg)
'?ulr{c urar{}nospue
aplû\ 'pnr]
-ua^oJolue ]ueq urtreur pcrde [€n]c" 'q]ee] u"lp
Ierluac ol PJe/v\]sBepu? uortar ueâusJJellpetr\t -euqns
oûu qly/t dgecrde eletu Jo LL 'luasqe et€ld
ot{l 'er,reurpuers 'puelEug Eurpnlcur adorng
prprtdd :(ltZ'ttg)JprI
FsBq uo rrer{3o ped esuep
lnoq8norr{} punoJ ere dr.IJ 'llârâTroC tuoîf)m r{tlm gS :(l 'ttZ 's8td peqoilq
ol€ru Jo e}rxocouo8
pue zlrlv\erotrAltsra(nazs'("razue4) DplntÛan.tponb
rslp geurs pue seuepode paler
5o xade :(ltZ'Etg)
'o>luaqcslpod
Dzrnesnra'a>1crpe11
suotrtaçcq'z1rtvre
-edas dlaprr* 'tuo1 qll^À el"ur Jo IS 'stsou7mq
-rol4tr Dsltxa (o17paur(soq) 'V 'sapa{g papryeuJ
'crddr
'(ZOO1'ueslerg)
'B6LI
Dplnzuxultpnnb'V sr u/v\ouT sr d8olorq Eurlsau
'Goplg
esor{^r.o716aur(sDcI
Io sarcadsdpo aql
'el€ru oql
Jo gSJoJIBqleseqoql uo rleqJo r{r}edasuep'atre1
oql ol turrregar 'uoxu] palelar e Jo errr€u e opûay4J
snld 'drreq dlesuep '('rC) sûoq '(Eopu(gg
'ruru
'a1eure;'nrru
'e1eur
ç'T,I-0'01
0'I I -0'6
ql8uâT 's 'sn1d1souoE5o
3o
çEue[ seur] g'U tnoq€
'snldlsouot
xade tururelle asodar ur turlg
'r 'lurodplur Jo
ol lelslp Z 11e]leur8retuqns tururof
ure^ luerrncar lsrrd 'b ']uesqe ol€ru Io LLJo sse
-cord r€lnpsrt :("lf Z 'ttd epp\ pue pertua^oralue
luaq urtreur pcrde l€nlJs 'qlaal u€rpetuqns'lroqs
'a1nceo/vuqlp\ oleru
xede lluesqe âl€ru
Jo
lJ
5o
yo aruld p1pfd4 'd 'eprrnse tuol sB serultç'Z-0'Z
'o
'eleurtrerue dgeseq o^l€^ sruad
Jo atprrg
'e?lâs peJâlt?f,srllr/v\ epl/v\ sz Euol s3 ecl/vulnoqs
e^le^ srued3o xedy 'u '(gZ 'El.d epl^Àsu Euol se
poqo[q
sorurl9 aleuryo snll.lsouoC 'ru '(ltZ'ttd
xady 'l '(\tZ
elrxooouoE
elgord ur âleru
Jo
3o
'Etg)
vorlez\lorâlcs
luesard
leurpnlrtuol'ezr€ruor
'peqo[q dplear*
Isre}€l
JIer{ lecrde uo urEreur
eleutllJo BS 3o xedy 'T '6tZ 'tlg) rslp go qrtuel
s€ Euol se serurl ç L' I-00' I euapode Jo {}Eue1
'ecuereqnlord ro etprr esJe^suerlpâzrloralcs
TJep
-ouotrAl
rezuedo?rrynenxu!ryrnb
sldv :sarcadsadda
'NZDI aqt dq
,,rsrr ua'uelrg,, âI{l r;n::ri::î'$R
aqt dq petepr[e^ul'ng1'd'[ZOSI-I0BI] 'aurrnf snlsv7
'GO1I lsrrq3 cau '964I rezued outQnn
sldv :sarcadsadÂ;
stdv - ) gO/ I razued DpptDzat.rponb
snuatqns ^Àerr o41$au[*e
'er1otuotrAtrurelse&\qlnos ol preÀusee pue ueeu
-srrelrpetr tr dpreur sr dnort srrll Jo uorlnqrrlslp
erIJ 'opoqrJC tuounz pue 'eserrg ruqaîam 'ueT
-JIV ol oglpom' o>luaqcs]ped sll !w! s'd>12
srvro>lzs
op€ lI
(zar)d
{lsmotlzsopn^t
osofint
l:r7loaîr'oluaqcslped
'raTJIV uausat"t{ '(srapunes) anonlorl 'zata4 oîDlllt
-o.tï1u 'o>lueqcslped Duunw 'esarrg xaruou 'opoq
-lJC DlnetJluî 'zltl.l,e-,tolN tuasuoq 'reuserr d Dlnst4tca
'eserrg ozJnlneîune'rerleladel sa{tsso.tc'rer1e1ade1
zat)d Dnltlqlz'a1crpe11
otDrnelo,'re1;IV saycsofiqTo'
papnîtuJ
sutuaanuav (uoua1sllop1)'V'sa1ca{g
'reqtua]
-d.sol rrdv ruo{,"t?1,,rî;Ë}il"iW,
eq] alersdas ol sJeIJBJBqJ poo8 ou tuor; 1v\ouT
'dgerpaur pe/v\orreu ss dlreau
]ou sr qclq^À BS
I
NrJs-I-rng scNsrcs svsNvx .{o ^rIsugAINn
gHI
9Bt
SvsrpuATrcs
AND PHvrocENy
oF THE ANtsopHoRINn
I
Bnns
+87
' sM er ^
Jor J el u e
pue rorrelsodo-ra1e1nlnu?uuray(uoposln) 'V Jo €llâqelC 'C 'C 'eleural 'nlnutwtaT
(uopos1p) V pue eleru nspnalpa (uoqdoqqun7{4) ntoq{oqtuv Jo sa1e1dl€rqr}rs€B 'fl 'V 'ç1 'C ur ruur
'fl 'V ur urru ]ueserdâr sâurl âl€ls 'uoposllJ pu€ otoqdoqqunï(4 u-raua8qns onqdoqruV '67, '
I0' pue
I'
OL{
'p)
'sdnorE o/vu pazru8of,ar ar{S 'sarceds
tsrg aqt
/ I pâpp? pue uopqsllo Jo uorlrugâp âq] peuâp€orq
'(OgO
t ) ^ourrrus Jo poqlâur srsLpue f,rurouo
-xe] aql Eursn '(6/
u.,{e{s^o>lrr€I4l 'slualawl7
61)
'{uru g -6
I
qr8ual 's 'sn1L1souo8se 3uo1 sz serurt Z'Z lnoqe
'snlLlsouofi
Jo xade tururelle asodar ur Eurlg
'r "lurodpnu 01 plslp
Sururol
Z
IIer
leur8-reurqns
ure^ luerrn)er]srlC'b'(c'qBZ'r8tg)'pado
'paua11ug
-le^ep epur
Jo LL Jo ssarord rulnpzr8
L11e-rtua^osrop 'qlaal u€rpetuqns T, rlll^ dgecrde
epur Jo urnrpr8.(4 'd 'âplm se 3uo1 se seturl t'e
'ateurS.r?ruâ
dgeseq
â^lel sruad 3o a8pufl 'o '(ege
'3lg) rluq -re8uol
Jo r{f,}ed punor e r{}rm l1p-rtuan
pue rlsq uor{s q}l^A peralol 'apryt su 3uo1 s€ sâurr}
g'I e^len sruad 3o xady 'u '(nAZ '8tg) o^{} âq}
uâ3/v\leq uorlssJsurâp ou sr eJâr{} r€r{} r.{f,ns3}rxoJ
'luesard
'a1uur
-ouo8
3o xadu qllm snoJolof,uof,
3r
'ru '(rg; '3lg) eqol râ]no 'pseqo
Jo snll.rsouog
-relel ralprus (rynuru.uay) auo to (n1nu,n/) olw1pus
âqol reuur 'lerrde 'a8rz1 qllm 'algord w (stlautuuaT)
NrJsT'rnfi scNslcs
pâqollrt âleru Jo etl
o] (nqouni 'fl
paqo[q
-xof,ouo8 Jo xady 'l (l 'lBU 's8tg) tuesqe uortez\t
'a1z1n8u€ ol pepuno; ut8rutu
-o.râlf,s
I€urpnlr8uol
'T '(tgZ
'peorq l.lpcrdz epur
Jo BS
Iere]zlocrde
'(o 'ur
'Etg) rslp se 3uo1 s€ sâtur]
B'-g' aurapode
-{BZ 'r8tg) rl€r{Jo pueq uerpâtu l€urpntr8uol rltlm
pue uorlralord pseq 'pra]e1 q]lm lslp 'tqol lesrop
p 1l ol l€s€q dpq8rls puu âqol l€rtue^ p r{tllnt rslp
go ur8retu leJelelocrde 'aleut8r€ure ,tptolleqs lnq
,(peorq xade 'peo.rq el€ru Jo LS Jo rslg 'f "(- 'r{BU
'r3lg) euo usrpâru 'psuq erour ,(pq8r1s 'ralpurs
e Jo epls râr{}Ie uo dunq a8-ru1 q}lm JIer{ rolr
-elu€ uo lsrp '1.11e-rtue^
pâxeg ur8.ruur pcrde 'aleu
-r8-reura .(gzrpauroctde elelrr Jo 9S 'l '(qOZ 'Stg)
palurod dgecrde 'ale8uole â1etuâJJo 'luasqe âFtu
yo aluld 1urqrlrs€g 'q '(aO1 a-rn8rg) srreq asounld
'aldulls srl€r{ pdocs q}l^
Jo prreq Jorrâtsod 3ur>1ce1
âleruâJ JO €IqI] pqg lpzr-ralsod parr-rnf,pu€ peua]
-lug 'aldurs âl€ur Jo snsr€]rseq pulH 'E ']uesqu
âleur Jo seqsnrq Sr1pl6 'J-p "sadrls se 3uo1 su
/' €ssol8e-rediprrqt l€s€q rzeu qlool rorrelu€ lno
-qllm 'aldurts urnluâul 'utnlueua.rd se 3uo1 se L'
svsNvx
do ÀJISus^IN{l
aHJ
BB?
SvsrBvrATrcs
AND PHvrocENY
furcatus group) includi ng A. furcata (P anzer) , terminalis Cresson , borealis Morawitz,
meridionalis
Fedtschenko, raddei Morawitz, pubescens(Fabricius) , dufourii Lepeletier, wegelini Friese, riuolleti
Pérez and urbana Cresson and the second (A.
parietina group) consisting of plagiata (Illiger) [ - parietina (Fabricius)] , nigripas Morawitz, orientalis
Morawitz,
siewersii Morawitz,
excisa Morawitz,
uulpina (Panzer), muscaria Fedtschenko , hortensis
Morawitz,
and
giacilipes Morawitz.
Marikovskaya's broad definition and diagnosis of
Clisodon and of her two included groups is based
on characters showing much variation. In my
is polyopinion Clisodon (sensu Marikovskaya)
phyletic and represents seven distinct lineages
which I place in other subgenera of Anthophora.
Clisodon (in its usual restricted sense) has often
been accorded generic rank (Patton , lB79; Popov,
1951). Its mandibular and basitibial plate differences from other Anthophora are striking, Do doubt
associated with its wood-nesting habits. It is,
however, only a small derived group of Antho-
o"!iï;;l;
i:#Ë::
f:f:ffii;:t'1,ï
$ru;l
the resulting paraphyly of Anthophora).
Biologlt. The adults are active from mid May to
September. Many of the uniquely derived characters of Clisodon, which easily separate it from its
close relative s Dasymegilla and Petalosternln, such
as the tridentate mandibles and basitibial plate of
the female, are presumably adaptations to wood
nesting. Wood as a nesting substrate is known
only for Clisodon in the Anthophorini
[except
perhaps for A. (Anthophoroides) signatal. Females
nest in fallen logs utilizing beetle exit holes, and
in rotten wood, stumps, telephone poles, fence
posts, partially decayed or pithy sticks, etc. t(W.
P. Cockerell, 1903, and Medler, 1964
terminalis); (Nielsen, 1902; Popov, 1951; Radchenko,
1984 - furcato)1. Because of the preferance for a
wood substrate, Clisodon is usually a forest
dweller. I have observed the dark form (pernigra
Cresson of terminalis) nesting in driftwood among
sand dunes on Mad River Beach in Humboldt
The females, which are
County, California.
somewhat polylectic, have a definite preference
for plants of the family Lamiaceae, especially
Stacfuts(Popov, 1951).
Included Species. A. (Clisodon) furcata (Panzer),
sinensis (Wr), terminalls Cresson and xinjiangensis
(W") . Clisodon is a boreal holarctic group which
ranges far into the Arctic both in North America
and the U.S.S.R. Popov (1951) found that the
distribution of terminalis besides being nearctic,
included East to Central Asia and that of furcata
was rnainly European to the Near and Middle
East. In the United States terminalrs is generally
oF THE ANrsoPHoRINn
489
Bnns
found wherever there are northern woodlands,
i.e., in the northern states and in the mountains
southward.
Subgenus
Heliophila
Klug
HeliophilaKlug, 1807, p. 227.
Typ. species: Apis bimaculataPanzer, 1798. Monobasic.
Latreille, 1809, p. 177.
Saropoda
Typ. species:Apis bimaculataPanzer,
1798.Autobasic.
Proposed unnecessarily for Heliophila Klug, 1807,
not HeliophilusMeigen, 1803.
Micranthophora
Cockerell, 1906, p. 66. New Synonym.
Typ. species:Anthophora
curtaProvancher, 1895 ( -A.
squammulosa
Dours, 1869). Original designation.
Diagnoses.Length 6.0-15.0 mm, usually 10 mm
or less in almost all species of Humilis
and
Estebana species groups; female face with pale
markings,. rarely all-bl-agk-ur.-il A.
poste.\rae9or;
rior margin of female hind tibia with simple hair
(FiS. 10d); male of Old World species usually
with 56 and often 55 with thick pad of hair in a
depression (Figs. 30a, 31e), pad covering much of
preapical ventral surface.
Description. a. Flagellar segment 1 of female
equal to length of next 2.0-3.5 segments together;
of male equal to next I.I-2.8 segments. b. Face
with white to yellow markings, rarely black; male
with paraocular and supraclypeal marks present
or absent, labrum of male usually apically trilobed with median lobe emarginate (ur in Fig.
301); scape of male with pale marks anteriorly;
female clypeus usually bearing inverted T-shaped
mark, paraocular marks of female present or
absent, supraclypeal mark of female present or
absent; maxillary palpus 4-6 segmented; labial
palpus 2 or 4 segmented (apical 2 segments
sometimes lost). c. Malar space of female 5-16
times as wide as long, of male 6- 16 times as wide
as long; flabellum apically with several finger-like
lobes; mentum without anterior tooth on basal
third, mentum .6-.7 as long as prementum (.*cept .3 in A. hololeuca);paraglossa about .50-.75
as long as stipes. d-e. Midtarsal and midbasitarsal
brushes of male absent. f. Middistitarsal brush of
male sometimes present. g. Hind basitarsus of
male simple (except flexipes with anterior tooth),
sometimes flattened and/or strongly curved; hind
tibia of male sometimes with apical, anterior
spine; posterior margin of hind tibia of female
with simple hair (FiS. 10d). h. Basitibial plate of
male usually present. i. 56 of male usually apicomedially emarginaje
30a),
Old World
ll
.(FiS.
species usually with dense pad of hair covering
much of basal half of ventral surface, basally
delimited by gradulus (Figs. 10f, 30a), pad occupying large, flat depression; in Old World
species 55 often with similar pad of hair and
depression, often deeply emarginate apically. j.
57 of male (FiS. 30e) not or slightly expanded
apically, emarginate to produced medially with
apex gently rounded; lateral margin with two
'urruq?-I 'T 's,.r^3r^
'Bs pu"
IpJlue^
/s
'y 'f 's,u,arrr
'alnsder
,1 ,11 .sr*arzr
,/I
,gS-tS
.C
.,,\Àer^
.l,I
eprs
pu"
epls
I"rlue^osrop
letruâC
lerlue^
'd 's^Àer^
'/S
'l ,C .sMer^ eprs pu?
,alnsdet
,3 ,g .srr,rerzr
pue
prtue^
lurtue^
BS
I?Jtua^os.rop
l€trueC
'/J '9S-6S 'V 'Dsolnutumbs('H) 'v'l-t
lop1ntmuq ('ù 'V,g-y .eruels Isruos"lâru ârltJo srâqunu eqt
ar" sreqrunN 'sdnorS sercadseueqals[ pu" srpumH Jo sapur 'WU$olpll snue8qns o.totflot1tuy.0g.CIù
c
'b ']urodpl*
ot Flsrp U 11ecpur8reurqns Eur
se Euol s€ g't-t'I
â^l€^ sruad yo xady 'u 'eplm
tsrld
-qol urâ^ luerrncor lsrrd 'eEprr u€rpetu lnoqlr/v\
s€ tuol se ç'B-Z'2, 'paua11eg ^r.ârl lesrop ruor;
'goqs e1etu snldlsouoC'ureldurrs
dgeuors€JJo
ro r{rl^À a1e1d lerpr8dd aletueJ :GOg 'Etg) q}oo}
So
'(tgt 'Etg) pâqolrrl ol pâqoilq dgensn ^Àer^ âpls
otq pe8uolord seturleruos 'padola^ep 11âr\dlensn
âl€ur Jo LL Jo ssacord relnper8 lsarcads uerdo
ur 'paua11eg elerrr Jo alrxocouo8 go xedy 't '(pOg
'ttd
-lqlg ur ]uâsârd dleuorsecco qleel pcrdeqns eroru
luasq€ ot ]uasard vortez\lorâlcs I€urpnl
'(osolnuuonbs pue ou"ta1o"{ -r8uo1 'papunor ro aleln8ue rer{trâ sur8reur pretel
Jo o1ûl dgeuortlppe
-ocrde lluasqe o] Teâ/v\ uorl€urEreura dgeuorseJJo
ur a1e1dprpr8dd) qtaat pcrde o/v\] qlyi'e1eur p LL
'd 'eleur8reura
'ateur8reruâ dgecrde el€tu
'{ 'fogolncoutq
dlpseq
ol
errlue
pue
eplÀ\
se
Euol
Jo BS
se ç'Z-0'I 'papuedxa dlesuq lou (gensn se^lel
ur (tOg 'Elg) cslp 3o qr8uel gI'I ldacxa] csrp Jo
'o 'Âgesropocrde pe/v\er^ eplù\
srued Jo atprrg
qrtual çL'-BZ' euopodeSo qrtuol laldurrs or seqol
Nrrs-rrnfl scNsrcs svsNvx do ^rISusAINn sHJ
06t,
SysrnuATrcs
AND PrryrocENy
recurrent vein joining submarginal cell 2 distal to
midpoint. r. Sting in repose attaining to surpassirg apex of gonostylus, about 2.2-2.7 times as
long as gonostylus. s. Length 6.0-15.0 mm.
Comments. An attractive idea is to separate
Heliophila as a genus from Anthophora. I long
considered this possibility, which could be justified on the basis of small size and somewhat
dissimilar male terminalia. However, the Vestita
species group bridges the gap between the two so
that recognition of Heliophila as a genus would be
highly arbitrary as well as making Anthophora
paraphyletic.
Humilis
Species Group
Description. a. Flagellar segment 1 of female
equal to length of next 2.0-3.0 segments together;
of male equal to next I.l-2.0 segments. b. Face
with white to yellow markings; male with paraocular and supraclypeal marks present, labrum of
male usually apically trilobed with median lobe
emarginate (ut in Fig. 301); scape of male with
female clypeus bearing
pale marks anteriorly;
inverted T-shaped mark, rarely all pale (ascending median arm of mark almost never absent or
indistinct as in an undescribed species from Cape
Province, South Africa), paraocular marks almost
always absent (except in an undescribed species
from Kenya and Tanzania), supraclypeal mark
present; maxillary palpus +-6 segmented; labial
palpus 2 or + segmented (apical 2 segments
sometimes lost). c. Malar space of female 5-13
times as wide as long, of male 6-16 times as wide
as long; paraglossa about .5 as long as stipes. f.
Middistitarsal brush of male sometimes present.
g. Hind basitarsus ôf male simple, sometimes
flattened and/or strongly curved; hind tibia of
male sometimes with apical, anierior spine. h.
Basitibial plate of male usually present. i. 56 of
(FiS.
emarginate
male usually apicomedially
3.0u), but s.ome southern and southeastern African species with apex entire (..g. , some
rufouestita and tetradonta); usually with dense pad of
hair covering much of basal half of ventral surface, basally delimited by gradulus (Figs. 10f,
30a), pad occupying large, flat depression (ptd
absent in some African species); 55 often with
similar pad of hair and depression, often deeply
emarginate apically. j. 57 of male (FiS. 30e) not
or slightly expanded apically, entire medially
(rarely emarginate and then only slightly) with
apex gently rounded; lateral margin with two
lobes, ventral more apical lobe larger and usually
more darkly sclerotized than dorsal one; disc
sometimes with lateral margins lobed midway (as
in some African species) but usually margins
gradually converging toward apex, length of apodeme .32-.6I length of disc [1 . 16 length of disc
(Fig. 30e) in bimaculatal. k. SB of male apically
emarginate, occasionally emargination weak; apicolateral margins either angulate or rounded,
longitudinal scleroti zation usually present (FiS.
30d). l. Apex of gonocoxite of male flattened, in
oF THE ANTHoPHoRINe
BBes
+91
side view bilobed to trilobed (Fig. 30c), in ventral
or dorsal views apicolateral margin either angulate or rounded, apex not projecting inward but
extending somewhat straight distally (Fig. 30b).
m. Gonostylus of male short, from dorsal view
flattened, 3.0-6.0 as long as wide. n. Apex of
penis valve I.3-2.6 as long as wide viewed apicodorsally. o. Bridge of penis valves not basally
expanded, either 1 .0- 1 .5 as long as wide and
basally entire, or brid ge 2.0-2.5 times as long as
wide and sometimes emarginate basally (Fig.
30b). p. Pygidial plate of male present, apically
emarginate, sometimes like that of Lophanthophora
(..g. , fraterna), or with a short tooth situated
laterobasally on margin of pygidial plate (..9. ,
ruJoaestita),or deeply emarginate (FiS. 10f) (..9.,
fuluodimidiata), more often with strong median
ridge; gradular process of T7 of male usually well
developed, sometimes prolonged into a tooth
(FiS. 30f); female pygidial plate almost always
flat, without median ridge [an undescribed species from Peshin Jut (Farah) India has a well
developed ridge]. q. First recurrent vein joining
submarginal cell 2 distal to midpoint. r. Sting in
repose attaining apex of gonostylus, about 2.+
times as long as gonostylus. s. Length 7.5-1 1.5
mm.
Comments. A. bimaculata is typical of a small
group, not
species group within the Humilis
formally recognized here, having distinctly different terminal sterna. The 57 of male bimaculatahas
long apodemes and SB is narrower apically and
more deeply emarginate than subsaharan species
(Figs. 30d, e). The rest of the species of the
Humilis group (mainly African) have 57 with
short apodemes and S8.apically broad with lateral
margins weakly emarginate.
The Humilis species group is a group with
characters between the Estebana
transitional
(: Micranthophora) and the Vestita species groups,
although it is most closely related to the latter
from which it differs by its small size, more
extensive pale markings on the faces of females,
lack of the two median projections on the female
labrum, apicolateral margins of the genital capsule more or less straight and SB of the male with
lateral margin deeply emarginate. The Humilis
group is similar to the Estebana group, being
usually by the synapomorphies
differentiated
(shared with the Vestita group) of the dense pads_
of hair on 56 and 55 in males, lateral margin of
57 of male weakly bilobed (ventral as well as
dorsal lobe present), SB of male with apicolateral
margins produced (not with apex produced as in
sclerotization
the Estebana group), longitudinal
of SB usually present and pygidial plate of female
usually flat (see character p).
Biologlt. The adults are active from March to
November in the Palearctic region and November to August in subsaharan Africa.
Included Species. A. (Heliophila) albella Gussakovsky, albifronella Brooks , albopicta Cockerell,
alfkenella Priesner, appletone Cockerell, bimaculata
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aamla'aserrg ouvxsunnJç'gara
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oToun1o{n^tpue ouazsunDrq
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relnroered !(xocao^t{''E't) TrBIq ge dlerer'(1reru
peua^ul
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t paue^ur Jo ruroJ ul dlerer
Jo xade
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lxeu o1 pnbe âleur Jo
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snDlq! 'gere>1coC Dîunpnr4at'(8.U) qpual 'rausarr4
sllnsJnt 'lsrouag lullos 'rauserr4 nsua"tfloqs'Jeuse
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ny4saao{nl'ua4;1y srcuanqqt,'op1e6-epeetr troaouÊ@
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lmaol'(elourdg)
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'(E.tX) Duuîguoe'lare>1co3 suotfipryuoc'(razue4)
Nrrsl-rnfl sDNsrss svsNvx do Àrrsus^rNn
sHJ
7,6+
SysratrATrcs
AND PHylocENy
oF THE AnTTHoPHoRINB BEns
+93
FIG. 3 1 . Anthophora (Heliophita) bimaeulata. Scale lines (on right side) represent . 1 mm in A, E , F . 0 1
s=seta. A, B, Hind tarsus of female. C, D, Lateroposterior and anterior
mm in B-D; a:arolium,
views of flabellum. E, Ventral view of male metasoma showing the dense pad of hair on 55 and only the
apex of the pad of hair on 56. F, Clypeus of male showing rugose-lacunose sculpturing and
apicomedian emargination.
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'l 'luâsq? uorl€zrlorelJs
pue âroJ pesrpoul dlppo pu? els8uole aql Euraq
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'pepunor dplearvt sercads e\eJ e ur
lseâl eql lou 'selnqrrlle al?tu lBrâ^es ur anbrun
I€urpnlr8uol
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ue sr sa(1xa{.V
, ]nq
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Jo {fua1
pue pepuedxe dgeretel 'peua11eg) aldurs eletu Jo
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Jo
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Jo
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ssoJc€ TJeru esre^su€Jl e ql1,v\ snaddlc '(aou,ozo
ur punoJ sr'sarceds
u€ 'ocna\o\ot1'v
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lueserd dgensn sTretu paddlcerdns
puu qlanleor ldacxa) ruesqe dlpnsn sTreru relnro
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>lreru pedeqs-a peuâ^ur lnoqlr/û dllensn sna
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-aqd pue
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'V ul aldurrs)
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elstuâprq elqrpueur ls8urlreu
ql1,v\ or€d 'q 'slueurtas ç'Z-0'I lxeu o1 lenba
(rer{}e8o}
s}uaurtas ç 'g-Z'Z }xau o1 lenbe
el€tu Jo
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eql râpur. sluau?il7c âr{} w ue^r8 sre}3?reqJ eq} (q
peler€des sr lr qcrq/v\ ruor; dnort srpurnH plro6
plo aql ot pelslâr dlasolc sr pus plroM /v\eN eql
ol pelf,lrlser sr dnorc €uBqelsg er{J 'sruaralaoc
'ruru
ç'tI-$ rptual
's 'snldlsouo8 s€ tuol s€ sârurl
'sn1d1s
7'Z-Z'Z
'r 'dnort
-ouot
Jo xede turure]le asodar ur turlg
srltunH ur se uor]?ue^ 'b 'aleurJsc oslB sur8Jeur
'eleuuec dltuorls o] peua]]?u
[ere]31 serurtaruos
'uerpaur qlyv\ alerueJ
tBrl/v\eruos e8prr leqpnlr8uol
Jo ereld lerp€dd lqtoot e se tueserd rc (n1uopof,t1cn{
pue 'ottonç.tota 'qlataslcocur s€) padolanap Âp1eam
dpo serurleuros '(aotnzo1os pue 'xnuaqd 'olngutîru
'aottnqwnloe 'aotuo.tqow
luâsqe) luesard uelyo âletu
p LLJo sserord repper8 i(osolnurannbs w luesqe)
suorlJeford pouelleg dgerlua^osrop'uerpaurqns
'lecrde o/vrl sB
lueserd el€tu Jo LI Jo ssecord
l(sarceds peqrrrsâpun euo pu€ 'nso1nu
Frprtdd
NIIsT-rng scNsrDS svsNv)
(ntoqdoqruDrztry- ) dnorC
sarcads eu€qelsg
.{o Àrrsus^rN.[I sHJ
+6t
SysreuATrcs
I
AND PsyrocENY
oF THE ANIHoPHoRINB
BBBs
495
I
I
,
I
I
-TroC] spuelsruearo u€rpul eq] '(uanr8 .(e1 prr
-red e '116I 'lara>1coC)elpq Eurlrqeqursarcads
roJ epeur uâeq e^eq (otorldotltuv ur paqrrrsap
dgeur8rro) sercads qTtEauVJo stsrl dldurs ro stol
-BIBO 'o116awou0z
peqrrf,seprer{unJ pu€ opfawy
Jo uorlrusâp peorq e Eursn uorldrrcsâpar poot
e ane8 (çlOt) TculryelT 'pepnlf,ur suorldrrf,sapar
pue sercads /v\eu [ere^os q]lû\ (++OI q orlrld
-oqtuv ur tueql pepnlcq eq) oppatuV uersdepl4l
ol sda>1e,re8 (gçOI 'i+6I) >lrurgârT 'n^toq(oqruV
anrt turaq eruos 'araql petuolaq 11e1ou q8noqr
ntoq{otltuv u\ pepnlcw ueeq dlsnorrrardpeq }€rlt
opfauV ur sorrodsdueur paceld (Oç01) nodo4
'(OgOy)rauaqclry dq ralel pue (tç01) a4elroqull
dq rorre srql o1 pell€o lsrg s€/v\uorlue]lv '(EOO1
)
uosuaqo'U dq u8rat leruoselour aql uo spueq
Fluaun8elur ellq/v\qllù\ sercadsuscrrârrrV qlrop
o^Àlro3 paldope se/v\qlfauV ârrr€u eqJ '(to4stg
'(pg 'ttg) ssacord
l€]eleTsoxâ
padole,râpilâ/û e dluo 'atprrq 1etâlâTsoxâ
ue e^eq
]ou seopsyanfiqulq npfawV ]€r{} sl I atqul (OZOI)
s(âueCJo uorlrârror V '(eg 'tlg) rnds FIql] pulq
reuur pue rrrnrroc eql ruor; rnds 1elqlt pqq relno
eq] Jo uorl€losr (q 'eg 's8tg) elalduror o] (p 'c9
'stt$ prsed aql se^[o^u (OtO1)eue3 l'q puno; ,
ol1fauV epureJ Jo retc€rerlr elqeTr€ruer euo
'(q 'ntt 'rttg) eldurrssr ro
(p 'rtg 's8t,q)(srxe agl punore gs 'e'l) e{rl-urTler
'(J 'r+t'sEt.E)
'e'l)
leldurrs eleru Jo snsrelrseq pulH 'E 'luesqe aptu
'G 'tB 'sttg) saqol eill
J-p
Jo seqsnrq rpr/v\ttlplft
-ra8ug pcrde qlyv\ unileqeg !(eç 'Elg) sreqruerlr
uaurnl rorrolue pu€ rorrelsod qtmt eept lsadrls se
Euol s€ 0Z'l- Lg'g Bssol8ered lurnluaurard se Euol
se çB'-09' runluoru '(ou"to{uo2u0u 'v ur ldecxa)
IZZ orn8rg ur se prlql l€seq uo qtoot rorrelue
qlp\ urntueur ieldurrs dllecrde elerrr Jo tunrqel
:(gg' ot d.) opûaunssol7 ur uorsnrtord ]saluart
Eurqceer lnq algord ur eda Jo qlpy'\ gg'-tt'
'luereqnlord
dlereuat
ot peuoll€lI
sneddlc
!>1ce1qII€ dlarer 's8urlreur l€]uetun8a]ur u/v\orq
ol /v\ollad-qslppor ro arlr{a o} /v\o[ad qlrr* al€ureJ
pue aleur Jo of,€d 'c 'sererrroga8eg 7'Z-I'T lxeu
ot elerrr Jo isereurogateg Z'i-g'Z lxau ;o sqr8ual
pawquror o] pnbo epural Jo I lueu8es rega8elg
'e 'uoq{t.rcsaq
'q 'ruur
lZ-B {}Eua1 dpofl
'(nç 'ttg) suetunl
rorrelsod pue rorrelu€ qlr/v\ eale8 :lZZ ern8rg o1
relrrurs prlrll 1eseq r€eu r{loo1 rolralue olEurs q}lr\À
runtueru :(p-ng 's8l.{) rnds lelql} rauul ruor; patel
-osl qcntu rnds 1elqll pqq râlno :(t 'ctt 'sttg)
ooueJseqnd passardde qlyv\ dgerauot Burossleur
'crgeleur serurleruos Jr€tI ialdturs el€ru
3o s8elprru
:(rO 'Elg) luâsq€ sIIorV 'stsouflotq
pue pqq
eserrt
oTTpEauUsnueC
'llâroTroc
(ary1-retlrea;
s/v\orelrsoddoo/r{.} tutrazzpue llere{lo C DUD?sxTa
qsn'd4srvro{zsope
A oTonsn{t"t1
'a>1crpe11
ur reqtre esorunld sl rler{ aled 'rolor aq} Eurcnpord
sn{ot1t1t1'srepunesruonaoqt'zlrtreJotrNszu.toez.t
']qEII
-rasqns'd>lsno4essnC
ot{}
dlqeurnsard
eq}
uo
satprr
srrerl
tceqglp
suDenuqns'F{oXnnbaoqns'A1sno>1e
'sttd satprr tur8re,ruoc dlpc
-ssnC taoyoîsaqs'garaTroC nsuat{as 'gara>1co3
or{J '(q,'ntg
T-cgg
lqqor
-lde qlyv\ 'paua11eg']uaurntatur eql ol pessardde 'z1rrvrerotr41
nuuot( 'pqcen nn4so{ 'z1rrcr-totr41nuasn4a(
'a1crpe11
'ga.relcoC
'e>1crpa11
ere srreq esarlJ 'rlsrl o8uero pu€ ueert 'a.lq
Dqntro
DuCIunu
sanortaaru
'D.totldotltuV 'pqce6
'z1u*erotr41
'z1vvrero141
t?aDau
onloïuow
Jo sanq cr[e]eru ur peroloc dpq8lrq ew
thtol
'o>luaqcslped tî"taqauo.tq'4ezseueg
orrrun 'ol1fauvJo sarcads
lqDzDI'z1r*tetotr\lsis
-uaqlq'zete1 nsuayqo(ot1aq'
ii#
##î:r|"
o4uer{Js}pedylnwta{'z1rr*e
-rotr^trruasx{'d>lsrrolessnCnsuauotlîta{'11anTJoCDsuaa
'(esarrg) ouDzstnop'1.>lsnolessn9
zanrouasap'z1rr*ero141
suqul?tuap'dlsnoless
nC fl sJvlta
eon' zate4 Depslq'd>lsrvro>1
-zsope1 stsuapaqDqlso'z1rrvrero141
nrqDlqntD'1ara1co3
'tr :pelpnts ]ed se lou ro pedorlsep uââq o^er{
o1cunfqo
Jo
'lsol are sadd] eq] ;o 'sueurrcadsuJo/v\l,laurerlxa Jo/pue
seprual ere saddloloq eql asneoeqsnuetqns ol se dlurel
-rer r1llm paceld âq louuer sercads Euuvrogo; er{I
srogs svrugcNr vuoHdoHINV
eql pue uorEar ueeu€rrellpatr tr aqt 'ecuJv urelsse
ur asorcods dgercodso sr oryflawV 'srlerlsnv pue
eer, uersdeptuopul eql'recsetepentr'sluaurluoJ
plroM plo IIe uo punoJ sr,o1178a?av'uolnqlqslcJ
'o8prr
l€ulpnrr8uol
uerpeur lnoqlr/v\ eletuoJ 5o ereld prpr8dd llueserd
serurloruos a8prr leurpnlr8uol uerpotu'Âgerluan
-osrop pâue{clq} sprrr{}-o^r} ol prrr{} I?sBq ro rl}aet
'qlee} earr{t o}
Jo es€q le eTrl-Jler{s sarur}âruos
o/vu qllù\ dllecrdeorpeu âl€ru Jo LI lluasqe aplu
'ru 'qlplù\ sB tuol se
B'e-9'0
Jo a1e1d prprtd4
'e1nce Jâ^eu 'papunor
dgecrde 'aleu
{fua1
-rEreure o] errtua dleseq sa^F^ sruad 3o etpufl 'l
'aplrvr se tuol
'aosJrns uo pâJetlef,s pue
llzurs eeles
se serur] L'Z-n' f dlpsropocrde pe/v\el^ se xade
e^le^ sruod '{ 'epyv\ se Euol se serurl 0I-1 luaserd
uer{/v\ i(o11ûawnrqaz ur tuasq€ dlerer }nq 'szs
-{ououtltv pve 'r}s 's opûrwv W tsorul€ pue 'rlfï
-aunssolg'o11tflauotelry'oqpflatuouozle'opûawopN)
'relsrlq e dpo ol
]uosq€ dleralduoc saurrloruos
pernper dpearE ue13o eletu 3o snldlsouog 't (gec
-lde pe/r{,oJJeuo} papuedxe pu" 'dgerluen pa^{ell
se pâqolrrt ot aldurs 'poue11eg dgurtua^osrop
ol punlor 'eldurrs (1enr1e1ar â1eur Jo ellxoJouo8
yo xedy 'l 'rslp go çEueI Lg'-+Z' 15 Jo euepodv
'q '(qOt 'Etg) asournld ur8reur rorrelsod
ldacxe
eldturs qtlÀ\ apureJ Jo €lql} pqq
srrer{ pdocs
'peleqelslur aq deur
uaurrcads eql lurnasntrry urlreg eql ur ueuncads
e uo pes€q sr 'ropencg 'pnbedeng ruo{ Dsolrua
's€rnpuoH
-wnnbs 'V
Jo prof,er lcunlsrp al8urs y
o] pre/ûrllnos '(tururodg
turpnlcur) eru-roJrleC ol
pre/vuse/v\ s€xeJ pue 3tuorl3p19 'sesue>I Jo ware
f,rrex ur punoJ are dnor8 eueqelsg âr{l w sarcedg
'llâreTJo1 onauDrlzaozpu€'gere>1coJ DrllqglqruDx
'gara1co3 DpnDetîsn'srnoq Dszlnuupnbs'e4e1raq
-utrrll ?ozrnzD1zs'reueqrlntrxn.tlqtrpltp'(11ara1coD)
xDuaqd 'gara1co3 o1lqdowd 'gara1co3 aouoytn(
' gara>1c
o J Dtuopo{q co{' garelc oJ Dptlâ1u' e1e1.req
-urll
DrJDnUora 'uossar3 suotrtpcout 'gera1co3
ocnaploq'ga-ra1co3 Duozsreun{'uosserC sadqxagf
'pren11 Dleuîcnao{' uosseJC nnîua'lare>1coJ ouoqat
-sa 'rauaqcr141 xnroqlor4l(ta '1are1co3 T â>lelroq
-rulJ aoxnquryot'e>lelraqurll
tllaraltoe'goro>1co3
Nlrs-rrnfl scNsrcs svsNvx do ^TISUSAINO sHJ
96t
SvsrEN,{ATrcs AND PHyr-oGENY oF THE ANTHoPHoRINn
BeBs
497
FIG. 33. Metasomal tergal hairs of Amegilla,dorsal view. Scalelines represent .l mm in A, C, E, and
subinsularisshowing simple red hairs (in life) and a close up
.001 mm in B, D, F. A, B, A. (Gtossamegilla)
of a single hair. C, D, A. (Zonamegitta)cinguliJera,showing the flattened metallic blue-green hair (in the
lower three-fourths) on apical margin of tergum and the sparsely plumose black to pale hairs (in life)
(in upper one-fourth) and a close up of a flattened hair showing oblique ridges. E, F, A. (Zebrarnegilla)
quod*io showing the flattened white hair (middle) and the sparsely plumose hair (top and bottom
margins) and a close up of a flattened hair showing the ridges and marginal teeth.
uzrpeu alturs e qlrm ro (lIS 'ttg) seqol
asnlqo lleurs Jo rred e uea/vueq eleu
-r8renra dy*orr?u dlanrlelar
BS Jo xedz
i(a,roqe se pornsearu eelet) erqrlâroJ
se tuol se serurl g'Z fsour 1" 'exocpnu
Jo etpe JorJâlue ol lsoru le esodar
ur Eurqceer 'gl8ue1 elereporu Jo eepC
o141âawosso4g
' (rlnq a8uero qU^
JIer{ rorra}uz }wel
te ro rler{ >pelq IIE r1tlm e8rat pruos
-31eru rreqt pu? r{}Eue1 w IrIIII g I }s?el
lv, eJ? r{Jlr{,v\ sarJads usrpul eeJql er€
arâq}) u"rs/t"Furopul Âpreur'sarceds
'ttg) seqol esnlqo a8.re1
Ieluorro :Ggg
Jo rred e uea/vqâq elsurtrerua BS Jo
xede l(sndpd dregrxeru ot xade tuory
pernsearu zap8) "rqrleroJ s? Euol se
seurrl t lwel ]e 'exocplur Jo elpprur ol
'Euo1 eoleC
ls"el 1e turqJeer esodâr ur
(l 'J 'egg
0I
's8tg) a8epuadde ate8uole ue ol ralsrlq
v, ruor; turdrerr '}uasard snll;souoC
' '(qgg 'JLg'sEl.D
tuesqe snldlsouoC
6
DII!îautDtqaZ' '
sarreds
usrPul o/v\] dpo qlP\ dnor8 u?3rJJv
dpruur lurur il-B {fua1 dpoq :(>1ce1q
ser{ qrlqa
serreds ueu€r{C
IIe IJ
e roJ ldacxa) rler{ Jo spueq pcrde
epd qtlÀ\ etra] Izrrroseleu il" :(pçg
'Etg) tuol se peorq se
'd1eru
tnoq"
-rou pepu"dxâ dlerelel ZS Jo xady
'6
'B
'r!!!'.*.':':rg
'rorurJ 'elpul 'ecg;y ]see
-rluou lurru BI-?I gfual dpoq lleq;o
spueq pcrdz a1edql1,v\tJ puz ga Âpo
:(tçg 'Etg) Euol uer{} rapeorq r{f,ntu
'd1pre1elpopuzdxa dlreert
/S Jo xedy
' (rln.{ e}lq/v\q}l^À
ollfauuuoz
slpul ur serJedseuo ldecxa) e8uero
Jo uoaJ8 'anlq crllelâur sdervrlz]sotule
etrat uo JIsr{ Ie{uos"leur lpaluaurtas
-9 sndled dregrxeru :(t tg 'ttg) lues
-erd eere r"lnJrrc lzf,rdeqnsleretel r{tlm
'(J 'nLg
ZS leprrvrsu Euol se tsoru te
'stt.l)
e
pernpor
retsrlq
ot
snldlsouog
npo(onsv '
rrlptetu
Je^eu 'rlut8 Jo >lJslq 'urvrorqe8ral 1ztu
-oseleru uo JIzr{ lpaluaur8as-g dlarzr
ro -ç sndpd drepxeur i("lOg'Etd eere
relnJrrJ pcrdzqns Ieralel lnoqll/v\ ZS
ieprrvrse Euol se seurrl aerr{t }seol}B '(E
'q6g 's8tg) padola^ep
[elvr snldlsouoC
'L
' sârrâds
L
IBluâlrO
rvrâJ3 tlll/v\ sdnor8 uzldolqlg ue Âpreru
lanlq Jrlplerrr ro^eu JIer{ I€ruoseteur
Jo rolor larrtua çS Jo urtreur 1erldv
'usrselert
g
-snv i(adocsorf,rur e repun elqrsrn l.1uo
uego) roloc ânlq crlpleur qlri* e8.re1
,errlue ,e1eu
Ftuos?lâru uo rrerl ueql
51
'ç
-rtreura dpeorq çS Jo urtreur pcldy
npfauqoN
dnorE u"rlertsnv
:(rgg'ttg) ]redse lertue^ ur paqo[q
dl8uorls BS Jo aseq :(peg 'Etg) qlool
'reln8uelrt Erl
Ieralel qtlm aurapode
-readde 'pepuedxadl8uorls 4SJo xâdv
'rls 's nl1fautv ' '
' dnort ptuerro pue rrtrrea1ud 'uerd
-oFltg :(tçg 'Etg) pelwod dlecrde 'tld
-ulls BSJo aseq :(qçg'ttg) qloor 1erelel
lnoqlr/v\aurepode'elerpenb turreedde
't
'pepuzdxa
dpo
dlalerepou
1SJo xady
e8epuedde
ç
atu8uola ue ot relsrlq e se (ruur 6-8 ere
sqÉual dpoq esoq/v\Dleutenaoru
uerpul
pue fuq 'V ueydoglg eql ur dpo ]ues
-qe) tuasard snldlsouot :(tOg 'tLt '{
'cAt 'sttg)
l,ltuor]s
peqoflq
BS;o xedy
all
,
-xoJouo8 uo relsllq " o] poJnPer sn1d1s
-ouo8 pue urur ZI twel le qr8uel dpoq
uegt luesard 5r 'luesqe sdurvrp]soruls
snldlsouo8 :(tgg 'lçg 's8t.l) paqolq
't
Lplzar* dran ro pepunor BS Jo xady
(t 'e 'cAt 'sttg) lred
B
-s3 [?srop Jo l?r]ue^ tuoJJ rep8us}Jar
turreadde runurets 'pacnpord d11el
-peuocrde saurrleuros'altJua Jo e]"ur8
-r?ure dplprruocrde 'apyo BSJo xady
(rt 'l 'cçt 'stt.f) rced
t
-s3 Issrop ro l"rtue^ ruoJJ ælntueFt
sreadde runuJeJs ]etll r{Jns pepunoJ
'z
ro peqoilq rârllre 'porvrousuBSJo xady
' ' (sapureJ)'sluaur8aspuuatuu
Z,l
a^[o/vu le8rat Isruos"lâur pasodxe 4s
'(sapur) 'sluarutas
puuotue ueet
Z
.I*rl'i:ïJ;:îii:ïff'":Ë
r
'(eço
I
'9
'1curgar1) (o"rotldotltuVânrl rllyr\) spuelsl dreu
-BC eql pue (OOO1'esarrg) eeurnC /v\eN '(ç96t
'626y 'garaTroC) Brl€rlsnv
'raueqcr141
gO
:ql
I
'ernssnes lztOl 'goro
'[(Âpo recse8epel
l) 0681
NIIs-rTnfl scNsrcs svsNvx .{o Àrrsus^rN^[I sHJ
B6?
SvsreuATrcs
AND PHvrocENrr
oF THE ANruopFIoRrNn
Bnns
+99
FIG. 34. Metasomal tergal hairs, dorsal view. Scalelines represent.1 mm in A, C, E, and .001 mm in
showing flattened hairs (white in life) in lower twoB, D, F. A, B, Amegilla(Dizonamegilla)sesquicincta,
thirds of Figure A and round hairs (black in life) in upper third and a close-up of a flattened hair
showing oblique ridges in Figure B. C, D, Amegilla (Zebramegilla)albigena,showing white round hairs
the shape of wheat tasslesand a close-up of a single hair showing the longitudinal ridges. E, F,
Anthophora(Heliophik) bimaculata,showing plumose (ochraceous in life) and simple (black or pale in life)
hair and a close-up of same.
-oseterulurur L'Zl tseal]e qr8ual Lpog
' '
(ued u\) D11!flaunrn747
' slueur8asg't-ç'f,
lxau Jo sr{}Euel
pewqtuor o] pnbe I âreurolateg
Jo {fua1 lurrr g'l }soru }" q}plm leru
-os"teru lurur ç'ZI tsoru ]u ql8uel dpog '02,
"
"
(ued u1) s1s(ouoraslev
' 'pr5lr pcrd" o] e1e1d
1elqltlszq uro{
l,gelsrp Eurpuatxesrrer{pdors >lf,elqJo
pu"q qll^À elql] pqq yo a8pe rorrelsod
srrerl pdocs Jo pu"q TtBIq lno
07,
-qrp\ e]$I,v\slqll pu5{ Jo e8perorrelsod ' 6 I
eJuacseqnda8uero ro rvrolad
lZ
'snoeJeJr{Jo'a}lga passerdde 'asuep
qlp\ pere^oc etral II3 ro ourosJo uorl
-rod elqrsh r{}1,v\rolpue rlerl Jo spueq
pcrde ]noqlr/v\ ut-ral ["ruos"leur etuos
' snoecerr{Joof e}n{/v\'rreq
6I
3o
.BI
spu"q pcrde qllm etret leruos"reur [V
opo(onsv '
1rd prrotual rorrelu"
le >[r"ur {r"p geurs roJ ]dacxa e1ed
dlelalduroJ snaddlc lsarcedsuer["r]snv
BI
'llg
Trelq [1ersoruleol gz snedllc ro (l
'sEtg)
,J, palre^ur Jo ruroJ ur Euqr?ur
plueun8alur e1ed qllr\À snaddlc lserc
-ads lstuerro ro 'crlcreepd 'uerdo5{lg .LT
B
Z
"
sluaur8esxrs rlllÀ\ sndpd dregrxeru :(t
'1Lt 'stl.l) tuaserd sTreru rzlnroered
' '
' '
'SlUaUI
LI
-tas x1s ro a^S qllÀ\ sndpd dregrxetu
'9r
:(pOg 'ttg) tuasqz s>[rerur"lnroeted
' '
'uerlerlsnv
(ued uù DIIlEaunuoT
pue uersde1etuopul 'uerpul 'uursy
lffieqtnos lsluanrtas B'Z-t'2, txau
eql Jo sqrEualpeulqruor eql ol pnba I
oreruole8eg lerqrlaloJ s" tuol se serur]
'exocplur
ç'7, {}Eua1
Jo a8pa rorre}sod
l$sedls"el le turqcear asodar ur "ept
isluaurEos 4s rlllÀ\ sndpd drepxe141
' '
oppauopN
dnor8
uerl"rlsnv lstueurtas I ' t-0't
lxeu
eqt Jo sqÉual Peqqruor egl or Fnbâ
I erâruolateg Jo r{l8ua1 ierqr}aro;
se Euol s" eJr/vulnoqe gl8ual 'exocprru
Jo a8pa JorJelue o] lsoru Ie Eurqcuer
asodar ur eeFE lstuau8as e^$ d11e
'çI
-uors?rro 'rno;
sndpd
drepxetr41
qllÀ\
(ued uù DnIîauo^{V' '
sdnort
ueJr1;V lluasqe s>lreru re[ncoercd
sdnor8 leluârro ro
çI
uerprlsnv ltuaserd sTreru relnJoered
f*1.f q.).Dl1Iîawnsso17
'+r
Iluasqe
*nrn-rrrr,ro*î?(-îiifltîi
-ilrx"ru ol xade tuor; pernseeru eapt)
"rqrteroJ Jo rptual sarurt g lnoqe qr8uel
'exor pulq;o e8pe rorrâlsod
ol elpprur
ruor; asoder ur Eurqcuer 'tuo1 "e1uC
sdnor8 ueJrr
,l
-JV ro 1eluelro 'uerlerlsnv
iluesqe
ro luesa.rd s>[r"ru relnJoered :(sndpd
dregrxerrr ol xade tuo.r; parnszaru Ba[
-st) "rqrleJoJ
Jo t{}Eua1 seurr} g'Z lsour
te r{}tua1 'exor pulrl Jo etpe rorrelue
ol lsoru te asoder ur turqceer 'qrtual
ur tror{s ot et"reporu dlpnsn eapC
' >lrelq ro ,(er8 'moged
9I
'a8uero 'urrnorq 'snoaJ?Jgoo o1 erplrvr
'orgelaru
lou z8"ral Ftuoselaur Jo JrpH
gl
atuero
-par Jo enlq 'uear8-enlq 'uaer8 '(adocs
-oJJrru e Jepun âlqrsrrr dpo sarurlauros)
rrlpletu
e8rat Ieruoselaur Jo rleH
n1773auofla74J''']uesqe a1z1dFlqllrseq
:(ngg 'ttg) aqol lnoqlrm aldurrs ur8reru
reuur 'papunor pue papuedxa lcadse
Frlue^ tuor; a}rxorouo8 yo xade :("gg
'Eld
dlelsrp
tur8"ra^lp
sprrq]-oml
ot JIeq lecrdz 5o sur8r€ur Fra]el r{}lm BS
'luasard
nppaun{y "
serurteuros a1e1d plqllrseq :(lgg 'ttg)
ur8reur Jâuur uo eqol 11"urs rl]lrvr lcadse
prlua^ tuog âtrxorouo8 3o xede :(tgg
'Etg) dlulsrp Eur8ra^uor sprrgl-omr
ot Jlerl pcrde go sur8reur I"retBI rltlÀ\ BS
'(rgg 'Etg)
ns{ououtyev "
aruereqnlord ]q811sqrlm dppeurocrde
ro arrlue ur8reur pcrdu qly/t BS :(qgg
'tt"f) qloo]
Frel"l padola^ep llarn r{}lÀ\
Jslp Jo uorloâsplru 'uorsu?dxa pJalel
padola^op IIal* qllÀ\ 'apyn ZS Jo xedy
'tI
'zr
.II
' '
(t'cgt
II
's8tg) seqol uerpetuqns esnlqo yo rred
uee/vuequorlsur8reura pcrde qllÀ\ BS
:(t 'pgt 'stt.{) qtoo} 1eral"l tq811sq}lÀ\
ro tnoqlr/v\ Jslp Jo uorlf,ssplul 'uors
-uedxa l€rol"l lerurou r{}lm 15 Jo xedy ' 0 I
qlpauz"ttxry ' '
' (rp.rsq
l"rlde apd qllÀ\ e8rat
Ieruoseleurpue {}Eua1 ur urur ZI tsoru
le âr? qcII^À sarcadsuzrpul oml are
erer{}) sarcadsuerdorq}g dpreur laqol
NrrsT-rnfl scNsrcs svsNvx do ^rrsus^rNn
sHJ
00ç
oF rHe ANrHopnonrNB BBes
Svsreltetrcs lr.lo PHYT,ocENY
2r.
22.
23.
24.
2s.
501
mal width at least 5.0 mm; length of
flagellomere I equal to combined
lengths of next 3.0-3.3 segments
. . . . Amcgillas. str. (in part)
hair with some black intermixed or
black and white intermixed . . . . . .
. . . Atrugilla(in part)
26. MetasomaeitherwithT4coveredwith
Pubescence of thorax brown to orange
(sometimes with scattered dark hair);
metasomal hairs usually almost atl
black, sometimes visible portions of
terga with appressed white hair from
. 22
apical margin of T3-T5
ochrawhite,
thorax
of
Pubescence
ceous, brown, black, or black and yellow (sometimes with scattered dark
hair); metasomal terga with aPical
bands of hair and/or appressed hair on
visible portions of T1-T5 or if T1-T3
(excluditg apical margin of T3) all
black, then scutum with white hair,
23
black intermixed. .
Maxill ary palpus with five segments;
body length at least 16 mm; length of
flagellomere 1 equal to the combined
lengths of next 3.0-+.2 segments
. Megamegilla (in Part)
Maxill ary palpus with six segments;
body length at most 14 mm; length of
flagellomere 1 equal to the combined
lengths of next 3.0 segments.
white hair, T5 with lateral Patch of
white hair and remainittg terga black
or with T1 or T2-5 with aPPressed
ochraceous to orange hair, not with
distinct apical bands of hair though hair
sometimes gradually becomes denser
27
near apical tergal margins
Metasomal T2-T5 with apical bands of
white hair, without appressed pale hair
on visible portions of terga.
. Zebramegilla (in Part)
Maxill ary palpus with frve segments;
pale mark on clypeus reduced so that
inverted T-shaped mark is small,
sometimes only represented by a meline; supraclypeal
dian longitudinal
mark small, almost absent
. Megamegilla (in Part)
. .
Maxill ary palpus with six segments;
inverted T-shaped pale mark of clypeus well developed; supraclypeal mark
well developed (Figs . 43f , l)
27 .
r;;; ," â!rK:#u:,'';^,|,ï?
28.3p..i.,
;; ";i#::'$::tî:tii]
P#";.,i.
eus with inverted T-shaped mark or
reduced to a median longitudinal line,
if clypeus all black then species from
. 2+
the Cape Verde Islands . .
Oriental species (Indian region); clypeus all black or with small mark
basomedially . . Glossamegilla(in part)
Scopal hair on outer surface of hind
tibia all white (sometimes with a small
patch of dark hair near basitibial plate)
25
or all orange
Scopal hair on outer surface of hind
. 26
tibia all black.
Length of flagellomere 1 equal to combined lengths of next 2.5-3.0 segments; scutum with all white to
ochraceous hair, no black intermixed
or species almost all black with orange
. Micramegilla (in Part)
scopa
Length of flagellomere 1 equal to combined lengths of next 3.+-3.6 segments; scutum with brown to orange
29.
30.
31.
Palearctic regions (East of 65th merid29
ian) . .
Species found in Ethiopian or Western
Palearctic regions (West of 65th merid..33
ian)..
T1 and T2 all black with no pale apical
bands or appressed brown hair (T2
sometimes with white Patch of hair
30
laterally) .
T1 and T2 with pale aPical bands of
hair (sometimes weakly developed on
T2) or most of visible portions of T1
and T2 with appressed brown hair
(sometimes very sparse or absent me31
. .
dially)
a
except
all
black
terga
Metasomal
patch of white hair laterallY on T5;
propodeum with black hair
Glossamegilla (in Part)
. .
T3 with aPical
sometimes
T+ and
bands of white hair; proPodeum with
Dizonamegilla (in part)
white hair. .
Length at most 9 mm; thorax Prewith white hair, some
dominately
'sÀ\âr^
'4J pue
'p 'mara aprs 'slnsdec
'y41'smarzrpJlue^ ./S pre
,.I
["Jlue^
9g
1etrue3
BS
';1 'rr,ror,r
'alnsdec
'f 'snsrzr
'gS
,I ,H .sÀ\ar^
pue
aprs
pue
pJlue^osrop
lerluâ^osrop
lerrueC
l"rluâ^
/S
'elnsder
'C 't 'peaH 'g 'sr*erzr
'/S pue
,C ,C .smâr^ âprs pu"
l€rrueC
lsJlrrâ^
BS
l"rluo^osJop
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'g 'V 'epur 'o1tuutnbsas
('d 'V'N-f :g ldecxe apur lp 'WoltnSupab ('ù .V ,1g,
l€lruâC
'.a1eu 'oua&q1o
('à 'V'q-y 'o1pântouonepue ''4s'sq1tSawy'rprîawo.tqa7etaua8qnson Sr*V.çg .CIt
e o1 pernpâr lnq (ruesqe TEuq 'V u€rrr;V eql
e sB Eurrrras pue uoxel palela-r B ol perlddu erueu
eql 'opifla747*td 'ecg;y yo à.rtnbe padirls aq] roJ
ldacxe) luosard âletu yo snldlsouo8 i(q 'nçg 'sEt.E)
popunoJ ol oleJunJlqns
ur8reur pcrdz Jeuur
D"rqaz'fo1ou(tg
eru€u uerurssdqV'(crrequry)
'prervrul
'paua11eg dllerlua^osrop
(n4o1ncncvl
e1etu Jo
lueq
elrxorouot 3o xede :(rçg 'tld dgelsrp Eur8ranuoc
errlue ldacxe) eleurtrerrre l.1pseq 'eplar se Euol se
'parvrorreu
sarurl g'e-0'I se^F,r sruadSo o8pufl 'l 'e€les perâl
JI"q ol prltll lelsrp uo surtr€ur lero]el
-leJs 'geus qlyv\ 'peorq se Euol sB serull L'Z-y'Z
dgzcrde eleru Jo BS Jo xade :(pçg 'Etd luasard
eete JelnJJrJ uerpâtuqns 'a1e1n8ue 01 papunor
e^le^ sruad ;o xedy 'T 'relsrlq ? s€ (ruesqe funl
'peqopr] ro pepunor pu€
ur8reu
ur) luasard apur
snldlsouoC 'l '(q 'eçg'stl.E)
1era]Blocrde
Jo
arrluâ oletuJo /S3o xede laleurtrcura (yvrolegs ol
aterunrlqns'peue11eg dllerluo^osrop dllq8rls dpo
'lunlg
'atre1 serurlaruos 'prervrur alEue aertap
'stsouflotg
errlua eleru Jo gS lurur ?I-B qltual
06 e lsorule le lueq urEreur Jauur uo eleur Jo
elrxorouo8 yo xady 'l '(Tcetq IIe IJ qll^À sasoqxuD
ldacxa) spusq pcrde elaldruoc r{}r/v\ gI-1a ierrlue
setulleulos aqol uelpetu qly/t ue{o qlâel ueâ^{.leq
'dgenua^os
'qlaal
(ued uù Dlllflaunuoztg
o^r]
ur8reur
qllÀ\
lecrde
'(nnlawos 'V)
-roP pauo{3lr{} sPrrr{}-o/vu o} prFI}
lurnlncs
qll^À dlln
11telq IF 7J-IJ
I€seq
-nsn alelnqer dlerpeurocrde eletu Jo LL'q '(tçg
uo rlsr{ Tr€lq r{}l/v\ PeKru e}lrlM
'Etg)
âruereqnlord
IerelBI Tee/û r{}1,v\sprnl}-o/v\}
ol JIeq pseq uo 'dgetsrp turEranuor JI€t{ o} prn{}
' '(nTcuxtorîD'V)
Eurreed
pcrde uo surtreur Ierel€l :(peqolrpenb
gJ-tJJo suoruod alqrsr^ pue 7a3o urt
-de
dilqpaurocrde'parvrorreu
pâqo11q_
.szsnqt"rlo)
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eleru Jo BS yo xady 'E '(pçg 'Etg) rslp;o çEue1
euepode
lluasard uorl€zl1
çç'-gt'
Jo
çtue1
f*y.f
yt.):r!!8aun^tqa7
-orelJs l€ulpnlr8uol lecuereqnlord leratel t{tl1v\ro
' rl"r{ apd
Jo spueq pcrde
aldurrs JIBII leseq uo urtreru [€ra1e1 ieqol [er]uâ^
lurur
ql1,t\
çJ-II
il lsoru te r{18ue1
punor e pu€ eqol Iesrop (aleurrec dlerer) punor e
'rleq aled
r{}y/t peqo[q serur]eruos 'a1e1n5ue Jo punor rer{}Ie
+t
'{celq
dgeralel pâpuedxa rslp Jo urtreur Isretelocrde
Jo spueq pcrde tnoqlr/v\
IF Z,f
ileqJo tlcled a8rel € qllû\ aldnrrs Âgensn ]nq qlxrs
pue IJ ]see1w :ruru çI ls"el te qltueT 'gt
pcrdz uo eloreqnl uerpaur e ro setprr Isurpnlr8uol
(uzd uù zn!îaun"rqa7
u€Ipeluqns o/v$ I{}lù\ setul}etuos f,slp Jo uorilod
papuedxe Jo elpplur i(nsoqu,p ul ldecxa) luesard
eote JulnoJrc 'luaserd ueryo €urJsc pcrdeqns IeJa
-lB[ '(uorleur8reura peorq r{tyv\ peqo[q sxlaDqnw
Ie^al /v\oleq IIefft >[r?ur Jo al8uz r?lnf,o
ldecxa) euo u€rpetu Jo epls rlree uo eqol atnce
'leurrou >[rzru relnJoeted ldlrorre]ug
dlleervr o] lq8rerls uttreur
IIBurs qlp\ paqolrr
u/v\orq r{rl,v\ edecg
Tr"Iu plueun8alur
pcrde 'errlue IS epur 3o xedy '3 'Q7uq ur drunq
'
(ued u\) un!îaunuoT
(oTcutcoaazu
posr€r rlllû\ urtreur lera]elocrde) prlt{t lecrde uo
'Eld al8ue paddlc se
usrpeur'uorssardtp
atprr
lecrdeqns
leurpnlrtuol
ù (rtZg
lo^el
'(oalntaoeqnspue '1ïuo1
ldecxe)
lerelel qlp\ rslp
âur?s dlreau w >lJeur Jo altue JelnJo
'aleur8retue
l.gecrde
dlmogegs
erlluâ
ol
leg rsrp
'aEre1
Treru relnf,oeted ll,lrorrelue Erl
-orpetu âleru Jo gS 'â 'eleru ur luaserd ereld l€lqll
'28
-TJeru leluetun8alur r*olad ql!v\ edecg
-lseg 'p '(sdacryuo6 'V vr. pel"rsnJur) realc s8uunr
laleura; pu€ âletu ur ]uaserd sTJetu relncoered
(ued u\) DllI&auossolg
'stur>1retu r*olled or ellr{À\ I{}T.,v\
âJuJ lpaluaurtas-7
spuuq
olq
Eursuapuor
4gecrde
sndled dregrxeur
pue pelueutas-g
sndpd
'e8ra1 go suoruod elqrsn
IBIq€l
IF ro aruos
lexocprur Jo etpa rorrelue ol exocoroJ Jo e8pe
uo rler{ umorq passarddu qtlm dlln
pulq Eururelle asodâr uI eep8 lsadrts se Euol se
-nsn 'r1e.I u/v\oJq Jo spu€q Forde qtrrur
lnoqe essoltered lurnluâurard se Euol se L' ]noqe
utrrnluetrtr 'r 'reqta8ol stueurtes ç'7,-t'l lxâu Jo
utre] Ftuosetâur
!pexruuâtur TJelq
'raqleEot slueur8es
0'1,-ç'Z
çEua1 o1 lenba al€ru 3o
rlllÀ\ serurleruos 'rle,{ peJ rlsrumorq
txâu Jo tll8ual ol lenbe elerueJ Jo 1 luaurtes
r{lp1,v\xsror{} luur U tseel te tp8ual
rela8elg 'q 'rutu +I-B qr8ual 'e 'uo4{ulsaq
'(>1ce1qp
rler{ epd Jo spueq 1erlde qrrm
Zt
II qlp\ szsaqulv ldacxe)
>lJulq
e8ret Ieruoseleur !paxruuelur
rler{Jo spusq lecrde olalduror r{tl/v\ çJ-ya lratsrlq
addl
*B''"'l'i'iiil
:Ï'::""îl;::i";;;,^'
r*1.f:t.).Dt!!l,y?'rry
.
;,;*:,; v)(r'ttn
t,ri {;:::'::f^Ëi:
Nrrs-r-rng scNsrcs svsNvx do Àrrsus^INn sHJ
209
SysreNaATrcs AND PHyrocENY
oF THE ANrHoPHoRrNe
Bees
503
-lquoc ^{.eu (11ara>1coC)ocyo71a,uorleurqtuor /v\eu
(qlp1trrS)osn{uoc'uorleurqruoo /v\eu (esarrg) nsua(oc
'(,{ll*S)
suo"rrtuoc(o11fawù 'V 'sapa{g paprytul
-nrqed
ruo';dsslpp'err}'rrqa*iilfitiËl'fii
eqt uI 'llrdv ol reqruoraq tuor; o^r}re ere stlnpe
eql Brlr;V ure}sea pue ureq}nos uI '6o7o19
Josrequeqeruoso1da1e anes
i:{:ii'
frràil
'oppauyo^tq?Zrreql ra8rel
ere .rls .s o11ûawV
'ereuatqns o/v\l eql
uee/vuaq delrerro ezrs
lsoru
e sl erer{t q8noqt ue^g 'xade s}r uo rler{ yo qcled
u"rparu asuop e turrreq IS Lq pue turpnrlord pue
aletuole euepode stl '(lçg 'std dgecrde pepunor
pu? pe/v\orr?u se dlreau tou Sureq BS eql dq
paleredes ers sa1eru eql qJrq/v\ ruorJ o11ûauotQaZot
pel"ler dlasolc lso(u srrls 's oyûauv 'sruazauoC
'errlua ol o]"urtreura l,lpuorq
.(1psuq 'a_plal se tuol sB serur] 9.T-Z.T se^F^
srued;o_atpug 'l 'luaserd a€tas IIBurs ,poqs idrct
-ropoJrde parvrer^s€ eplrvr se 8uo1 se serurl
I.Z-r.I
e^le^ sruad Jo xedy 'T '(ralsllq e o1 pernper
'lueserd snldlsouo8 nenalo"rqeo
pu€ nsua(oc
]nq
ldecxe) ]uasard ruls eelas eTrl-reldtsouo8 qtyr\
'luesqe dpuaredde ol"ru
Jo snld}souog
{pryn
'f '(E 'lçg 's8t.{) ereJrns
uo asredi
ler}uenocrde
ol esryord rreq 'algord w pa/v\er^ sB tuesq€ ol ]ues
,1ced
-ard aJBJJns pJluel Ja/v\o[
Jo eJueraqnlord
-s3 Isrlue^ ruo{ pâ/v\er^ se l,lprerv\}no alep8ue
]erlÀ\etuos prSI] pcrde go ur8reur Fre]el ,elgord
w peqo[q aprrr Jo e]rxocouo8 3o xady 'r 'rrlle]etu
'rleq
lou
Jo spueq pcrde aleldruoc qtyv\ etrel
eletu Jo LL
Ieutuos€leru ipaqopq dlprpeulocrde
'q '(tçg 'tlg)
pcrde 3o urEreur
tqtrerls lsorule
'paqo11q dppem dranJIer{
serur]aruos ,punor
IerelBI
dgecrde ssel ro erorrr epur Jo BS 'E '(qçg .tlg)
rsrp Jo rp8ual ffi'-gt'
euepode 5o {}Eue1 ,uor1
-ezrlorelrs
leurpnlrtuol qlp\ cslp iqtool uor{s q}1,v\
JIBI{ FsBq Jo urtrenr lerelel l(nsua{oc ur luasqe)
eete relncrrr pezrlorelrs dpq8rl pcrdeqns qlyv\
dgerolel xade leqol palcalord dlere]elosrop re]no
ot lerlue^ aqol paonpord dplee/v\ aroru reuur , (r.rt
-ua{oc w eqol elturs) peqo[q dplear"r dlleralelocrde
'rleq
Jo qrl"d uerpaurocrde qlyv\ pue (eleur8rerue
dlrvroleqs lnq dpeorq osn{uoc pue o1n"t^tofl)err}ua
dlpcrde epur Jo ZS 'J '(ocueraqnlord uerparuor
-lde
IIBurs e qlyv\ osn{uoc pue o1n^t^roi)dgerpaur
-ocrde eleur8r€rua dy*ogeqs pue dy*orreu apru
'e 'el€ld
'p '(orotcso{t"t
Jo 9S
l"lqplsgq qll^À aletr\tr
-ponb euros ur pe]ecsn;ur) realc sSuvrr ipedole^ep
'luesqe eFureJ
[Ia/u\ elsur Jo
Jo sTrsru rslnJoered
'(nsua(oc 'V ul rvrogad) stuqreru
elrr{/v\ rl}l^À eceJ
lpaluanrtas-7 sndpd plqel lpelueurtas-g sndpd
drepxeur lexocp1ur Jo elppltu ol a8pe rorrelue tur
-ur"tls esoder ur eap8 lsadrls se tuol sB
00'I-1g.0
essolEered lurnlueurard s? Euol se çB'-Z,L' runl
-uetr\tr'c 'raqtatot sluaur8es
B'I-2,'J lxau 3o qfual
o1 lenbo eptu ;o 'raqlatol sluetutas 9'g-0'S lxau
yo çtuel
ot pnba eleureJ Jo I tuetu8es rega8elg
'q 'rulu g'9I-0'I
'e 'uoty(ucsag
I qr8ual
nsua(oc 'V u\.) luesqe a1eru;o snldlsouo8 lagord
ur uoes sB dlpclde peqogq dpleam e1etuJo all
-xocouo8 :($S.'Elg) pepunor ssalro eroru dlfecrdri
'81.f) eete r€lncrrr pazrlorelrs
fp.i8l
BS i(Use
pcrdeqns
r{}yv\ (uoqfitcsaq aàs) airiuS
leratel
dlpcrde /S iurur ç.9I-0.I f qi8ual -s1s6ûo1g
'uorleu8rsâp
[Bur
-EFo'
o;,;}!t,
68L1'srasrn,r,:';;i"rt{K
:rtTJïï
'LL7,'d 'eIt6I '1ara:1coC
dq pel€u
-Ersep'6811 'sra1p1 ep Dfntso{tqonbsldv:sercadsaddl
'd '1,681'asarrg ol1pauV
BI
'Jls 's eserJg oTpEauV snuatqng
urâr{}noso} uortar u-?euerro}rpetrN
eq} ;ilr}
lnoq8norql punoJ sr dnor8 etrel sHI .uor]Bu
-lqutr.Io?
/v\eu(1era1coc)aÛzuuzz
pus ,uorleurqtuor
zuor}
1v\ou (11ara>1coC) 4s{tapuoa '(zen6)
s1^to7o7-"wqutrroc
/v\eu
(ge-ralcoC)
oat^ta{o^@nr,(rer1e1eda1)
Dalruaoeqns.'(uoraueC) ouoTso1t(s,(rarleladri)
t(ufuans'(z1r*retory) aoxalps,(lara1coC) sioruqit
'uorleulquroc /v\eu (lere>1coC)
o1o^tponb,(ra>1p6)
suotrttcund 'uorleulquror
/v\eu (reusorr4) olsiwo
'uorleurqtuor
/v\eu
(gare>1coC) ylapfio,(aserrg)
s(anmcsqo'uorleulquroJ
/v\eu (lera>1coC) ou,iqnû
'uorleulqrrroc ^reu
(esarrg) ns.toTttùru,uorleurq
-ruoo À\eu (garalcoC) stlanrlezw,uorleurqruoJ /v\ou
-rvreu
(garelcoC) Dluopouqau'uorleurquoc
(11ere
-rpo3) DraeouDlara'uorleulquoc /v\eu (gara>1cô3)
o{n.toryaw'uorleulqruor
/v\âu (arnssneg) rssinpow
'uorleurquoJ
petwolttol (uoll
/v\eu
(gera>1coC)
,uorleurqruoJ /v\eu
-surqluoc /v\eu (lera>1coc)
18uo7
'(ue4JIV) pltUotl ,uorleurquor
IlereTro J Dexsowrotl
^au. (aserrd) sdanpuoÊ 'uorleurqtuo)
/v\ou ir.rl
-JIV) pcaot7 'uorleulquror
/r{,eu (gare>1coC) o1cuti
-oua{'(eserrg) rrnr{'(qll*S)
*ttr{,uorràurquroc'
/v\e_u(esarrg) Dut4ua'uorleurqtuoc /v\eu (11are>1coC)
oy{nrslp'uorleurqtuor
/v\au (garelcod)
s1iuao1o1
-!p'uorleurqtuoc
/v\eu (E"U) CIalzre,(snrcrrqeg)
ol.Dqntrzt 'uorleurqruoo
^reu (11ara1coC) s!îoanc
'(rerlaladrt)
sualut'(11era1cod) fraonbaq'iuort
-eurquroc
1.eu (gere>1coC) szsuauontltaq,uorleurq
-tuof, /v\eu (gara>1co3) s$pqzqD'(arnssneg) ouawqui
'uorleulqruor
/v\eu (eserrg) oaco(ocsoqp,(rertela
ouafuqTn(opûauo^rqa7) 'V 'saya{g papqrul
-drt)
;iiiï;,Ëii#f frî {f"#i ïf,î'"î,î:'f
'6o7o19
Lg slFlnp"aq] Brl{V uersr{Bsqnsq
's op4îawVrcg sruaww.Ceql ur uorr.rt,rrrrnr"Tfl
aql dq snua8qns lul{l ruo{ peleredes eq uer pue
'rls 's ol1faaV ot pe]eler dlasolc s1ollfaraorqaz
('rlr s ototldorltuv
ro1sruazauo7aas) (d-rescoy11)
suaSdld'V dq pecelder
eq lou peou pu€ arueu pll"^ e sr (zlrmerotrAl)aomlos
'V '{,l8g urorJ olpflauo"tqaz sarcadsrnoJ o1da1 e
Jo
ane8(nt 'd'gZil ) uaTJIV 'serprusdreuorsr^erJo
esrnoc egl ur peqsllqelseaq Âlqeqord ru/r\ dur(uo
-uds orotrAl'eleredas ot lpcgrp pue repurs are
nryû
auo"tq
az ro sercads'yâi:,
lou ''eq ;o spueq pcrde eleldurocut;i'ïÀ1i
â:î;#^"i:iiir:,f"*
'ttg)
padrrls aql o] sri;ar' ô1p*flariùqag
tuesqealerrralJo sTrsru sgl Jo seurrros"leru
[eruos€]aur:(rçg
'snua8
relncoered
l(relsrlq
ol
pernper
wnalorrlto pue
eureu erIJ
Nlrs-r1nfl scNsrcs svsNvx do ^rrsus^rNn
slq] ur serueu snorJe^JoJ ruels
sHJ
?0ç
SvsrnuATlcs
AND PHvrocENY
nation, farinosa (Kl"S) , garrula (Rossi) , incana
mauritanica (Benoist)
(Klug) new combination,
new combination, ochroleuca(Pérer), quadrifasciata
(de Villers), socia (Kl"S) new combination, and
xerophila (Cockerell) new combination. Members
of this subgenus are found in eastern and southregion, Central
ern Africa, the Mediterranean
Europe, eastward to northern India.
Dizonamegilla
new subgenus
Erichson and Ktg,
Typ. species: Megilla sesquicincta
lB+2 (- Apit bicinctaFabricius, 1793, nec Schrank,
1781).
Diagnosis. Length 14-18 mm; apex of 57 of
male medioapically entire, produced bluntly and
expanded laterally without lightly sclerotized submedian circular area (FiS. 351); apex of SB of
male narrowly tapered, weakly bilobed, lateral
margin almost straight (FiS. 35k); gonocoxite
apically expanded, dorsoventrally flattened (Fig.
35j); gonostylus present, reduced to a blister.
Description. a. Length t4-l8 mm. b. Flagellar
segment 1 of female equal to length of next
3.2-3.5 segments together, of male about equal to
length of next 1.7 segments together. c. Mentum
.7 as long as prementum; paraglossa about as
long as stipes; galea in repose surpassing slightly
anterior edge of midcoxa; maxillary palpus with 6
segments; labial palpus with + segments; face
with white markings, paraocular marks present in
male but small, and in female well developed;
wing infuscated. d. Basitibial plate present in
male. e. 56 of male narrowly emarginate apicomedially (FiS. 35n). f. 57 of male medioapically
entire, produced bluntly, apicolateral margin with
large lobe, submedian circular area absent, carinate C-shaped area on apicosubmedian margin;
scleroti zation; length of
disc with longitudinal
apodeme about one-third length of disc (Fig. 351).
g. SB of male with apex narroq bluntly bilobed,
lâteral margin almost straight (Fig. 35k). h. T7 of
male medioapically shelf-like, dorsoventrally flatridge, and apitened with median longitudinal
cally trilobed, median lobe about one-third length
of lateral lobe (FiS. 35n). i. Gonocoxite of male
flu!apically simple, expanded, dorsoventrally
tèned, (Figs. 3 5j , -) . j . Gonostylus of male
apparently absent. k. Apex of penis valve about
1 . B times as long as wide viewed apicodorsally,
with small scattered setae. l. Bridge of penis
valves about 1 .1 times as long as wide, basally
emarginate.
Etymologlt The name Dizonamegilla refers to the
two whitebands of hair on an otherwise all black
metasoma of the type species.
of Dizonamegilla
Comments. The relationship
with other subgen era of Amegilla remains obscure.
However, its affinities seem to be with Amegilla s.
str. and Zebramegilla frorn which it is separated by
the limitation of apical bands of metasomal hair
to the last two or three segments, by the laterally
expanded apex of the male 57 and by the apically
rounded apices of the male gonocoxites.
OF THE ANTUOPHORINB
BEES
505
The name Apis bicincta Fabricius, 1793 is not
available for the type of Dizonamegilla because of
homonymy. The next available name is Megilla
sesquicinctaErichson and Klug, IB+2. The supposed type material of sesquicinctais in Berlin but
none of the four specimens there is from the type
locality nor of the proper sex. These specimens,
however, are the same as A. bicincta Fabricius as
Since sesquicincta and
interpreted.
traditionally
bicincta Fabricius are the same species judging
from their descriptions, the distinctiveness of the
species, their type localities (both from India),
and the Erichson and Klug material of M. sesquicincta in Berlin, I here designate a neotype of
Megilla sesquicincta Erichson and Klug (see Neotypic Designations).
Biology. The adults are active from November
to April.
Included Species. A. (Dizonamegilla) sesquicincta
(Erichson and Klug) new combination, somalica
and ,tetrataeniata
new combination,
(Magretti)
(Gribodo) new combination. Each of three !pecies of this subgenus occurs in a different region.
A. somalica occurs in Somalia, sesçluicinctain India
and tetrataeniata in Timor.
Megamegilla
new subgenus
Fabricius, 1793.
Typ. species:Apit acraensis
Diagnosis. Length 13-19 mm; apex of 57 of
entire, laterally expanded,
male medioapically
without lateral subapical circular area (FiS. 36d),
median section of disc narrow; SB of male apically
lateral
broad, with apicomedian emargination,
margins either diverging or parallel distally €ig.
36c); apex of gonocoxite of male dorsoventrally
flattened (Fig. 36a); gonostylus of male pres-ent;
basitibial plate of male absent; maxillary palpus
with five segments.
Description. a. Length 13-19 mm. b. Flagellar
segmenf 1 of female equal to length of next
3.0-3.7 segments together, of male about equal to
length of next 1.6-2.0 segments together. c. Mentum about .75 as long as prementum; paraglossa
about as long as stipes; galea in repose attaining
anterior to posterior edge of midcoxa; maxillary
palpus witti 5 segments; labial palpus with_ 4
legments; face with white markings, paraocular
màrk present in male, present to absent in female; wing infuscated. d. Basitibial plate absent
in male. e. S0 of male narrowly emarginate and
sometimes ventrally flexed medioapically, at base
a transverse depression someof emargination
times prèsent. f. 57 of male apicolaterally expanded, entire; median area of disc narrowed
sclerotization;
greatly; disc with longitudinal
length of apodeme .2+-.+0 length of disc (FiS.
36d). g. S& of male apically broad, emarginate
medioapically, lateral margins either parallel or
diverging distally (FiS. 36c). h. T7 of male
mediôapically bilobed. i. Apex of gonocoxite of
and expanded
flattened
male dorsoventrally
(Figs . 36a, b) (inner margin with lobe in acraen-
'^\er^
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pedolarrap IIaÀ\ e Jo ârueserd pnsn eql dq ueql
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dpeorq
dgeseq 'apyl se tuol se sârult B'I-B'0 sa^1"^
sruad Jo aEprrg 'l 'eeles pereuecs geurs r{}l^À
i{.lpsropocrde peù\er^ se eplÀ\ se Euol s€ se(ur}
'T 'luasard aleru
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9 lnoqe) alu8uole ot (peorq sB Euol ss lnoqu)
pepunor '({gg 'tl.d a}rxocouoE tuor; pelecreru
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ol snouerqrueru eprugo snldlsouoC 'f '(lgg 'ttg)
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'l 'paqolrr]
Jo alrxocouo8 yo ur8reu lerrde rauul
ol peqopq dgecrdeorpetu eprrr Jo /J 'g 'tuesge ol
:(tgg 'Et.f) JIerI
luesard uoltezltorelrs leqpntrtuol
FsBq lB Peqol ol lq8rerls dlerrrleler JISII pcrde 3o
ur8reur FretBI 'pepunor dlmorreu pue pacnpord
dgerparuocrde ol 'aleurtreura dyvrorreu o] dpeorq
ruo{ elqerru^ urtrenr pcrde 'xede peorq rl}l/r\
elsur Jo BS 't '(tgS 'ttd
rsrp Jo t{l8ua1 0+'-62,'
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peqrrrsepun uB) (tgg 'ttg) JprI Fwq uo qrool
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eleptue
urEreur 1ere}el
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xade qlyr\ epru Jo IS I 'eleur8rerrre dyvrogeqs
dgerpeurocrde eprrr Jo 9S 'e 'luasard o] ]uesqe
elerrr Jo a1e1d 1elqllrssg 'p 'pelersnJur ol realc
turrtr i(ogcutc ur ruasqe) eletu ur padolanep ila/vt
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stloratpu
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dl.rnrorreu 'odecs
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eletu eql uo s>lrsru aled Jo ToBI 'rlet{ Jrlle}eur
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Jo Tre[
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'dnor8 s${t
Jo seaq oq} Jo ozrs atrel
aq] ol sreJer opûauo3a747 'uoxel pa]€ler B Jo erueu
âqr 'opfayy snld 'e8re1 '('rC) soflayg '(îoput(1g
'dgeseq aleur8rerue dpeorq
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se
tuol
se
sârurl
0' I -g'0 se^le^ srued
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NlJ,s-r'rng scNsrcs svsNv)
do Àrrsus^rN.o sHJ
90ç
Systsl{ATlcs
AND PHyrocENy
,y
oF THE ANTuoPHoRINB
c
BnBs
507
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,4I
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FrlueO'g'V'eurats
l"ruoseleru
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SysrnuATlcs
AND PHyrocENy
and having a much more apically narrowed male
SB. Zonamegilla differs from Notomegilla by having
apically strongly emarginate 55 and 56 in the
males, lacking a tooth on the apodeme of the male
57 and having the male SB basally simple rather
than bilobed. In addition, Notomegilla is restricted
New
to Australia and Asaropoda to Australia,
Guinea and the Bismarck Archipelago.
Parts of Zonamegilla have been treated by Dover
(Oriental - 192+), Rayment (Australia - 1939,
t944 [not L9+2], 1947) and Lieftinck (Korea 1975). Some of the Rayment species are probably
synonyms but are listed below pending a revisional study.
Traditionally
Zonamegilla have been referred to
"blue
as the
banded bees" (Rayment, 1935) since
almost all members have blue or green metallic
hair in apical bands on the metasomal terga.
Biologlt. Michener (1960) described the nesting
biology of A. salteri. Cardale (1968a-c) described
the nesting biology, the lawa, and associated
parasites and predators of pulchra. The species fly
year around.
Included Species.A. (Zonamegilla) adelaidae (Cockerell) , andrewsi (Cockerell) new combination, asserta(Cockerell), atripes (Friese) new combination,
atrocaerulea (Dours)
new combination,
australis
(Raymettt), berylae (Rayment), binghamf (Schulz)
new combination, borneensas
(Cockerell) new combination, buruensis (Cockerell) , calceifera (Cockerell) new combination,
cinctofemorata (Dours),
cingulata (Fabricius) , cingultftro (Cockerell) new
combination, comberi (Cockerell) new combination, custos(Dalla Torre) new combination, dohertyi
(Gribodo)
new
combination,
dulciftra
(Cockerell), elegans(Smith) , fabriciana (Rayment),
ferrisi (Raymettt), flammeozonata (Dours) new combination, hackeri (Rayment) , holmesi (Rayment),
insignita Brooks , korotonensis(Cockerell) new combination, longmani (Rayment), longula (Rayment),
mimica (Rayment) , murra)ensis (Rayment) , niaeocincta (Smith) new combination, parapulchra (Rayment), parhlpate Lieftinck, perasserta (Rayment),
perpulchra (Rayment) , pulchra (Smith) , puttalama
(Strand) new combination, salteri (Cockerell) , samarensis (Cockerell)
new combination,
sapiens
(Cockerell) , shaferyella (Rayment) , stantonf (Cockerell) , subsalteri (Rayment) , ternatenses(Cockerell)
new combination,
thorogoodi (Rayment) , aegeta
(Bingham) new combination, walkeri (Cockerell),
wallacei (Cockerell) , whiteheadi (Cockerell)
new
combination,
whiteleyella (Rayment)
and zonata
(Linnaeus). Zonamegilla range from Australia (absent from Tasmania) to southeast China and
India.
Notornegilla
new subgenus
Smith, 1854.
Typ. species:Anthophoraaeruginosa
length 9-12 mm; hair
Diagnosis. Australian;
having a metallic green tint (sometimes only
visible under a microscope; except male of A.
chloroqtaneawith white hair); 56 of male simple,
convex without lateral depression, margin entire
. to shallowly emarginate medially; apex of S7 of
oF THE ANtHoPHoRINe
BBBs
511
male greatly expanded laterally suggesting Ack'
monopsis, without lateral subapical circular area
(FiS. 3Bd); apex of SB of male narrowed, truncate
or rounded, similar to that of Amegilla s. str. (FiS.
3Bc); apex of gonocoxite of male dorsoventrally
flattened, bent inward and apicoventrally (Figs.
3Ba, b); gonostylus of male absent; maxillary
palpus with 4 segments (chlororyaneawith 5).
Description. a. Length 9-I2 mm. b. Flagellar
segment 1 of female equal to length of next
3.0-3.1 segments together, of male equal to length
of next I.2-1 .9 segments together. c. Mentum.T
as long as prementum; paraglossa about .B as
long as stipes; galea in repose attaining anterior
edge of midcoxa or slightly less; maxillary palpus
with + segments (5 in A. chlorocyanea); labial
palpus with 4 segments ; face with white or yellow
markings, paraocular marks of male and female
well developed; wings clear. d. Basitibial plate
present in male. e. 56 of male simple, without
lateral depression, apically entire to shallowly
emarginate medially. f. Apex of 57 of male
greatly expanded laterally as in Ackmonopsis, entire and rounded to flat medially, with dense
transverse patch of hair, sometimes with median
weakly
apicolaterally
subapical protuberance;
bilobed with small upper lobe and large outer
lobe or with only large outer lobe present, outer
lobe sometimes with transverse carina mostly
paralleling margin on distal half; lateral margin
with toottr on basal half (aeruginosa rn addition has
tooth on basal third); lateral subapical circular
area absent; longitudinal sclerotization present;
length of apodome .38-.67 length of disc (FiS.
3Bd). g. Apex of SB of male narrowed, eithel
round ôr tiuncate; lateral margin on apical half
converging distally (FiS. 3Bc). h. T7 of male
apicomèdially with two teeth, apical margin between teeth straight. i. Apex of gonocoxite of
male dorsoventrally flattened, bent inward and
then basally laeruginosa also with apex laterally
expanded on basally directed apex (Figs. 3Ba, b)].
j. Gonostylus of male absent; ventral lobe of
gonocoxite of male present. k. Apex of P.lis
valve 1.8-2.1 times as long as wide, subapically
constricted, as viewed apicodorsally, with small
and scattered setae. l. Bridge of penis valves
1.0-1.5 times as long as wide, broadly emarginate
basally, not constricted but lateral margin round
(Fig. 3Ba).
Etymologt. Notos (Gt.), south, plus Megilla, a
name of a related taxon. The name Notomegilla
refers to the Australian distribution of this subgenus.
Comments. Notomegilla is closely related to Zona'
megilla and Asaropoda. It is separated from these
two by the light green metallic hair (not blue or
bright green), the expanded apex of male 57 with
a tooth on the apodeme and the apex of the male
SB more or less simple.
Some of the Rayment species are probably
synonyms but are included below pending revisional study.
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pcrde 'Qqcuu{aqw ruasqe) eqol papunor B r{lr/v\
eptu Jo âlrxooouot ;o ur8reur reuur :GgS 'tlg)
dgersrp turtrer'uor dpqtqs ol turtre^rp d1tq8r1s
surtreru 1ere]Bl'peqo11qdpunlq 'peorq optuJo BS
'8lg) seqol o/vuo] euo qllÀ\ ur8reur
;o xede :(rgg
uerpul
FJelelocrde'pepunor'(saptoqwoqopnas('tr
aql w eleurtreure dpeorq) arrlua eleur
Jo
lS Jo
xade ispueq se dllecrde dpo ro l.parra scsrpptral
'd1arr1an
turrarroc 'passardde
JIBq
l€ruoseleur
'asuap 'poqs rlerl JrJerorll '(syuawwoC
ees) u/v\orq
'stsouîotg
âcuarseqnd inru 1Z-tI tlltuel
os6
1'larercoc
i:K
addl
îil#:,,;î'o'
'erlerlsnV or rrruepue st:Dllllauoyo1g'(ruou
-dnU) aopqt(spue '(luanrde6) {nunu '1uaru,(e11)
olpxmau '(luarudeg) yawof '(ruetude11) opafuâ
' (gara>1ro oauo(to.tolrle' (qlt-S osowïn.rao'(tuavt
)
C)
-dnU) Dllaswnpo(opfauosollfi 'V 'saleadg
papqeul
Nrrsaang scNsrcs svsNvx do Àrrsus^rN.clsHJ
ZTç
SysrnuATrcs
i
,
a
i
I
a
AND PHyrocENy
censis),inner margin with one or two lobes (Figs.
3Bg, h). j. Gonostylus of male absent. k. Apex of
penis valve I.5-2.2 times as long as broad, with
small scattered setae. l. Bridge of penis valves
0.8-1.5 times as long as wide, shallowly to deeply
and broadly emarginate basally.
Etymologlt. Glossa (Gt.) tongue, plus Amegilla.
The name Glossamegilla refers to the long tongues
characteristic of the bees of this subgenus.
Comments. The relationship of Glossamegilla to
the other subgenera-of Amegilla remains uncertain. It seems to be closest to Micramegilla. Glossaby the following
megilla can be recognized
of characters: the usually brown
combination
appressed pubescence on the metasomal terga, 57
of the male wide apically with two blunt apical
lobes and the apex of the gonocoxite of the .male
with one or two lobes. Besides the brown coloration, these bees have short, velvety thoracic and
appressed metasomal pubescence and have long
tongues, attaining as much as 11 mm for the
galea and 23 mm for the glossa.
A. aiolaceais essentially all black with patches of
white hair along the lateral margins of the metasomal terga, fimbriata has white and black hair
intermixed on the thorax and a black metasoma
with small lateral white patches of hair, and
pseudobomboidesis probably a bumble bee mimic
having a yellow thorax, a black scutum and the
metasôma all orange (male) or T1-T2 yellow,
basal half of T3 black; apical half of T3 and
subsequent terga orange (female).
Lieftinck (19+4, 1956) described some new
species, redescribed many others, and gave a key
to the Malaysian members of Gbssamegilla
Biologlt. The adults fly from Mty to March.
Inchfud Species.A. (Glossamcgilla) amymone (Bingbouwmani (Lieftinck),
ham) new combination,
cinnyris
new combination,
brookiae (Bingham)
cyrtandrae (Liefnew combination,
(Lieftinck)
tinck) , elephas(Lieftinck) new combination, feronia
new combination, i,mbriata (S$th)
(Lieftinck)
new combination, florea (Smith) , gigas (Friese),
combination,
new
hanitschi (Meade-Waldo)
himalaj enses(Radoszkowsky) new combin ation, insularis(Smith) new combination, jacobi (Lieftinck)
new combination, lieftincki n. sp., luzonica (Cockerell) new combination, malaccemes(Frigse) _neY
combination, mesopyrrha (Cockerell) , pagdeneLieftinck, pendleburyi (Cockerell) new combination,
pseudobomboides (Meadeproboscidea Lieftinck,
subinsularis (Strand)
Waldo) new combination,
new combination , subrussata(Cockerell) new comurens (Cockerell)
bination, sumatrana Lieftinck,
new combination, uigilans (Smith) new combination, aiolacea (Lepeletier) and lunnanensis Wu.
Glossamegillais distributed from India to southeast
Asia intô the Indomalaysian area where most of
the species diversity is found.
Subgenus z{sa ropoda Cockerell
AsaropodaCockerell, 1926, p. 216.
Smith, 1854. OrigiTyp. species:Saropoda
bombiformas
- nal designation.
oF THE ANrHopHoRrNn
BnEs
513
Diagnosis. Australian species (including New
Guinea and the Bismarck Archipelago); length
13-24 mm; pubescence brown to gray (mostly
black in A. aurata from New Guinea and the
Bismarck Archipelago); maxillary palpus with 5
segments (see Description); apical margins of male
metasomal sterna modified, 54 usually produced
medially, rounded and having a thick brush of
hair (hair omitted from FiS.) (FiS. 39a), q5
broadly and deeply emarginate (FiS. 39a), 56
shallowly emarginate medially with one to two
dense patches of hair laterally (hair omitted from
FiS.); 57 apically somewhat quadrate, medioapically emarginate (FiS. 39f); SB apically narrowed (FiS. 39c); apex of gonocoxite of male
bilobed with long, narrow, upper lobe and small
ventral lobe (Figs. 39b, S); gonostylus of male
well developed (Fig. 39b).
Description. a. Length t3-2+ mm. b. Flagellar
segment 1 of female about equal to length of next
2.8-3.1 segments together, of male equal to length
of next 1 .3-2.0 segments together. c. Mentum
.6-.7 as long as prementum; paraglossa 0 .7-1.1 as
long as stipes; galea in repose attaining anterior
edge of forecoxa to anterior edge of midcoxa;
maxillary palpus with 5 segments (sixth is fused to
fifth segment) (/. qurata^lrom Bismarck Archipelago occasionally has 6); labial palpus with 4
ségments; face with yellow to reddish-yellow
màrkings, paraocular marks present in male (FiS.
39e), absent in female (FiS. 39d); wings clear to
infuscated. d. Basitibial plate present in male. e.
Apical margins of male 54-56 modified, 54 with
median tenth produced, rounded, with dense
hair, 55 wit!
patch of black to reddish-brown
deep, broad median emargination and with I or 2
dense patches of hair laterally, 56 apically flexed
ventrad with shallow, median emargination and
rounded ridge on apical
median, longitudinal,
sixth of disc, laterally with dense patch of redbrown hair (hair omitted from drawing) (FiS.
39a). f. Apex of 57 of male somewhat quadrate,
apicomedially entire or with small, shallow emargination, apicolateral margin rounded, lateral
subapical carina often present, circular area abridge on
sent, often with median longitudinal
apical half; lateral margin with tooth on basal
hâlf, longitudinal sclerotization present; length of
apodem e .41-.46length of disc (FiS. 39f). g. SB of'
male apically
-fourth narroq bilobed; lateral margins of
to half converging distally, margin
apical
angulate or straight (FiS. 39c); SB basally more or
lesi rounded (except dautsoni basally trilobed with
lobes basally directed). h. Apex of T7 of male
apicomedially with 2 widely separated teeth ("*cèpt dawsoni shelf-like without teeth), apical margin between teeth straight (Fig. 39a). i. Apex of'
[onocoxite of male not dorsoventrally flattened
6ut bilobed, with large, apical, elongate upper
lobe and small blunt to acute, ventral lobe as
viewed ventrally (FiS. 39b), it profile bilobed
(FiS. 39g). j. Gonostylus of male present, well
developeâ, 5-10 times as long as wide. k. Apex of
'â
'elnsdzr
'g ',u,errr
'l,S 't 'peeH 'g 'q 'a'rsr,rprluazr 'gg 'p
âr^
âprs
plruâC
lenue^
'^ âI^
'elnsdec
'g 'r'rau
'(prrrl-o rtnq)
I"rtuâ^osrop
l€lruâC
l€rluâ^
Z.t pue gS-gS Igedrcurrd Sur,uoqs
Bruos?larugo xady 'V 'ele,u 'g-g iapuray 'q lapu '3'-V 'q*ofiquoq (opo{onsy) qf&*V .6g .CIt
I
I
I
I
I
;
Nrrs-r1nfl scNslcs svsNvx do ^rISusArN.o sHJ
it9
SysreldATrcs
AND PHvrocENy
penis valve | .B-2.2 times as long as broad as
viewed apicodorsally, with small scattered setae. l.
Bridge of penis valves 0.6- I .6 times as long as
wide, broadly emarginate basally.
Comments. Asaropoda is most closely related to
Zonamegilla and Notomegilla. It is separated from
the former by the lack of metallic hair, the
presence of a well developed male gonostylus, the
pale clypeus and characters of the male 57, SB
and genital capsule given in the Comments on
Zonamegilla. It is separated from Notomegillaby the
larger size, brown or gray pubescence, somewhat
quadrate apex of the male 57 and presence of a
well developed male gonostylus.
This group of bees has been given generic
status by some authors (Cockerell, 1926; Cockerell, 193lb; Rayment, 1951), but is clearly only
of subgeneric import considering many of the
derived characters it shares with all other subgenera of Amegilla. Rayment (1951) treated the species of Asaropodabriefly, describing six new species
but giving no k.y to species or good diagnostic
characters for the group. Some of the Rayment
species are probably synonyms but are included
below pending a revisionary study. There are
several species, both described and undescribed,
which look superficially like ,4. bombiformis.
Traditional characters for Asaropoda have been
the red-brown coloration and loss of the last two
segments of the labial palpus, but it appears that
these segments are easily broken off and in life
there are four segments. Michener (1965) defrned
Asaropoda more clearly and recognized its affinity
within Amegilla, giving it subgeneric status and
building upon the work of Rayment (1951).
Biologlt. This group of bees flies year around.
There is an apparent absence of Asaropoda frorn
the dry interior of Australia, but this rcray be from
lack of sufficient sampling.
In Anthophora turret-making
is found only in
species of the subgenus Melea; in Amegilla it is
characteristic only of some species of the subgenus Asaropoda. A single female of A. rickae from
Bolgart, Western Australia (Rayment, 1951) and
a big aggregation of dawsoni from 160 miles (258
k*)
northeast
W. Australia
of Carnavon,
(Michener, 1965) both were found with turrets
over their nest burrows. Cardale (1968b-d) gave
some biological notes about an unidentified species of Asaropoda and described its lawa (Cardale,
19 6 B c ) .
Included Species. A. (Asaropoda) albiceps (Ruyalpha (Cockerell),
ment), albigenella Michener,
anomala (Cockerell) , aurata (Friese) new combination, bombiformrs (Smith), calaa (Rayment) , cygni
(Raymettt), dawsoni (Rayment) , dentiaentris(Ruymen|,
epaphrodita n. sp., grisescens(Rayment),
houstoni n. sp., imitata (Rayment) , meltonensis
punctata (Ruypreissi (Cockerell),
(Rayment),
ment) , rhodosclmna (Cockerell) , rickae (Rayment),
rubricata (Rayment) , rufa (Rayment) , rufescens
sordida (Rayment),
(Friese) , sc)mna (Gribodo),
and uictoriensis (Rayment).
sordidula (Rayment),
oF THE ANrsopHoRrNn
515
Bens
This
group is Australian
(including
New Guinea
and
the Bismarck
Archipelago
Tasbut
not
mania).
Auecnte
TNcERTAE snDrs
The following
species of Amegilla cannot be placed
with certainty into a subgenus because the males are
unknown, the types are lost, destroyed or have not been
studied yet: A. aduenula (Cockerell) new combination,
annos (Vachal) new combination,
aspergina (Cockerell),
atriceps (Radoszkowsky),
bucharica (Gussakovsky),
cafra
(Friese) new combination,
caldwelli (Cockerell),
camelorum (Cockerell),
new combination,
cana (Walker)
candens (Pérez),
didella (Priesner),
candida (Smith)
new combination,
centralas(Cockerell)
can-
new combination,
deltoides (Buysson) new combination,
expleta (Vachal)
new combination,
glyqtrrhizae (Gussakovsky),
gussakouskyi (Popov) , hypocyanea (Cockerell)
new combinanew combination,
tion, karakumensis (Gussakovsky)
katangensis (Cockerell) new combination, liberica (Cockerell)
new combination,
lutulenta (Khg),
macroleuca (Cockerell),
maculicornis (Lepeletier)
new
combination,
marqueti (Pérez), mephistophelicana (Strand) new combination, nigropilosa (Friese) new combination, niuea (L"peletier) new combination,
nonconforma n. sp. , potanini
puluuea (Walker)
(Morawitz)
new combination,
new
combination,
rapida (Smith) new combination,
ruficornis
(Dours) new combination,
rufipes (Lepeletier)
new com-
(Friese) new
bination, semipulaerosa (Dours), senegalensas
new combination,
combination,
simbana (Cockerell)
subtorrida (Cockerell)
sjoestedti(Friese) new combination,
new combination,
torensis (Priesner) new combination,
tubifera (Cockerell) new combination
and uestitula (Cockerell) new combination.
CLADISTIC
ANALYSIS
Using a list of variables for which polarity
could be determined with reasonable confidence, I have developed a list (Table 1) of 70
apomorphous tribal, generic and subgeneric
characters found in the tribes Anthophorini
and Habropodini, the genera Anthophora and
Amegilla and the subgenera of Anthophora. A
similarly constructed list (Table 3) of 17
apomorphic characters was compiled for the
subgenera of Amegilla. The distribution of
these characters among the taxa, i.e., the
data on which the cladograms arebased, are
given in Tables 2 and + respectively.
Cladistic methodology is discussed in the
section on Methods. Because decisions on
what taxa to reco gnize were made after the
cladistic analysis, some species groups, not
now recognrzed as taxa, were included as
separate entities in the study. Table 1 therefore includes some characters relevant to
those groups but not used as subgeneric
characters.
'opûatuy
anbrun
ol
sr
pârâqruerp
..uârunl
epn9p,, eq'I,'ueurnl pap8 morreu e18urse e^Eq I[" seâqeurprrluâr pu" âurrârne'eunuor11a141
['(qç
'8tg) uaunl MorJ€u a1Sursz qrrrvreepgl '(t) (rç '3rg)
'çI
,,ueurn1 pâJaqrueqr alqnop,, " rllrr\ "â1eC
'op$auou1t1gur dpo punoJ sr Âqdrourodelne
srqa ['aldurrs ro pâqol ^1daapul9retu F3ydB
tltli\À_run11aqelg] '(t) (qgZ '3rg) urS.retupcrde uo seqol flerus snorerunu qly* 'Euol urnllaqelg :71
'opt8ewoq\uy dqdrourode enbrun u" sr snII
1o
['saqo1a4t-ratug 8uo1 Isre^as qlrru dlerrde runllaqeyg]
'(t) (pçZ 'Ërg)
paprs lepered prrc pa^\orreu sprul-o^ l lecrde 'aldurrs Âgeerde tun[âqelt 'gI
't1s 's otot$otpuy
ol enbrun sr dqdrourode tqf ['lt prr€ €I selqerre^ ur s? ro alduns ro pâqol ilsnoraurnu puz dldeap
ur8.reur pcrde qlrm tunllaqelg] '(I) luo"J ur ruo{ pâ^{âr^ se pcr1d11e'aldurrs z(gecrdeurnilâq"ld 'ZI
ur Surlseu o1uorlzldepe rrc sr pue uopostlColenbrun sr alqrpue,., et'tueprJt eqa ['qtool 1""rdrqr,rÏJ,ffr
'(1) qleel prrduqns râ^{ol puu roddn
'oleluepul
tnoqlr/\^ 'slzlueprg rlqlp*Wl
elqrpuel4l 'tI
çr,rl
'qloor relnqrpueur raddn u? qtr^ sâIeur a^Eq rurpodorqell pue rurroqdoçuy
reqto 'o1ufioqag auos
go (qdrourode u" sr srql ['qroor pcrdeqns qlr^ âpruJo alqrpueyq] '(1) eldurrs â1eruJoelqrpupl l '0I
.rugoqdoqruy
erp ur opûauroqtuy pue r11ûauoutrlg 11e'oe7cy47
arnos ur ,(po puno; dqd.rourode snor,rqo ue sr sgudqlnoru
aqr;o qfual pâs?ârcur ârp roJ areds apr,rord ol aoeds rel"ur oql ;o uolte5uole aqa ['8uo1 ueç
râPrM serull âroru ro ezrg 'reaurl âFurâJ Jo ereds rzpl4i] '(1) 8uo1 u"ql JeprÀ\sorurl aarql 'epraa,eleuray
go aczds *py.t '(Z) 3uo1ueql .raprÀ\sarull eotrp uuqt ssal 'ale8uola d1Euo.r1s
âlzruâJJo acedsrep;41 '6
'opUdolpU umuJv
dnort e;o dqd:ouodeuds y ['eldurrs
Jo
ol€ureJJo uruqe1l '(nZg 'StS) uorlcalord el€lncreqnlrq pcrdeqns uerperu qlrâÀâ["urâJJo urnrq€rl 'B
'eeuuoqdoqluy
râqlo ut punoJ tou sr 1nq rurroqdoqruy eqr ur err^,u ts"al l" uesr.redpuepuadapur szq ruruq€l peqolFr
Âleorde
eq; ['arrluadlerrdeet'.', Journrqul]'(lçz 'rr-r)psqolrr^ty,ii.TirxfËrri,ffi,r*;*
",
pe1ur1dlqrssod sr uoueuroueqd peardsapr,n rliqa 'szposso1floæ;çgo
saroadsauo irue ointpuy go sirceds o,r,rl
's1douyn1'('ururoc 's,radarno141)alqmday4J
vettreurv qlnos Jo snua8qns paqrJJsâpunue '(wnryttlluop)
Dsnqtorl w 'oytg(oqeqp serredsduuru u1 'rpo{otqop 1o sercedsoa,rtur ''ds 'u oxop,to{ opûautF ur punoJ
osle sr uorlrpuoJ pe^rrâp e^rlrurlsrp srqa '(sarcads6out) ototldotlluoto1stT4T
ol enbrun sr dqdrourodeuds
srql rur.roqdoqtuy eql u1 ['eetas pezrrnodpueS 'pezrlercadsun qtrm lu€râqqord l18uor1s o1 dlele.raporu
eleurâJ Jo eceg] '(1) (qft 'Std sâpsrrq pâuâlt"U lf,âre lnols Jo ^Àor âsrâ^srrerl ? 'sl l"qr 'uaped
e qfrrvrdgeseq uâUo pue azles d,rea,r.ropalooq dgarrde '1ceraqlrar sneddp 'peua11egeFtuâJJo âJ€t 'g
dqdrourode ue sr dlqeurnserd slruur âsâql Jo eeuesard eqt âroJeraql '(slreru esâql tsol â^"q
o1pïauty lllal e) opôawy rcl ldacxa slreur relneoe.red apd gtrirr eeugoqdoqluy âql w sâprueJ ou lsoru1e
ârE âroql ['ruasqe âleurâJur sllr"ru r?lnf,o?Jed ep4] '(1) luesard epureJ q sTr?ru r"lnf,o"red ap4 'g
'sTreru r"InJoe.red epd eql
Jo ssol âq ot sueas
ruuoqdoçuy oql unprilr e1e1ssnoqd.roruode aqt 'ero;ereqa 'sTreru rulnf,o€red epd er{t â^eq qcrrl,rÀ
sâ^rlulâr osolr e^€rl saJEJTf,"lq qlln se1etuâ^"r{ l"ql sarcedsge l€rll sruaeslI 'sTJeur.relnroered apd
arreq 're,rainoq 'seleur aurroqdoqlûz pu" eurpodorqeq II€ lsou:rlv 'sEâJ"J€lmoe.red 1ce1qazreqsaurJeJne
epru Ip lsorul€ prre seurruolrlâru ât"ur 11" 'seleur aursdoleuroxe dueru âJurs ur?ueJun sr âlqzrre^ sql
'(1)
ruasqe eleurJo Tr?ru r€lnf,o€red ap4 '7
Jo,bFelod aq; ['luaserd eleurJo Treru r"lnro"Jed epd]
'ototl{ot1\uy eraue8qns snorre^
1o
uI uaslre z(puapuadapur s"q râlr"r"rlf, strt;- '(tptt4o!tag) o.roq{oqtuy p* on ît*V Jo sâpurâJ I1e lsou{u
ur punoJ e.re sturl.reur I?lJeJ âIed 'a1e1ssnoqdrourorseld âql sr srql dlqeurnserd puu 'ua,rotrlun a.re
sepurâJ ur sturlreru ["rJ"J sââqeursdoleruoxaeqt tuoruy 's8urryzu rlJns o^Eq saurpodorqeqgo saroads
na; e dpg 'rurroqdoçuy aql u1 tdacxa eeurroqdoçuy eqt ur erer sr 8ur1"reurprce; pruarun8etu qlu
âIerrrâJV ['1ce1q aprueJ 3o aceg] '(1) snaddp uo Sunlreur plueurn5elur apd qrrar "**"{Jl,iil*.rir""
sSurlreur prcu; irnogedezreq'saleur pece;-apd â.Àeqqf,rqa seaq]soru se IIem se 'aepFoqdoqruy crlrsered
-uou 11ulsou{V ['s8uqreur no11edqtr,u apru Jo aceg] '(1) s8urlreru 3tnl^ qlri e1etuJo âr"ù 'Z
'a1e1scrqdrourorsald aql pârâprsuor âroJârerp q qrII À araurole8eg
lsrg uoqs dle,lleler e â^"q I1e saeq eursdopuroxe pu" eunuotr1aru 'euuorne 'aurpodorqell ['(qZ '8tS)
sluaur8asEurpeeocnsrâ^ÀeJro eerqlJo qfuel pror ot pnbe epruâJJo ; tueruSesre11a8e1g]'(1) slueurSas
Eurpaecrns rnoJ ol eeJql u"qt aroru Jo qrtuel pror o1 pnba epurâJ Jo 1 tuaurSes reqetelg 'k) @t
'Erg) sluaruSesSurpeeccnsrnoJ ueql âroruJo r{rtual pror o1 pnba epruaJJo luaur8as re1p8e1g '1
1
'rurpodorqull pue
rur.roqdoçuy soqrl eql rrref,uor 0/-09 salq"u€r"otot1{otpuVulpr^r InJesnselqerre^ er? 6ç-I '0 s€ papor
pu€ sle1f,€rq ur pâlsrl âr" sel?ls snoqd.rotuorsal4 '(1) raqurnu el€ls snoqdrourodeui(s;o uone,urep
rerlunJ e $ âroru ro (7)yo âlels V '(1) sz papoc sr e1e1ssnoqdrourodeur(sr (Of-l) âlq?rr"^ qr"e rot
.INIqOdOUSVH
ONV INIUOHdOHJNV
'I
T'ItrVI
dO \rUSNSCSNS dO SS'IS\ilIUVA TO JSI'I
STSIUJ
SHI
CNV VUOH{OHJNZ
Nrrs-r1nfl scNflrcs svsNvx do Àrrsus^rNfl
sHJ
9Iç
SysrnuATrcs
AND PHyrocENy
oF THE ANrHopHoRINn
Bnns
517
16. Sixth segment of maxillary palpus equal to about half length of fifth (1). [Sixth segment of
maxillary palpus about equal to length of frfth.] Among those tribes of the Anthophorinae which havea
six-segmented maxillary palpus such as exomalopsine, melitomine and habropodine bees, the sixth is
about equal in length to the fifth as it is in Anthophora.Amzgilla has a reduced sixth segment, an
apomorphy.
17. Bàsistipital processbilobed (Figs. 25a, 25e,25f) (1). [Basistipital processwith one projection
(Fig. 5d).] Hàbropodine, centridine, eucerine and melitomine bees have a basistipital processwith one
projection. Anthophorahas a secondary projection on the process.
18. Scape antèriorly with dense pilosity such that integumental surface is hidde! (1). [Scape with
pilosity
anteriorly sparse to absent.] The derived condition is found only in sorne Hel'iophila.
19. Mentum with long anterior tôoth near basal third (Fig. 22k) (1). [Mentum simple (with single,
short tooth only in,4. (Dasymegilla)excrsa)near basal third.] This strong synapomorphy unites Melea and.
Anthophoroidrs.
20: 57 of male as wide as long, rather square in ventral aspect, sides almost equal in length and
lateral margins almost parallel (Fig. 18d) (1). [S7 of male not square in ventral aspect,_width less than
length, lateîal marginiconvergin{ apicâXy.l This apomorphy is found only in Càranthophora._ _
21. Midtarsal brush of male present (Fig. 2h) (1). [Midtarsal brush of male absent.] This
apomorphic character is found almost exclusively in Pyganhophoraand its relatives; except for a few
oiher speciesof Anthophora,it is unknown in other Anthophoridae.
22. Midbasitarsal brush of male present (Fig. 29b) (1). [Midbasitarsal brush of male absent.] See
comments on character 21, except this has independently evolved in Caranthophora.
23. Middistitarsal brush of male present (Fig. 2h) (1). lMiddistitarsal brush of male absent.] See
comments on character 22.
24. Hind basitarsus of male modified with tooth and/or ridge (1). [Hind basitarsus of male simple.]
Most Anthophoraand ù1 Arnegilk males have simple hind basitarsi. This modifrcation of the hind
basitarsus apparently has arisen once in the Anthophorini and is an apomorphy'
25. Basitibial plate of female pointed apically (Fig. 29b) (1). [Basitibial plaf: of female apically
rounded.] This is a unique character found in Clisodon.
26. Basitibial plate of male poorly defined to absent (1). [Basitibial p!a1e g'f male. present, well
defined.] Presumâbly the genetiè expression in the female to produce a basitibial plate is car-ried over
and expiessed in the male. Though the polarity of this character is uncertain, I believe that the loss of
the basitibial plate in the male is an apomorphy.
27. Hind ti6ia of female with posterior margin with simple hairs (Fig. 22h) (1). [Hind tibia of female
with posterior margin with plumose hairs.l This character is unique to three small groups of
Anthophora.The remàining Anihophora, all Anugilln and Habropodini have plumose posterior zones of
hair ôn their hind tibiae and thus the simple hair in this zone probably is an apomorphy.
28. Hind femur and tibia of female with white appressedplumose hairs almost completely obscuring
outer surfaces(Fig. 321)(1). [Hind femur and tibia of female without appressedplumose hairs on outer
surfaces.] This apomorphy is unique to some Heliophik.
29. Hind basitarsus of female with posterior distal process acute (Fig. 22h) (1). [Hind basitarsus of
female with posterior distal process rounded.] This is an autapomorphy of Melea.
30. Apex ôf hind tibia of male with anterior spine, hind tibial spurs separated such that one spur is
inserted near apex of spine and the otler in the corium (Fig. 2aj) (1). [Apex of_hind tibia of male
simple, hind tibial spurs normal, in close approximation and both inserted in the corium.l -In the
and rarely seenin other Anthophorinae. Such a
Anthophorini this is an autapomorphy of Anthophoroid.es
peculiar
modifrcation of the male hind tibia must be an apomorphy. .
31. Posterior inner surface of female hind basitarsus with longitudinal cavity (Fig. 18h) (1).
[Posterior inner surface of female hind basitarsus convex or angulate.] This character is unique to
Caranthophora.
32. Hind femur of male dilated, bulbous (Figs. 24i, 24j) (1). [Hind femur of male normal, slender.]
Such a peculiar modification of the male hind femur is not necessarily correlated with a toothed hind
basitarsus. This unusual modiûcation must be an apomorphy.
33. Pygidial plate of male with carinate margins, dorsally flattened (Fig. 13c) (1). [Pygidial plate of
I am
male, if present, without carinate margins.] Carinate margins are unique to Pyganthophora..
uncertain as to whether the presence or absenceof a male pygidial plate is apomorphic. Following the
reasoning given in charactei 26, a pygidial plate on a male may be an apomorphy. Males of Amegilla,
eucerine,-àelitomine and exomalopsine bees all lack pygidial plates but habropodine bees possess
them. Even though tÏe Habropodini are the sister group to the Anthophorini, almost all the rest of the
male Anthophorinae lack pygidial plates and thus I believe the presence of one is an apomorphy. On
the other hand, the presence-of a male pygidial plate could be an apomorphy indicating,sister group
relationship of the Ânthophorini and Habropodini. The loss of the pygidial plate would then be an
apomorphy
within the Anthophorini.
S+. fygiaia plate of male without carinate margins but dorsally rounded (Fig. 15c) (1). [Pygidial
te algord ur 'a3:e1 aszqouog] '(l) (egZ .8rg) tuol uerp râp!^Àqcnur elgo.rd ur .gerus â$qouoC .çç
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BI9
SysrpuATrcs
AND PnyrocENy
oF THE ANtHopHoRINn
Bess
519
least half as long as wide (Figs. 19b, 30i).] All habropodine and eucerine bees have large gonobases.
The gonobase has presumably become apomorphically reduced in various anthophorine groups.
56. Female constructs turret of mud over nest entrance (1). [Female leaves nest entrzrnce
unmodited, without turret.] Even though the old literature gives the reader the impression that most
Antlnphora build turrets, they are actually unique to Melea.
57. Stinging behavior absent (1). [Stinging behavior present (1).] An obvious apomorphy of Melea.
58. Batesian and/or Mûllerian mimicry of bumblebees (1). [No apparent mimicry of bumblebees.]
Anotlrer autapomorphy of Mclea.
59. Nest constructed in wood (1). [Nest constructed in earth.] Only Clisodonnests in wood, with the
possibleexception of Anthophora(Anthophoroides)
signnta.Since all melitomine, eucerine and habropodine
bees nest in the soil, soil nesting presumably is primitive.
60. Gonostylus fused to gonocoxite (1). [Gonostylus articulated with gonocoxite.] Melitomines,
eucerines and habropodines all have articulated gonostyli. Presumably the fusion ofthe gonostylus to
the apex ofthe gonocoxite is an apomorphy found in anthophorine as well as centridine bees (except
Camonomada).
61. Bridge of penis valves basally elongated and often emarginate (1). [Bridge of penis valves
triangular, basally simple and either square or bluntly pointed.] All Anthophorinae -have the
plesiomorphic type ofbridge ofpenis valves except the Anthophorini which have the bridge ofthe penis
valves elongated to varying degrees.
62. First recurrent vein of forewing meeting posterior margin of second submarginal cell near its
midpoint (1). [First recurrent vein meeting or nearly so with first transverse cubital vein.] Melitomine,
eucerine and habropodine bees have the first recurrent vein of the forewing meeting or nearly so with
trst transverse cubital. Anthophorines have the apomorphic state.
63. Third submarginal cell of nearly equal length on posterior and anterior margins (l). [Alterior
margin of third submarginal cell about half lengh of posterior margin.] All bees in the outgroup have a
third submarginal cell which is narrowed anteriorly. All Anthophorini have the apomorphy._ ^
64. 56 of male strongly convex, apically entire, apical margin strongly bent ventrad (1)- {Si of male
gently convex, often apically emarginate, apical margin not or only slightly bent ventrad.l The 56 of
male centridine and habropodine bees (except pachymelines) is strongly convex, apically bent ventrad
and curving only on the lâteral margins (few Anthophorahave the apical margin bent ventrad but the
disc is not strongly convex).
65. One gonoJtylus on gonocoxite of male (l). [Two gonostyli on gonocoxite of male.] All bees in the
outgroup hàve two gonostyli per gonocoxite. One has apparently been lost in the Anthop!_otiqi. - 66. Gonostyli much flattened and wide, paddle shaped (l). [Gonostyli elongate, usually qlghtly
clubbed o. oniy slightly flattened.l Melitomine, eucerine, centridine and anthophorine bees all have
the elongate gonosiyli if present. The flattened, paddle shaped, ventral gonostylus is unique to the
Habropodini.
Oi. Stip"r *ith apical concavity, housing the stipital comb, one-fourth length of stipes (1). [Stipes
with long apical concavity housing stipital comb, broadly and shallowly emar,ginate,extending at least
one-third lèngth of stipei.] Among the non-parasitic Anthophorinae only the Habropodini and the
Anthophorinf have a short round notch housing the stipital comb. This is a strong synapomorphy
uniting these two tribes.
68. Forewing-thèn
apically papillate, papillae simple, not tipped with hairs (1). [Forewing apically hairy
paiillJe tipped with hairs.] Most bees have_hairy wings. In Anthophorinae.the
or if papillate,
hairs bn the papillae have been lost twice, once in the Centridini (Ctntris) and again in the habropodineanthophorine lineage.
69.-Tibial scopiconsisting largely of simple hairs on outer surface (1). [Tibial scopa consisting
largely of plumoie hairs on outer-surface.] Almost all non-parasitic female Anthophorinae \ay9 tfe
tiblal icopâ consisting mostly of plumose hairs, except the Habropodini and Anthophorini in,which the
scopal hâirs are largèly simple, iometimes with some plumose hairs along the anterior and posterior
margrns.
70. Jugal lobe of hind wing one-third or less length of vannal,lobe measured from wing base (l).
[Jugal-lo6e of hind wing at-least one-half length-of vannal lobe.] Àll Anthophorinae. except-theÉa6ropodini and Anthophorini have relatively long jugal lobe one-half to three-fourths t!'relength of
have the jugal lobe nearing one-half the length
the assôciatedvannal lobès. (Large speciesof Pachymclus
of the respective vannal lobe.) The ihortening of the jugal lobe must be an apomorphy'
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SysrnuATrcs
AND PuyrocENy
oF THE ANrHopHoRINn
BBns
s21
8. A large lateral tooth near end of apodeme of male 57 (Fig. 3Bd) (1). [Apodeme of 37 of male
simple, lateral margins more or less parallel.] This tooth is found only in Notomegilln.
9. Lateral margin of 57 of male witJr a tooth on basal half (Fig. 36h) (1). [Lateral margin of 57 of
male without a tooth on basal half.] The Habropodini lack this structure but some groups of Amegilla
and Anthophorahave it.
10. 57 of male with width of disc on basal two-thirds equal to about half of width of disc on apical
third; basal two-thirds almost parallel-sided (Fig. 35h) (l). [S7 of male with width of disc varying on
basal two-thirds such that basal two-thirds is not parallel-sided.] Eucerini, Habropodini andAnthophora
do not have 57 with a long parallel-sided disc which is slightly expanded on apical third. Such a
structure is presumably an apomorphy.
11. SB of male witl two well developed basal processes(Fig. 38c) (1). [SB of male simple, without
any basal processes.] This is an autapomorphy of Notomegilla.
12. Median apical lobes and lateral lobes protruding from apex of S8, with margin between them
strongly emarginate (Fig. 37i) (1). [Median apical lobes weakly developed or lateral'nlobes absent;
Micramegilla.
margin between them only weakly emarginate to entire.] This structure is only found
13. S8 of male apically narrowed, lateral margins converging apically (Fig. 37c) (1). [S8 of male
apically broad, lateral margins more or less parallel (Fig. 36c).] Almost all males in the outgroup have
broad eighth sterna. The apical tapering of this sternum is presumably an apomorphy.
14. Gonobase in lateral view much longer than wide (Fig. :69) (t). [Gonobase in lateral view about
as long as wide to wider than long.l This apomorphy is only found in Ackmonopsis.
15. Gonostylus greatly reduced to only a membranous blister or absent on gonocoxite (1).
[Gonostylus present, from 3 to 10 times as long as wide.] The absenceof a gonostylus is an apomorphy
since all bees in the outgroup have gonostyli.
16. Penis valve with strong laterally protruding lobe on apical half (Fig. 36f) (t). [Penis valve with
small lobe or no lobe laterally on apical half. This character is unique to Megatnegilla.l
17. Pubescence metallic (1). [Pubescence not metallic.] None of the non-parasitic bees in the
outgroup have metallic hair.
TABLE +. MATRIX OF CHARACTERS OF
THE SUBGENERA OF AMEGILLA LISTED
IN TABLE 3.
Variables 1-t7 are representedby the columns
from left to right. Explanation as for Table 2
exceptP-1&2.
Anthophora
Amegilla s. str.
Zebramegilla
Micramegilla
Ackmonopsis
Megamegilla
Aframegilla
Dizonamegilla
Notomegilla
Zonamegilla
Asaropoda
CLADISTIC
11111111
123+567
r234s67890
00000000000000000
10010100
000000001
00100000010010100
00100000100100100
100
00000000100001
11000000000000010
1
10000000
20000000
10100000000010100
11
01010101
0P10000
2 0 1 0111 0 1 0 0 0 1 0 1 0 1
0101 010100010000
RELATIONSHIPS
The relationships of the AnthophoriniHabropodini to other Anthophorinae are not
clear. Michener (L97 +) placed the Anthobephorini (which included Habropodini)
tween Centridini and Eucerini, positioning
ericrocine(and
the
centridines
the
"terminus"
of the
rhathymines) at the
seems
placement
This
Anthophorinae.
unjustified now that Caenonomadahas been
reassigned from the Exomalopsini to the
Centridini (Snelling and Brooks, 1985). The
many plesiomorphies of Caenonomadaindicate a more ancestral position for the Centridini, possibly near the exomalopsines. If
this is correct then the venational apomorphies hitherto used to associate the Anthowith the Centridini
phorini-Habropodini
(i... , shortening of the second abscissa of
M + Cr, loss of hairs on the alar papillae and
shortenitg of the marginal cell) are convergent between the more derived members
of Centridini (i... , Centris and Ptilotopas) and
These wing
the Anthophorini-Habropodini.
attributes may be associated with large size
or rapid flight. Probably the closest relative
lineage is
of the Anthophorini-Habropodini
the Eucerini, the two groups at least sharing
a small stigma. The former has diversified
and probably arose in the Old World, the
latter diversified and most likely originated
in the New World.
In the following paragraphs, character
numbers in parentheses are from Table 1
except as otherwise indicated.
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AND PHvrocENY
phora is the bilobed basistipital process (17),
which is unique in the Anthophorinae. Even
ignoring these characters, I can find no
subgeneric group of Anthophora from which
Amegilla might have evolved.
There are four main lineages of Anthophora
shown in the cladogram (Fig. 40). The first
is a group consistitg of Anthophora s. str.,
Pyganthophora and Lophanthophora characterized by long first flagellomeres (1) and
"brushes" of hair on
males which often have
their midlegs (21-23). The second group,
consistirg
of Petalosternon, Clisodon and
Dasymegilla, has males with a trilobed labrum
(7) and 57 with a strong lateral tooth on the
"deapodeme (+2). The third and most
rived" group contain s Rhinomegilla, Anthomegilla, Melea, Anthophoroides, and Paramegilla.
Their male synapomorphies include an apically narrowed SB (43) and an elongate,
basally emarginate bridge of penis valves
(49). The fourth group is Caranthophora,and
Heliophila. The females of this group have
white to yellow facial markings (3), males
have short apodemes on 57 (41) and most
have a small gonobase (55)
Figure 41 is an alternate cladogram which
gives separate origins for the RhinomegillaAnthomegilla and Paramegilla groups, mainly
because the former has males with simple
hind legs (2+). In addition the Melea-Anthophoroidesgroup is shown as the sister grouP to
Heliophila because they share a small gonobase which would have to be reversed in the
Estebana species group of Heliophila (55). I
prefer the cladogram in Figure 40 since it
appears that Rhinomegilla and Anthomegilla
have their clos*T relatives among Melea,
Anthophoroidesand Paramegilla as suggested by
their similar termin alia. The unmodified
hind legs (2+, 32) of the Rhinomegilla-Anthomegilla group probably indicate that it is
more ancestral than the Melea-AnthophoroidesParamegilla group. Even though both cladograms are about the same length, the alternate one has five more reversals than the
preferred one.
There are two main groups of Amegilla
(FiS. 42). Six subgenera have SB in males
narrow apically (13, Table 3) and triangular.
In the other five this sternum is broad
apically and quadrate. The small number of
OF THE ANTHOPHORINB
characters
reflects the
results in
change if
covered.
523
BNBS
used in the analysis of Amegilla
homogeneity of this group, and
a cladistic pattern that might
additional characters are dis-
BIOGEOGRAPHY
we have only two fossil
Unfortunately
"Anthophora. " Cockerell derecords for
scribed A. melfurdi (1908) from Florissant,
Colorado. There is no reason to differ with
Cockerell's placement of this Oligocene fossil but not all of the diagnostic wing veins are
clear (for further discussion see Zeuner and
Manning, I97 6). The placement is therefore
"Anthosp€culative. The single specimen of
t
50
14
+44
+13
+7
)
55
37
27
92
1
2
I 32
8 31
3 22
34
28
26
18
10
7
,2
37
8
2 0 48
3 2 40
2
49
58
57 54 48
56 30 34
32 32 32
29
26
2t
1
35
27
6
24
7
49
47
43
26
FIG . +1. Alternate cladogram shown for certain
subgenera of Anthophora, using the same characters as for FiS. 40 but making different assumptions in the main right branch of the cladogram
about which characters are convergent. Explanation as for FiS. +0.
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SysrnuATlcs
AND PHyrocENY
ranges in North America. Lophanthophora
may well have had the same evolutionary
past since it has a similar distribution to that
of Pltganthophora,but lacks arry South African
members and has a widespread range in
both western and eastern United States.
Again like Pltganthophorait does not range
into boreal regions and the New World
dismembers are not morphologically
tinctive, as a group, from Old World members.
The Clisodon group is mainly temperate
Palearctic. The most widespread subgenus,
Clisodon, has only two or three species. One
speciesis Holarctic (Popov, 1951) and inhabits part of northeastern China as well as
northern North AmÇrica. The other one or
two species are Palearctic. Vicariance is an
unlikely explanation of Clisodon's range,
since one species is boreal, occurring well
into Alaska, and thence to northeastern
China. Most likely this species also occurs in
the intervening areas of Siberia. Since the
other members of the Clisodon group are
strictly Palearctic, presumably that is its area
of origin. Petalosternnnpossibly originated in
the Mediterranean region since 18 of its 2l
species occur there.
The Melea group includes five subgenera
with diverse distributions . Rhinomegilla and
Anthomegilla both occur at high elevations in
Asia, mainly in Xizang Province, China
(Tibet) and in neighboring areas. Rhinomegilla is restricted to Xizang Province and
Sikkim . Anthomegilla additionally occurs in
Mongolia and the forest zorte of Siberia.
Paramegilla is widely distributed in the western Palearctic and Ethiopian regions. In
addition, it has two species in the southwestern United States. This distribution
may be explained by vicariance, or by former wider distribution. Neither the Old nor
New World species are boreal and the two
New World species are similar to their Old
World counterparts, not unique as a subgroup in any morphological aspect. Paramegilla has great species diversity from the
Mediterranean region east to the Aral Sea
(at least 40 species). In the Ethiopian faunal
region a single derived species group (about
10 species) of Paramegilla predominates. It
seems clear that the area of origin for Parame'
oF THE ANTHoPHoRINE
BBBs
52s
gilla was in the Palearctic. Anthophoroidesis
found only in western North America, south
to Honduras. There are 12 species (includittg six undescribed ones) which are mostly
limited to the Mojave and Colorado Deserts.
Two species have apparently dispersed to the
south, one reaching Jalisco, México and the
Since Anthophoroides is
other Honduras.
strictly New World and is the sister group of
Melea, both r.rraywell have originated in the
North America. Indeed, of the two species
groups of Melea, the one with the most
ancestral characters is the Bomboides species
group, which has three nearctic and one
palearctic species.
Of the Heliophila grouP, the member with
the most ancestral characters is Caranthophora. It is Palearctic with two disjunct areas,
one Mediterranean and the other Asian.
The Vestita species group of Heliophila is
subsaharan and has its center of species
diversity and probable origin in southern
Africa where the most primitive members of
Heliophila are found. Therefore Heliaphila
probably had its origin in Africa. The Estebana species group is largely restricted to
the southwestern United States, but ranges
to Honduras. (A single specimen in the
Berlin Museum is labelled Guayaquil, Ecuador, perhaps in error.) Southern California and Arizona is the center of species
diversity of the Estebana species grouP (26
species plus one undescribed). Since one if
not all of the species ranging into the neotropics also occur in the southwestern
States, they probably dispersed
United
southward from a possible temperate origin.
Mlstacanthophnra occurs in several disjunct
areas. Its largest, more or less continuous
range is in North America where one species
of the Urbana species group and three species of the Montana species group live. A
monophyletic subgroup of the Montana species group is Antillean. The five Antillean
species are more closely related to two American species than to the other neotropical
members (seven plus one or two undescribed
species). This suggests that Mystacanthophora
invaded the Antilles from Florida. The possible invasion route via the Yucatân peninsula is also less likely because the single
Mexican species is more closely related to
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9Zç
-
SysrnuATrcs
AND PHylocENy
oF THE ANIHoPHoRINB
527
BBns
,al from Asia to Australia could account for
the present Australian fauna.
Eiiaentty major barrie rc to Anthophoradis-
1927.Ueber einigeArten derAnthoptnra
- Gruppe. Entomol. Mitt. 16:120q.uadiJasciata
The only large southern hemisphere Anthophora launa is in southern Africa which is
âccessible from the North via the east Af-
àa.. à"i principaux genres'd'Apoiâea.Anais
da Faculdadede Cienciasdo Porto 56:.l-117.
Banaszak,J. -197.!._
Observations.on.!n1hop_!to1g
,
genus, Mystatanthophora,
regions; it is found in both North and South
AÀerica and tropical speciesexist now in the
Antilles. To accôunt fôr the subgeneric dis-
Batra, S. W. 1'. igOZ. Crop pollination and the
flower relationships of the wild bees of
Ludhiana, India (Hymenoqlela^:.A_poidea)'
Jour' KansasEntomol' soc' 40:164-177'
;-
phora must have moved between eastern and
westernhemispheresat least six times. Most
such dispersal^*as p.esu-ably in the North,
otËer bees. Jour. Iirtrr"r' Eniornol. Soc.
45:208-218.
.1977' Bees of India- (Apoidea)' their
*l
,.
.'
'
'
I.
?
',,'.
'
.
,.
.
;
,
'.
,.
^,!3;^
persathavebe"r,
o""a',u'dlo*Iu,,â;;"pt.;.^ff:fj-fi:::l?#r*e.";i,?Ji;,iL*,m::
ricanhighlands.
otherwi,.o,'rf-o".-'"u- f#:#"i!:i.Slff*l':rA
has crossed tropical
h"rrrirpt""r",-'À*i;- *.;lriâ;l'#;r:*::,'Tià;lr:I:rÏ;rirl*i
persar
in thenorthern
probably
across
theBering
su.it *"., r"iii
involved subgenera that include species that
range into cool to cold climates, The Heliophila case (Humilis
and Estebana species
gro,rps) ho*ever, involves groups ,roï ,"-
j
r
r,
3:l:A:i:';lrumiifff
tothesenirlev
1978. Aggression, territoriality, mating
and nest aggregation- of- some_solitary-.bees
(Hymenoptera:. Halictidae,_ Megachilidae,
c^olletidae' Anthophoridae)' Jour' Kansas En-
,-t'icieito warmand "o,r,r'oJj ;;il ;;
-"*"]if,r"ô.il.i,i;?i3;"titarybee,Anthophora
gions. Vicariance across tJre Atlantic or
more accurately dispersal acrossthe Atlantic
*hen it was much narrower than at present
may well explain the present distribution of
this subgenus. The presence of endemic
Anthophoraspecieson oceanicAtlantic islands
showsthe pôtential for dispersal acrossmoderate ocean barrieis.
The fact that the biogeography of Anthophorini can be largely, or wholly explained
as dispersal, with little evidence of vicariance
th.ough continental movement, suggests
that this is a group most of *rt..i "".i""ù."
occurred after the continents had more or
less attained their present positions.
an6ope,in Punjab, India. Jour. KansasEntomol. Soc. 53l.112-114.
Solitary bees' sci' Amer'
,*.iro!nli:
*""r"r"Ï.
i., C. F".rièr. and O. W Richards.
tggZ. 1.tt. gânericnamesof the British Hymelist of British
nopt€ra Aculeata,with I ch_eck
81-149. Iy'fle GenericNamesof
1ngcjgs,-pp.
British Insects,part 5. Royal Entomol. Soc.,
LTTERAT'RE
crrED
L. .I. R. 1847. Nomenclator Zoologicus'
Agassiz,
-co"ti"""",
.
"
fiî:i:b."*
rià*i"u iv""îliià
ô"nerurÀAni-
malium . . , tur.. 1'2, Indecim Ù;i;rJ;.
j;;ECræ.àÀ,
Soloduri, viii+ 393 pp. (fqt
o" àutJ seeBowléy and Smith, tàOS
.o**"rrt,
and Kevan, 1970).
los HimenAlt;-ilma;.
f i6zl. crta"_go de
opreros de éuba. Instituto C"ubanoàel Libro,
Éavana, 218 pp.
Alayo
-3;;;;b"banos.
Soto, n. iôgZ. Observaciones en Himenl. Apoidea y Sphecoidea.
Pà"yurr., No. 241, 7 ppi
rszo.'.pËii."g "rrr Kenntnis der
Al*eri, ;.'o.
Sen.k"nbe.gi.ttu
Bieneifa'na """ Âgypd.
"
8:96-128.
1919. Stammesgeschichteder
tohiTlt"".
Hauiflûgler. Biol. Zentralbl. 39:145-186.
R. and H. M. Smith. 1968.The dates
Bow-ley,.D.
of publicationof Louis Agassiz'sNomenclator
Jour' soc' Bibliog' Nat' Hist'
t',tftt-tulît'
Brooks,R. W. 1983.Systematicsand bionomics
and speof Anthophora:^The-!ômboidel
F_t_oup
îî"'-.:îiïiiff*î,i:'i9lÏ'Ëiirloi",iii:
Entomol' 98x-+ 8.6^ppt
of the p$oq!o11e
1986.classifrcation
bees.(Hy_meno?tera: Anthophorini). Ph. D.
IJniv' Kansas'- vi+ 325 pp'
-Br9w1,
-}:tit' F. M. 1978- The origins of- the West
Indian butterfly fl"qu. .1.,:F. B: Gill. kd..)'
Zoogeography in the Caribbean. Special Publ.
Acad. Nat' Sci. Philadelphia, pp.
f9a^13,
l-128'
Cane,J. H. 1979. The hind tibiotarsal and tibial
.pr1. artic rlations in bees (Hymenoptera: ApSoc'
Entomol'
Kansas
9i^a91)-'.^Jour'
^52:123-137'
Caldaler,J. 1968a-.Nestsandnestingbehaviourof
Amcgilk (Anugilla) pulchra (Smith) (Hymenop-
-drrcsap qllÀl, 'o"tot1doqruV
srrua8 aql yo sercads
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SysrnuATrcs
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Bnns
s29
. Heft 107,
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SvsreuATrcs
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SysrnuATrcs
AND PHyrocENy
length 9.1 mm. Structure. Inner orbits diverging
above; shortest distance between eyes .7 frontal
length of eye; malar space short; head wider than
long; clypeus protuberant, half eye width; labrum
oF THE ANtHoPHoRINn
Bens
533
apically simple; flagellomere 1 equal to combined
length of next two flagellomeres, two-thirds as
long as scape (excluding basal bulb) and twice as
long as flagellomere 2; flagellomere 2 three-
A-D, A. (M.) bohanimale; E-H, male, I, female,
FIG. 43. New speciesof Anthophora(M2stacanthophorQ.
A. (M.) balumensis.A, B, Genital capsule, side and dorsoventral views. C, D, 57 and S8, ventral views.
E, F, Genital capsule, side and dorsoventral views. G, H, 57 and S8, ventral views. I, Head of female
(hair omitted).
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sHI
it9
SvsrpprATrcs
AND PHyrocENy
times as long as second abscissa of M * Cu; jugal
lobe about one-third length of vannal lobe. Colora'
tion. Brownish black except mouth-parts (including basal two-thirds of mandible), legs, veins, and
metasomal sterna reddish brown; clypeus with
pale integumental mark (FiS. 43i); wings clear.
Pubescence.Ochraceous Buff, except paler on face,
pleura and thoracic sterna; a few dark hairs near
posterior ocelli, center of scutum, or visible
portions of T2-T6, and dense around pygidial
plate; pile on head, thorax (except propodeal area
where length is slightly longer), T1 and sterna
about equal in length to scape and erect, Posteriorly inclined; hair on T2-T6 shorter than scape;
apical bands of hair on terga absent. Punctation.
Face dull; clypeal, paraocular and supraclypeal
punctures
ground
between
areas lacunose,
strongly microareolate; labrum rugose, ground
between rugae shiny, smooth; upper face somewhat shining, punctulate, ground between puncscutum and pleura dull,
tures microareolate;
finely microareolate; propodeal area weakly shining, with scattered punctures, ground between
punctures microareolate; T1-T4 dull, puncticulate but apical margin impunctate, width of impunctate margin becoming wider from T1-T4,
ground between punctures microareolate, T5-T6
punctate, punctures almost confluent to 1 puncture-width apart, ground between punctures microareolate; 51-56 shiny, with scattered punctures
becoming more densely spaced from 51-56,
ground between punctures microareolate.
Male: Length 10.7 ffiffi, forewing length B mm.
Agrees with description of female except for sexlimited characters and as follows: Structure and
1 equal to combined
Pubescence. Flagellomere
length of next 1.6 flagellomeres together; hind leg
simple, basitibial plate absent. Pygidial plate
absent, apparent medioapical margin with two
blunt lobes; 56 broadly emarginate apically with
a medioapical protuberance and lateral depression; 55 apically entire with four patches of black
hair, a smaller submedian pair and a larger patch
lateral to submedial ones (Fig. +4ù; basal twothirds of disc from gradulus distally with a thick,
dense pad of white hair; 54 apicully entire; 57, SB
and genital capsule as in Figures 43e-h. Coloration.
Basal third of mandible, labral, clypeal, paraocular and supraclypeal areas and scape anteriorly
white. Punctation. Labrum shiny, lacunose, with
ground between punctures smooth.
TyPe materiaL Holotype female: Bahama Islands, New Providence, Nassau, 28 June 190+,
collected by Allen, Barbour, and Bryant [Cambridge]. Allotype male: Bahama Islands, Nassau,
British Museum 1931-343, collected by G. Salt
[London].
species is named for the
Etymologlt. This
Bahama Islands.
Comments. This species is closely related to ,4.
tricolor (Fabricius) from which it is separated in
the female by a pale mark on the clypeus and in
the male by a pale mark on the mandible and the
oF THE ANrHopHoRrNn
535
BBns
strong median, longitudinal rounded ridge on 56.
The shape of 57, SB and the genital capsule
distinguish A. bahamensis from other Antillean
Anthophora. The species is restricted so far as
known to New Providence, Bahama Islands (24.
tricolor does not occur there).
Anthophora
(Anthophoroides)
new sPecles
signata
Diagnosis. Female with white, integumental,
inverted T-shaped mark on clypeus and triangular mark on labrum; male with greatly modified hind l.g, tibia dilated with large anteroapical
spine and basitarsus gre atly flattened with large
basoapical spine (Fig . 44e).
10 mm; forewing
Description. Male: Length
length 7 .4 rnrn. Structure. Inner orbits diverging
above; shortest distance between eyes about equal
to length of eye; head wider than long; clypeus
protuberant, in profile about equal to eye width;
flagellomere 1 equal to combined length of next
1 .6 flagellomeres together, .52 as long as scape
(excluding basal bulb), I .6 times as long as
flagellomere 2, and .92 times as long as last
flagellomere, flagellomeres 2-10 about equal in
length and about 1.6 times longer than wide;
distance between posterior ocelli equal to 1.5+
median
from
distance
distance;
ocellocular
ocellus to posterior ocellus equal to .69 ocellocular
distance, labrum with medioapical notch; hind
l.g modified (FiS. 44e), with femur and tibia
dilated, bulbous, apex of tibia with large, anterior
spine with outer tibial spur inserted on its apical
third, hind basitarsus gre atly flattened and with
large blunt tooth and apicoanterior small acute
tooth; base of inner hind tibial spur partially
enclosed, enclosure forming a small, sharp tooth
against spur; second submarginal cell receiving
first recurrent vein slightly distal to midpoint;
marginal cell length .67 distance from apex of cell
to wing tip; hind wing with cu-v 1.2 times length
of second abscissa of M + Cu; vein M 3.0 times as
long as second abscissa of M * Cu; jugal lobe .+
length of vannal lobe; terminalia as in Figures
44a-d. Coloration. Black except apices of tarsomeres and claws ferruginous; basal half of mandible, labrum (except translucent basolateral spot),
clypeus (except anterior tentorial pit), paraocular
and supraclypeal areas, and scape anteriorly
white; paraocular area extends from malar space
to upper margin of antennal socket; wings clear,
veins reddish black. Pubescence.Hair white except
a few scattered black hairs on vertex; pronotal
lobe, scutum, 52-56, T5-T6 medioapically, T7 all
white, and ferruginous hair on inner surfaces of
tarsi. Length of hair on face, foretarsus, foretibia,
hind femur, thorax, metasomal sterna and Tl
slightly longer than length of scape, hair on
midleg about length of scape or shorter, hair on
forefemur and scopa long, about equal to length
of hind tibial spur, hair on visible portions of T2'
T5 shorter than length of scape and directed
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9t9
SvsrnuATrcs AND PHvrocENY
rated by 0.5-1.0 puncture-widths, ground between punctures smooth to weakly areolate; S1-3
with scattered punctures, 54-56 with punctures
becoming denser near apical margins, apical
margins impunctate, ground between punctures
microareolate.
Female:Length 10-11 mm; forewing length 7-B
mm. Agrees with male description except for sexlimited characters and as follows: Strzcture.Inner
orbits diverging above; shortest distance between
eyes about equal to length of eye; clypeus protuberant, in profile .75 eyewidth; flagellomere 1
equal to combined length of next 2.7 flagellomeres, .67 as long as scape(excluding basal bulb)
and 1.4 times as long as last flagellomere,flagellomeres 2-9 about equal in length and slightly
longer than wide; distance between posterior
ocelli equal to 1.25 ocelloculardistance,distance
from median ocellus to posterior ocellus equal to
.56 ocellocular distance. Coloration.Black except
apices of tarsomeresand claws ferruginous; clypeus and labrum with white integumental marks,
"T," of
that of clypeus shaped like an inverted
T6 surrounded balabrum triangular. Pubescence.
sally with black and laterally white. Punctation.
Face shiny; clypeus with large punctures separated by 0.3-3.0 puncture widths, ground between punctures weakly microareolate; clypeus
with median portion lacunose, laterully rugulose
to more finely punctate 4t. margin, ground between punctures near margin microareolate; rest
of face foveolate, ground between punctures
smooth.
TyPt materiaLMale holotype: California, Los
Angeles Co., 2.5 mi. (4 k*) S. Pearblossom,
3500 feet (1067 *), 14 May 1978, collected on
Salazariamexicanaby R. W. Brooks; female allotype with same data [Lawrence]. Paratypes:
CALIFORNIA: Inyo Co; Argus Mts., 1 female,
31 Muy 1940 (D. Meado*r), 2 females,4 June
1939 on Penstemon
(R. Bohart); San Lucas Canyoû, Inyo Mts., 5800 feet (1768 m), 2 males,22
June l97B (T. Griswold); Surprise Canyon, Panamint Mts. , 4 males, 24 April 1957 on Peucephyllum
schottii(J. Powell, P. Hurd), 1 male, 28 April 1953
(G. Marsh), 1 male, 9 Muy 1958 (L. Stange);
Wild Rose Canyon, Panamint Mts., 1 male, l+
Muy 1969 (P. Opler). Kern Co: Last Chance
Canyon, El Paso Mts., 2 males, 15 April 1964
(R. Snelling). Los AngelesCo: Desert Springs, 2
females, 5 May 1956 (J. Powell); Little Rock, I
female, 22 April 1950 (C. MacNeil); Mojave
Desert, Highway 138,3600 feet (1097 rrr),2
females, 28 April 1947 on Salaia dorrit (P. Timberlake); Palmdale, 1 male, 15 April 1956 on.
Eriogonum(R. Erdmarr); 15 mi. (25 km) southeast Palmdale, San Gabriel Mts., 2 females, 22
April 1972 from burrows (D. Bixler); 2 mi. S.
(3,2 k*) Pearblossom,3500 feet (1067 *), 1
male, 18 April 1977 (R. Snelling), 3 females, 1-2
Muy 1977 on Salazariamexicana(R. Snelling); 2
females, 17 Muy 1977 on Salazariamexicana(R.
Snelling); 4 mi. (6.+ ktn) S. Pearblossom, 2
OF THE ANTHOPHORINB
BNNS
537
femalesand 3 males, 13-14 April 1960 on Salaia
(R. Snelling); 5 mi. (B k-) S. Pearblossom, I
male, 23 April 1956 on Tamarix gallica (8.
Linsley); 6 mi. (10 km) SE. Pearblossom,1 male,
16 March 1972 (R. Snelling). RiaersideCo: Blythe,
1 male, 2 April I94l (J. MucSwain, E. Linsley);
Joshua Tree National Monument, Quail Springs,
1 male, 12 April 1950 (P. Hurd) . San Bernadino
Co., 10 mi. (16.6 km) S. Adelanto, 1 female, 3
May 1939 (P. Timberlake); Apple Valley, 1 female, B May 1958 on Gilia latif,oradaayi(P. Hurd),
1 female, 9 Muy 1958 on Phacelia distans (P.
Timberlake), 1 female, 9 Muy 1955 (W. Richards); Hesperia,1female,30 April 1939 on
axillaras(P. Timberlake); Kramer Hills,
Tbtradymia
7 mi. (11.6 km) SE. Boron, 1 female, !!_Ap_ril
1958 (I Powell), 1 female, 10 Muy 1955 (W.
Richards); Lucerne Valley, 1 female, T May 1938
(C. Dammers); Morongo Valley, 1 female, 18
April 1955 (W. Richardt); San Bernadino Mts.,
3800 feet (f 158 m), 1 male, 15 Muy 1937 (E,
Linsley);7.5 -i. (12 km) S. Twentynine Palms, 1
female, 7 Muy 1948 on Salazaria mexicana(P.
Timberlake); 3 mi. (5 k*) SW. Victorville, 6
females, 6 May 1939 on Daleafremontii saundersii
(P. Timberlake), 2 females, 12 _Vly_1939^.on
Haplopappuscooperi(E Linsley). NEVADA: Clark
Co: Charleston Mts., Lee Canyon, 2 females, 25
Muy 1940 (G. Bohart). Ten paratypes deposited
at [Berkeley], t7 [Brooks], 1 [Layrence], 5
[Logan], 13 [Los Angeles], ? [Ottawa], _1
-[Puùmân], 12 [Riverside] and 1 [San Francisc_o].
Ety*olàgt: This species is named {gnqta because of the white clypeal mark on the female.
Anthophorasignata and the subgenus
Comments.
Clisodonare the only known speciesof Anthophora
to nest in wood. At Pearblossom,California, the
species nests in old Joshua tree stumps (Yucca
breuiftlia)(R. Snelling, personal comm.) but apparently also burrows in the ground (see d-ata
from 15 mi. SE. Palmdale). This speciestogether
(restricted to New Providence,
with 24. bahamenszs
Bahama Islands) are the only known New World
Anthophora(except Heliophila) whose females have
white integumental facial marks . A. signatacan
since it does
easily be separatedfrom bahamemas
not have apical pale integumental bands on the
metasoma. A. signatais most closely related to
another undescribed speciesand A. phaceliaelrom
which it can be separatedby the unique shapesof
S4-SB, the genital capsule and hind l.g of the
male (Figs. 44a-e). Especially notewgrtly is the
combinaiion of characters of the male hind leg
(FiS. 44e), that is the apical spile on the tibia is
bidèntate, the tibia is strongly dilated, the basitarsus greatly flattened with the long basal tooth
wide, almost bidentate at apex.
Anthophora (Anthophoroides) phaceliae
new specles
abruptella,Marston, 196+, p. 101 (misiAnthophora
dentification).
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SvsreNIATIcs
AND PuvrocENY
ffi:*:ï,ï;:::'ii#lliï;
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upper anterior surface concave; terminalia as in
Figures 45a-c, e. Second submarginal cell receiving first recurrent vein slightly distal to midpoint;
marginal cell length 0.80 distance from apex of
cell to wing tip; hind wing with cu-v 0.77 times
length of second abscissa of M + Cu; vein M 2.72
times as long as second abscissa of M * Cu; jugal
lobe .4 length of vannal lobe. Coloration. Facial
markings as in A. signata male. Black except basal
half of tarsal claws ferruginous; tegula amber;
wings clear, veins reddish brown; legs, 51, 52 ?nd
morè or less 53 dark ferruginous. Pubescence.H.air
pale except a few scattered dark hairs on supraclypeal areabetween antennae and vertex. Scutal
pubescence dense, pale with a few scattered dark
hairs; rest of thorax, fore and midtibiae all pale;
pubescence of head and thorax about equa] to
length of scape; that of fore and midlegs all black
except a pale patch on outer apices of tibiae; hind
l"g 6lack except pale hair on outer surface of
coxa, a pale patch on outer apices of hind femur
and tibia; hind basitarsus with a few long pale
hairs on outer surface, outer posterior edge of rest
of tarsomeres with some pale hair. Metasoma
with sparse erect black hair up to length of
flagellomere 1; terga with a few pale hairs laterally, T4 and T5 with a sparse weak row of long
pale hairs ulglg b.as{ edge.of apical impunctate
margin; medioapical margin of T7 with some
pale hair. Punctation. Face as in A. signata, weaHy
shining. Thorax dull, punctation confluent, propodeaf area weakly shining. Metasoma as in 24.
signata except terga weakly shining.
Female: Agrees with ynale description except for
sex-limited characters and as follows: Length
about 11 mm; forewing length about 8.6 mm.
in profile .67 eye
Structure. Clypeus protuberant,
width; flagellomere 1 equal to slightly more than
combined length of next three flagellomeres' .79
as long as scape (excluding basal bulb) and 1 :59
times as long as last flagellomere, second flagellomere .31 length of flagellomere t , flagellomeres
3-9 about equal in length, longer than wide,
individual segments .37 length of flagellomere 1;
distance between posterior ocelli l.l4 ocellocular
distance, distance of median ocellus to posterior
ocellus equal to .32 ocellocular distance. Colora'
tion. Black except basal halves of tarsal claws
ferruginous and tegula amber. Pubescence.Upper
third of pleura and propodeum with dense pale
hair, rest of thorax with dense black hair; that of
legs all black except pale patch on outer apices of
tiSiae; appressed scale-like pubescence of basitibial plate golden. T1 with sparse Pale pubescence
about length of scape; rest of metasoma with
sparse erect black hair up to length of flagellomere 1. Punctation. Face as in 24. signata female.
TyPt materiaL Male holotype and allotype: California, Solano Co. , 3.2 mi (5 k*) W. of Mix
Canyon, (about B km) NW. of Vacaville, 21 April
1976, both collected on Phacelia egena (R. W.
Brooks), holotype caught at 0900-0915, allotype
OF THE ANTHOPHORINN
BNNS
539
caught at 1330- l3+5 hrs. [Lawrence]. Paratypes:
CALIFORNIA: ContraCostaCo: Mt. Diablo, 1
female, 2I Muy 1937 (E. Ross). FresnoCo: 6 mi
(9.6 km) S. Coalinga, I female, 30 March 1970
0. Powell); 5 mi (B km) S. Coalinga, 1 male, I
female,17 March 1975(J. Powell); 4 mi (6.3 k*)
N. Orange Cove, 2 males, 1 female, 6 April 1957,
on Phacelia(R. Snelling, M. Stage). Kern Co:
Woody, 1 female, 29 March 1951, (J. MacSwain) . Madera Co.'San Joaquin Experiment Station, 1 female, 20 April 1953 (H. Childs).
MariposaCo: Exchequer Dam, 1 male, 1 female, 9
(R. Snelling),
April 1960, on Phaceliaramosissima
4 mi (6.3 k-) W. Echequer Dam, I male, 10
(R. Snelling); Mariposa, 1
April 1960, on Phacelia
female, L+ Muy 1960 (R. Snelling); Miami
Ranger Station , I male, 2 June 1937 0.
Kliewer). MontereyCo.'Arroyo Seco Camp, 5 mi
(B km) W. Greenfield, Elev. 1000 feet (305 m), 1
female, 15 Muy 1976, oD Eriodictyoncal'ifornicum
(R. Brooks), I female, 1 Muy 1957(P. Torchio), 2
females, 2 males, 5 M.y 1956 (D. Ribble, P.
Torchio). Riuerside Co: Idyllwild, San Jacinto
Mts., 1 male, 18 June 1952 (M. Cazier, et al.).
SantaBarbaraCo: Gaviota Pass, 1 female, 23 April
1966, (J. Powell). San Benito Co: San Benito, 1
male, 24March 1971(E. Cane). SolanoCo: 3.2 mi
(5 km) W. Mix Canyon, Elev. 1700 feet (518 m),
4 females, 17 April 1976, or Phaceliaegena(R.
Brooks), 6 fernales, 18 April 1976, 4 females, 1
male, 19 April 1976,6 females, 1 male, 21 April
1976, 1 female, 30 April 1976, 2 females, 3 May
1976; 2 mi (3 km) W. Mix Canyon, Elev. 1000
feet (305 m), 1 female, 16 April 1976, oû Phacelia
egena(R. Brooks), 3 females, 17 April 1976; 1.5
mi (2 km) W. Mix Canyon, Elev. 1000 feet (305
*), 2 females, 10 April 1976, 3 males, I female,
14 April 1976, 1 female, 16 April 1976, 1 female,
17 April 1976 (R. Brooks), 1 fem ale, 2 Mty 1976
(5. Armbruster) . Stanislaus
on Collinsiaheterophylla
Co: Del Puerto Canyon, 1 female,20 April t949,
(P. Hurd); Knight's Ferry, I male, 10 April 1961,
on Phacelia(R. Snelling), 1 male, 1 female, 15
April 1961 (R. Snelling, T. Duncan). TulareCo:
Lemoncove, 1 female, 14 April 1950, (I. MacSwain). Yolo Co: Putah Canyon, 2 males, I
female, 1 May 1940, on Phacelia(G. Bohart).
Unknown Co: Midway, 1 female, 24 Muy 1936,
(M. Cazier). One paratype deposited at [Beqkel.yl , 45 [Brooks], 1 [Cambridge], 1 [Davis], 5
[Lawrence],5 [Logut], 12 [Los Angelesl, 2 [San
Francis.o], and 3 [Urban?].
MEXICO: Baja CaliforAdditional specimens:
nia, Sierra San Pedro Martir, Rancho Viejo,
Elev. 7000 feet (2134 m), 1 female, 13June 1953
(P. Arnaud).
Comments.The female from Baja California
differs from Californian populations in the admixture of several black with pale hairs on
scutum, pde pubescenceon upper two-thirds of
pleura, lower third of pleura and sterna with
black hair, outer apices of tibiae with a patch of
amber hair, fore and midlegs with pale hair
except anterior margin of tarsus with black hair,
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SysrBruATIcs
AND PHyrocENY
oF THE ANtHoPHoRINn
Bnns
5+l
7i,n:,:|;
Ti,;
ii
('r'ûli
ffi
ffi,:,*i
Tllt:mn
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FIG. 46. Anthophora(CaranthophorQspinacoxa,male holotyPe. A-C, Outer views of fore, mid and hind
legs. D, Anteriàr view of hind femur, base of femur at top. E, Hind coxa, side view, base of coxa at toP
orirightside. F, Head, frontalview. G, 57, ventralview. H, S8, ventralview I,J, Genitalcapsule,
dorsoventral and side views.
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9S pue el"ul8r?rue
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-paurocrde ?S ialeptue
pue padolarrap 1e/r\ LL
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padolenep [e/v\ e gJ Jo ]eqr 'qtoo] )Fe/v\ s o]ul
padolar'ep çI Jo ssacord repper8 :(8-pm 's8l.f)
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sHJ
ztç
SYstnvtATIcs AND PHYTocENY oF THE ANTHOPHORINB BNNS
543
and without median subapical process; flagellomere 1 equal to combined lengths of next 1 .6
flagellomeres, 0. 5 1 as long as scape (excluding
basal bulb) and 1.9 times as long as flagellomere
2, second flagellomere 0.69 times as long as
flagellomere 3, flagellomeres 3-10 about equal in
length, each | .22 as long as wide, flagellomere 1 I
subequal in length to flagellomere l; distance
between posterior ocelli about 1 . 15 ocellocular
distance; distance from median ocellus to posterior ocellus 0.65 ocellocular distance; labrum
simple without median apical or median subapical protuberances; cu-v of hind wing about
equal in length to second abscissa of M * Cu; vein
M 3.53 times as long as second abscissa of
M + C.t; jugal lobe about one-third as long as
vannal lobe- 54-36 with dense pads of hair; 57,
SB and genital capsule as in Figs . 4Bc, f-h.
Pygidial plate slightly emarginate apically without sublateral tooth (Figs. 4Bd, e). Coloration.
Black except facial marks bright yellow (FiS.
*Ba). Pubescence.Head and thorax on lower half
white and upper half yellowish-white; posterior
half of scutum with a few scattered dark hairs.
Foreleg all white on outer surface, but dark on
inner surface; mid and hind legs with hair all
black except white on coxae and trochanters and
fuscous on posterior surface of midtibia. T1-T6
with appressed orange hair but white to yellowwhite laterally; metasomal sterna with white hair,
becoming fuscous medially from 53-56 . Puncta'
tion. Punctures on clypeus and paraocular area
fine and separated by 0.5-2.0 puncture widths but
fine and nearly contiguous on rest of head.
Thorax and metasomal terga dull, weakly shinpunctation contiguous to nearly contiguous_.
itg,-Female:
Length 12 mm, forewing length 8.2
mm. Agrees with description of male except for
sex-limited characters and as follows: Structure.
Shortest distance between eyes 0.91 length of eye;
flagellomere 1 equal to combined length of next
thrèe flagellomeres and equal to 0.58 length of
scape (excluding basal bulb); flagellomeres 2-9
about equal in length and individual segments
about as long as wide; last flagellomere 0.77 as
long as flagellomere 1; labrum simple without
median apical or subapical protuberances. ColoraAnthophora (Heliofuhll") miclwneri
tion. Facial marks as in Fig. 4Bb. Pubescence.T5
new sPecles
with median patch of dark hair; S1-S3 with sparse
apical fringe of white hair; 54 with apical fri1g9 gf
Diagnosis.Body almost completely covered
with orange hair similar to that of Anthophora long white-hair and sparse subapical band of dark
haii; 56 with dark hair medially, white hair
aestitaSmith; mid and hind legs black except
laterally.
trochanters and coxae with long white hair (male)
male and allotype:
Tlpt materiaL Holotype
or mid and hind femur with mostly white hair
Kenya: 20 mi. [32.2 km] NE. Magadi, 3500 feet
(female); middistitarsis of male without lateral
16 June 1967, C. D. Michener
fringe of black hairs; male and female with bright
[1066.8 *],
fringe of black hair (FiS. +7j); hind leg with all
black except white on inner surface of tarsus
(from apex of basitarsus to claw). T1-T3 mostly
with black hair but Tl with a few scattered
orange hairs and T3 with appressed narrow
apical white band; apical half of T4-T6 with
white, T7 with orange-brown hair; 51 with pale
brownish hair; 52 and 53 with brownish hair, 54
with short brownish hairs apically and laterally
but median area (not reaching apical margin)
with dense mat of reddish brown hair; 55 with
short brownish hair laterally and a narrow band
apically but median area (reaching apical margin)
with densemat of yellow orange hair; 56 laterally
and broadly apically with short brownish hair but
median area (reaching only to apical third) with
densemat of yellow orange hair. Punctation.As in
A. micheneri.
Tyfe materiaLHolotype male: Angola: 45 mi.
172.4kml S. Santa Combre Dâo, 3 April 1970,
Edward S. Ross [San Francisco]. One paratype
male: Zaire (8. Congo): 18 mi. [29 km] NW. of
Tshinsenda, B February 1958, 1330 m altitude,
E. S. Ross, R. E. Leech collectors [Lawrence].
This distinctive specieslooks superComments.
ficially much like Amegilla (Zebramegilla)atribasis
(Cockerell). There is no other speciesthat resembles it in Africa. There are differences between
the Angola specimen (holotype) and the Zaire
specimen (paratype). The holotype has a distinctive apically emarginate pygidial plate (FiS.
+7ù and the gradular process of T6 developed
into a tooth whereas the paratype has an almost
apically entire pygidial plate (FiS. 47d) and the
gradular processof T6 is rounded and not developed into a distinct tooth. The genitalia, howand although these
ever, are identical
morphological differences are striking, I believe
that jh.y are within the range of variation of this
specres.
Erymologlt This species is named for Joetta
Weaver in recognition of her 25 yearsof painstakirg editorial and secretarial contributions for
Charles D. Michener and his many students and
associates.
yellôw markings on face (Figs. 4Ba, b); 57, SB
and genital capsule as in Figs. 4Bc, f-h.
Description. Male: Length 13 mm; forewing
length 7.6 mm. Structure.Inner orbits diverging
above; shortestdistancebetween eyes0.79 frontal
length of eye; malar spacelinear; head wider than
long; clypeal protuberance 0.+2 eye width; mandiblt with subapical tooth; labrum apically simple
-[Lawrence].
Commenis. This species looks very similar,
though smaller in size, to Anthophora aestitaSmith.
The male of uestita has a well developed lateral
fringe of black hair on the middistitarsus and a
deeply emarginate pygidial plate, whereas michen'
eri [acks the fringe of hair on the middistitarsus
and has a shallowly emarginate to almost apically
sprs
pu"
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I
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'Tcelq
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yo s8al pqq
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[[" eæ zrauaqztw
sade ueeûuaq erustsrp tseuor{s lanoqe turÊraÀrp
eql seeJer{/v\'saceJrnsJeuur eqt uo atuero pu?
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-eroJ lurur gI tnoqs rptual :apry 'uoq(ulsaq
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It9
SysrnuATlcs
AND PuYrocENY
ocellus to posterior ocellus 0.60 ocellocular distance; cu-v of hind wing equal to 0.67 length of
vein M 2.3 times as
second abscissa of M*Cu;
long as second abscissa of M * Cu; j"S4 lolg
about one-third as long as vannal lobe. 57, SB
and genital capsule as in Figs. 49c-f. Coloration.
except facial marks dark yellow
Brownish-black
(yellow markings on holotype and paratype reddened, presumably due to cyanide exposurg) and
tarsi amber-brown. Pubescence.Lower half of head
and thorax ochraceous, upper half red. Foreleg
with ochraceous on outer surfaces and brown on
oF THE ANtHoPHoRINn
Bnns
5+5
inner surfaces; midleg with ochraceous hair except midbasitarsal brush and hairs on posterior
margin dark-brown and lateral hair fringe on
middistitarsus fuscous, inner surfaceof basitarsus
brown; hind leg with ochraceoushair, but basitarsus dark brown on inner and outer surfaces
(except apex ochraceous). T1 and T2 covered
with long, shaggy orange hair; T3 and T+ on
apical third with light orange-ochraceoushair and
a few scattereddark hairs; basal two-thirds of T4
with short brown hair; T5 and T6 with mostly
long, shaggy, dark hair, laterally with lolg,
adamsorum,male paratype. A, B, outer views of hind tarsus and mid
FIG. 49. Anthophora(Pyganthophora)
tibia and tarsus. C, 57, ventral view. D, SB, ventral view. E, F, Genital capsule, side and dorsoventral
views.
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dpsoru rleq i1ce1qIF elerual Jo ered 'stsou7otq
lB euo ldacxe BoueC le pelrsodap sad(lered 11V
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001,-00? le ]sntnv q petrallor
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000I-009'868I
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p1euo'U roJ petueu sI satcads sHI '(îo1ow(gg
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41
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dlreeu pue JeuUrlontu seJnlJund leare JelnJoered
pue snaddlc uo sqlpr/v\-ernlcundç'0 dq palerudes
ol snon8rluoo dlreau sernlJund 'uoxlopund 'rletl
snoeJerr1oo'd8Eeqs '8uo1 qllû\ euJels letuoseleur
lrreq atuero uogs qllrv\ LL lrreq snoacerr{ro
Nrrs-rrn8 scNsrss svsNvx .{o Àrrsus^INn sHI
SvsrnuATlcs
a
^
!
I
I
AND PHvrocENY
ble Burnt Sienna; mouthparts and lateral surface
of flagellum Cinnamon- Rufous. Thorax black,
wings
legs and wing veins Cinnamon-Rufous,
clear. Metasomal terga and sterna with visible
portions of discs black, margins CinnamonRufous . Pubescence.Hair on face pale but mixed
with some black on vertex, hair densely spaced
obscuring paraocular area but rest of facial surface visible. Scutum with pale and black hair
intermixed; pleura almost all pale with occasionally only a few black hairs; thoracic sterna,
propodeal area, mid and hind coxae, trochanters
and femora pale; forefemur with long Fuscous
hair; tibiae and tarsi Mars Yellow. Metasomal
terga Mars Yellow with lighter apical bands of
'I5
with a dense median patch
Warm Buff except
of Fuscous surrounding base of pygidial plate,
hair on terga appressed with some semi-erect hair
inclined at most as a 45o angle; metasomal sterna
Mars Yellow. Punctation. Face weakly shining;
labrum shiny, rugose, rrgae smooth; clypeus
spaces again micoarsely areolate-verrucose,
nutely microareolate; rest of face somewhat more
shiny than clypeus, less coarsely microareolate,
spaces smooth. Scutum weakly shining, less dull
in center, densely punctate at margins, punctures
apart, but in
contiguous to .5 puncture-widths
middle of scutum punctures separated by 0.5-6.0
punctures
ground
puncture-widths,
between
finely alveolate; pleura, scutellum and metanotum weakly shining, densely punctate, puncapart,
tures contiguous to .5 puncture-widths
ground between punctures finely microareolate;
propodeal
area dull, finely alveolate. T1-T4
shiny, punctate except apical margin, separation
ground
of punctures 0.5-3.0 puncture-widths,
between punctures minutely microareolate; T5T6 coarsely areolate, ground between punctures
minutely microareolate; sterna shiny, punctate,
separation of punctures I .0-5 .0 puncture-widths,
apical margins and gradular area impunctate but
these areas as well as ground between punctures
minutely alveolate, alveoli often in rows, appearing striate.
UPt materiaL Female holotype: Cape Verde
Islands, Boa Vista, December 1897 , collected by
Leonardo Fea [Genoa]. Paratypes: Twelve females with same data and eight additional females collected in February 1898. All paratypes
deposited at Genoa except two paratypes at Lawrence.
Commenfs.This species is similar in appearance
to A. africana and capensis but is much smaller in
size. Besides being presumably endemic to the
Cape Verde Islands and being presumably the
only yellow Amegilla there, it is the only yellow
species of Amegilla with an all black female face.
Etymologlt; This species is named after the Cape
Verde Islands.
Amegilla
(Afrarnegilla)
robinae new species
Diagnosis. Face of male with pale yellow
markings as in
(Aniline Yellow) integumental
OF THE ANTHOPHORINN
BBNS
547
Figure 50f; T1-T5 with apical hair bands pale
Russian Blue; terminalia as in Figures 50a-d.
13 mm; forewing
Description. Male: Length
length 9.2 mm. Structure. Inner orbits diverging
above; shortest distance between eyes .B length of
eye; malar space linear; head wider than long;
clypeus protuberant, in profile .6 width of eye;
galea short, reaching slightly posterior to forecoxa; flagellomere 1 equal to combined length of
next 2.7 flagellomeres; .B as long as scape (.*cluding basal bulb) and 1 .5 times as long as last
flagellomere; flagellomeres 2-10 about equal in
length and about 0.9 times as long as wide;
distance between posterior ocelli equal to 1 .1
from
median
distance
distance;
ocellocular
ocellus to posterior ocellus .+4 ocellocular distance. Marginal cell length equal to distance from
apex of marginal cell to wing tip; vein cu-v of
hind wing .B length of second abscissa of M * Cu;
vein M 2.9 times as long as second abscissa of
M + Crr; j.rgul lobe a little less than half length of
vannal lobe. 55 broadly and shallowly emargiapicomedially
nate, 56 shallowly emarginate
(FiS. 50e) but apical margin only slightly deflexed
ventrally otherwise disc gently convex; 57 apically round, disc with well developed lateral tooth
on basal half (FiS. 50d); SB apically rounded (Fig,
50c); T7 apparently bidentate medioapically but
margin between bases of teeth weakly produced
medially appearing weakly tridentate (Fig. 50e),
without median longitudinal ridge; penis valves
small; gonocoxite with well developed inner lobe;
genital capsule from dorsal view rounded in
outline (Figs. 50a, b). Coloration. Black except
integumental facial marks Aniline Yellow (FiS.
50f), mouthparts, tegula and S1-S2 Burnt Sienna. Pubescence.Face with pale hair except vertex
Brown
Buckthorn
Brown. Thorax
Buckthorn
with a few black hairs intermixed, sterna pale;
coxa pale; foreleg black except inner surface of
tarsus Burnt Sienna, outer surface pale; midleg
like foreleg except only pale on outer surface of
tibia and basal half of basitarsus; hind leg like
foreleg except only pale on outer surface of tibia.
T1- T5 with apical bands pale Russian Blue with
a few scattered dark hairs; rest black. Punctation,
Clypeus dull, coarsely punctate, punctures separated by 0.5-1.0 puncture-width, ground between
punctures coriarious-areolate; labrum shiny, rugulose-lacunose, ground between pits alveolate;
rugulose-lacunose,
shiny,
areas
paraocular
ground between pits weakly microareolate; supraclype al area to posterior ocelli coarsely, densely
punctate, punctures separated by 1 punctureScutum and scutellum
width to contiguous.
weakly shining, coarsely punctate, punctures septo contiguous;
arated by 0.5 puncture-width
pleura shiny, coarsely punctate, punctures separated by 1.0 puncture-width to contiguous; metadull, finely punctate,
notum and propodeum
punctures contiguous. T1-T5 dull, puncticulate,
ground between pu-nctures microareolate, fPical
margin impunctate but microareolate; S1-S2 with
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AND PHvrocENy
oF THE ANrnopHoRrNB
Bees
549
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SYstEIraATIcs AND PuvrocENY
bles, posterior surfaces of legs and wing veins
Claret Brown, wings slightly infuscated, not
clear. Pubescence.Face, including gena, on lower
two-thirds covered with white hair such that
surface is obscured but surface of labrum visible,
white hair with some black around vertex of head;
scutum and upper fourth of pleura pale with
black hair intermixed, rest of pleura and thoracic
sterna Fuscous; legs Fuscous except outer surfaces of tibiae light Orange-Yellow, apices of mid
and hind tibiae with white hair; propodeal area
and T1 covered with long pale hair, rest of
metasoma black. Punctation. Face weakly shiny;
clypeus, paraocular and supraclypeal areas microareolate; labrum areolate-rugose, rugae minutely microareolate; vertex punctate, punctures
scutum
separate by 0. 5- 10.0 puncture-widths;
dull, alveolate and within and between alveoli
minutely alveolate; pleura and scutellum dull,
obscurely variolate, ground between indentations
minutely microareolate; propodeum dull, posterior two-thirds microareolate, anterior third minmetasoma dull, punctate,
utely microareolate;
punctures separated by 1 .0-6.0 puncture-widths,
ground between punctures weakly imbricate,
with narrow apical impunctate margins.
Female: Length 10.7 -13.2 mm, forewing length
7.8-9.2 mm. Agrees with male description except
for sex-limited characters and as follows: Stracture.
Flagellomere 1 equal to combined length of next
three flagellomeres together, .78 as long as scaPe
(excluding basal bulb), and 1.6 times as long as
to
Pubescence. Legs black
last flagellomere.
Fuscous, with pale hair on outer basal surface of
midtibia, and posterior to pencillus on hind tibia;
T1, in
scopal hair Fuscous to Cinnamon-Rufous;
addition to covering of long pale hair, with
narrow band of appressed pale hair on apical
margin, sometimes T2 and T3 with similar appressed bands of hair but interrupted medially.
Typt materiaL Male holotype: Cape Verde Islands, Sâo Tiago, Orgâos Grandes, Muy 1898,
collected by Leonardo Fea; female allotype with
same data [Genoa]. Paratypes: With same data
except 2 females collected in March, 18 females
and 1 male collected in April, 19 females and 3
males collected in Muy. All paratypes deposited at
Genoa except two males and two females at
Lawrence.
Commenls. This species is similar in apPearance
to A. godofredi but is smaller in size and the scopa
is pale, not orange. Besides being presumably
endemic to the Cape Verde Islands, it is the only
Amegilla which has a black face in both sexes and a
black body with pale hair on the head, thorax and
T1.
Etymologlt:This speciesis named after a similar
looking species, Amegilla modesta(Smith) (: A.
godofrediDours), which is also endemic to the
Cape Verde Islands.
Amegilla (Glossamegilla) lieftincki
new sPecles
Female
with Beryl Blue mixed with
Diagnosis.
oF THE ANrHoPHoRrNn
BBBs
551
black and white hair on head, thorax, outer
surfaces of fore and midtibiae, and apical margins
of T1 -T4; face of female with only small triangular pale mark on apicomedian half of clypeus; Beryl Blue pubescence of male only on apical
margins of T1-T5 and hairs scattered near wing
base and on outer surfaces of hind tibia, T7 of
male with apicomedian trilobed margin (FiS.
51i), 57 of male with apical half much expanded,
apodemes small (FiS. 51h), SB of male broadly
and deeply emarginate (FiS. 51g).
Description. Male: Length l+.6 mm; forewing
length Il.2 mm . Structure. Inner orbits almost
parallel (Fig. 5lj); shortest distance between eyes
.B length of eye; malar space linear; head wider
than long; clypeus protuberant, in profile.6
width of eye; galea in repose reaching posterior
edge of hind coxa; flagellomere I equal to combined length of next 2.3 fl,agellomeres, 1.5 as long
as scape (excluding basal bulb) and 1 .4 times as
long as last flagellomere, flagellomeres 2-10 about
equal in length and 1.3 times as long as wide;
distance between posterior ocelli equal to 1 .3
from
median
distance
distance;
ocellocular
ocellus to posterior ocellus .2 ocellocular distance.
Marginal cell length 1.1 times distance from apex
of marginal cell to wing tip; cu-v of hind wing
subequal in length to second abscissa of M + Cu;
vein M 3. 1 times as long as second abscissa of
M + Cu; j,rgul lobe about one-third length of
vannal lobe. T7 with apicomedian margin trilobed (FiS. 51i); 56 with weak apicomedian emargination (FiS. 51i); 57 with apical half much
SB broadly and deeply
expanded (FiS.51h);
emarginate, disc constricted in basal two-fifths
(FiS. 51g); gonocoxite apically simple from dorsal
it profile basally
or ventral view (FiS.51e),
constricted (Fig. 51f); bridge of penis valve long
and wide (FiS. 5le). Coloration. Black except pale
integumental facial marks (Fig. 51j), coxae, trochanters, femora, and metasomal sterna Burnt
Sienna; wing infuscated. Pubescence.Face with
mixture of black and white hair; gena with white
hair; hair on scutum black with a little white hair
intermixed, ratio of white to black hair gradually
increasing ventrally on pleura to thoracic sterna
which is nearly all white, propodeum laterally
with long white hair; foreleg with white hair on
coxa, with black and white hair on outer surface
of tibia and tarsus, femur and inner surfaces of
tibia with dark hair, inner surface of tarsus Burnt
Sienna; midleg with white hair on coxa, outer
surface of tibia and apex of femur with black and
white hair, rest of midleg with dark hair; coxa and
trochanter of hind l.g with white hair; outer
surface of tibia with white, black and Beryl Blue
hair intermixed, its length about equal to length
of scape but long at apex where it is about equal
to length of mandible, rest of hind leg with black
hair; visible portions of T2-T5 with black hair on
basal two-thirds, apical margin with transverse
band of pale and Beryl Blue hair intermixed; S154 with pale hair, 55-56 with dark hair. Punctation. Clypeus dull, coarsely punctate, punctures
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lâeJ 0026-000g ,s1p11rruel
'ee.re
'1ou4rw
'>p.ra4 'ersÂzptr
.repEuel.rt
pesse.rdap
uerparuocrde
qtrm
a{ta
I :e1euraddroloH
.Sururgs ,(11ee,r,r.
sneddlc lecuelsrp relnrollâf,o Zt'g snllâro rorr
91-91
-elsod ol snfleco uelPelu ruo.ll ef,uelslP :âJu?lslP
juorleredas rpprrr^-ârnpund ç' ot luanguor dFzeu
'esr?oc sernlcund 'Sururqs lerJnâruos x"ror{I
ælnJollâJo 9Z'I ol pnba r11acoroualsod uea,tl
'uotlopund'rreq
-eq
arrrclsrp
lepral
se
3uo1
se
pue
qr8ue1
lnoqz
Tr€lq qtr^ "rrrâls pue gJ-çI :çI
-ZJ a1eurJo reqr âTI 1I-II
ur pnba rnoqp 6-Z saraurolla8eg larauo11a8eglsel
i1ae1q s8e1uo rr"q
se Euol s? sârurl I'Z pu" (q1nq pseq turpnpxa)
;o lser lrreg enlg 1d.regpue Tr"lq qll^À e€rqnprru
adecs se 8uo1 se y6'g 'sarauogaEzg ç'g lxau
pue a.ro; lrreq elrq,r,rqtp{ exor purq treq 1ce1q
pu? etrq^{ qtlm â€xocprur pu? âroJ lrreq 1ce1q
Jo rÉual peurquoc ot pnbe 1 areurollaSeg iprrç
pu€ ettllrÀ qll/\À 3âr€ papodord lrruq anlg 1rfuag
Ps"q uo qlool rorrâlu" lnoqlrr\4,urnluau lglool
pcrdeqns qtr^Àalqrpu?u ia,(a;o qtprm g7'g agord
pue 'atrqa,r.'1ce1q 3o ârnlxrru çr,rn erneld pue
rn ecu?reqnlord peddlc 'teg ece; ituol ueqr
runtnrs lrreq alrqan qll,r'l,JIBII ra,no1 lpg "reddn
raprn p?aq lreeurlaeedsrepru ledego{fua1 19'6
uo rr"q anlg 1rfueg pu" etrqi qlrm eue8 lanlg
seda uaaalloq âruelslp lsâuoqs la,roqe Su8.rerrrp ldr"g p* 'elrqal 'rpe1q Jo ârnlnru " rrcrl [€rJ"d
'etu?s?qnd'{r€lq lsorup ol suuars lurng s8al 'sna
dpqEgs slrqro reuul 'anpruts 'urur g'11 qfual
-dr(pJoJleq uelpâur uo Tr?ru ptueurn5etur epd
8ura,r,e.ro;lurru ç1 qr8ue1 :tloruat 'uolldyraq
'(sercads
'ds
nsouïug
aes)
gerus ldacxe )pel16 'uoqDroloC'ævel
rep8uerrl
terqt
1o
'uoxopon{optàtwyoterue.readdeur.rzlurrsd.rerr
-srp .relnoolleJo€I'snllâf,o.ror.relsod o} snllef,o
lprrç pseq uo qlool JorJetue lnoqlr/r urnluaur
u"rpelu uro{ oJrr"lsrp :âJu"lsrp nlnJollâJo
lada;o qrpral çt'0 eJrr?râqnro.rdpeddlc algord ur
7'1 o1 pnba rllâJo rolrelsod uaa,rnlaq ef,u"lsrp
le.reruolpSeg1se1se tuol s" 6'I Pu? (qpq pseq
:(pZC'StdelulduerperueSuruuo;seSpuBurSra,r
-uoc dgecrde qlr^À urruqe.I 's{r€ur ["rr"J l"tuâru
Surpnlcxa) adecs se Euol se 9'1 'sa.raruoqet V ïe
-n8elur epd lnoqlrrn 'T"rlq 11nateg 'srsou7mq
txâu Jo rp5uel peurquroe ol pnbe 1 a.rauro1e8zlg
sercads
*awnuto{uocuou
optscuy
"'#,ffi'f_"'&'S"j"':;1ff"j*iïii.J",j3:Èilili
r' €I'F8o'ltTjâil'".îË,lt"
.scqeruarsds
H#;"ffî"*u,
aurroqdoqlue ol suorlnqrrluoJ luâllâf,xa dueur srq
roJ'TlulrJâl.I'VsnEI
I Tqpu_al{poo5_ete1 du;o
uorlruSocar urnpârueu sr sercedssyqa'iâ0l,9!trCtg _
_
^âFru aql;o sSurqruu paddp apd
pâf,npar fpn.ri pu€ ec"J elpurâJ{celq lsou{e eql
dq p,te azrs ra8rel aql dq 'uuroselâu âql se llâir,r
s? x"_rorll âql uo rrurl enlq rrll€lâru ;o eruesa.rd
eç dq,'ra,remoq'r11tîtwouo2;o segcadsduz uror;
pelzredes Âlrseesrt1lal,{ay'y'qpâawouo2 dueru;o
rlpïawosso\g unoun
rptlr e{lFltrn rou trcq anlq
çy't'
dpoeqrsrsarcads
sunp's
'\qL '"*K,,
-^{s-I le gJ"â sel?urâJo ^l Pu" saleru oan1ldecxa
uâprâ.11epalrsodap are seddlered 11y '&nqe1pua4
'l\l 'H dq pelcaqoc ere^{
_suorulJeds ge. pefec
-rpur asrmrarllo se ldaexg '(uep8e4 f 'H) tlOt
eunfç '(ur tgil) real 00gt 'lllH s.res?rd 'Steqed
(apurag 1) pr:r- lo1to1otot7uo (uap8e4 f 'g) OlOt
iGenuef Ig 'elelf qeuza 'spuzlq8rH uoJaru"C
'(ur
'Suer1ed (epruag
t) :ggOt 4t"f çZ
91çy) æa3
''rtr41lzpuâd 'spue1q8111
uoreru?C '8*qtd
0l,Iç
(apruag t) :OgOt I'zIN Ll '(u 1161-1991)ree;
,eteloareorcriu sunicund uàe,r1aq pundr-5
9g-7g
-,rioirâcunà
aropils ot âsreof, pâreilers qlr^
.durqs gg-1g :arftoàr?orrrru ,(lasreoc erou sirnl
-cuirâ u5e^iaq pirno.r$ tâacxi z$ral Surpaecerd
e1{ uonepund ,Eururqs d11eem 4J-gJ ,erzl
-oàieoprur (1eug sarriljrind ueemraq puno-$ ,e1e1
-n1pu.rà dles.reôt ,111,p(p*q leq-5pd Sur.reeq
ur5ieur pci.fe Eurpiilcxe) suôrfod elqrsr,r qrln
1a-1a :iegu4s 1nq ruhlnrs u€rll râug uôfenirnd
t.ttq* nrr,âia iqroo-* sarnlcund uââ^ leq punor5
_n# ":*ilî,il':ï[i
,il:l ffily"ï.l#3il:
sernpund ueâlÀlâq punort 'uzdu sqlpr,u-arnl
-cund 7 ol tuenguor sa.rnlcund 'uoltelcund aug'
ol âsrzo3 qlrr* ,r(urqsxeue,r lsarnlcund luenguoc
Â1.reauas:éoc daap qlr,l,r,'durqs ea:e paddperdns
:ilelosreorcrur ,(1eug sa.rnlcund uaaarlaq puno.rt
'durqs searer"lnf,o
'sa.ràlcund,r,rolpqsâsruor
çr,u
-ured lalzloe.reororru sârnlJund ueea,rleqpunorS
,luenguoc dgeau ,i*o1pqs serntcund 'durqs 1eq,u
-eruoJ ruruqe1 ieluloareonrru sarnpund uââ^{}âc.
puno.rE ,rredé sqrprar-erntund0'Z-ç'0 pue deep
Nrrg-r-rng scNslcs svsNVN do Àrrsus^rNn
sHJ
zçç
SvsrnuATlcs
oF THE ANTHoPHoRTNB Bees
AND PnvrocENY
white hairs anteriorly. T1 with white hair on basal
two- thirds, T2-T3 all black, S2-S5 with apicomedian band of short orange hair, in addition S3-S5
laterally with white. Punctation. Like Amegilla paradoxa except face dull, although shiny near posterior ocelli and apicomedian
area of
triangular
clypeus.
Anthophora
(Megamegjha)
new sPecles
553
paradoxa
Diagnosis. Clypeus of female completely flat
medially, with short dark bristles. Upp.r half of
thorax with red hair, lower half white; T1-T3
with all black; T4 with white hair on apical half;
T5 covered with white except black apicomedially; legs essentially all black except some white
hair on outer surfaces of tibiae.
Description. Female: Length 17 .6 mm; forewing
length 13.8 mm. Structure. Inner orbits more or
less parallel; shortest distance between eyes 0.84
length of eye; malar space linear; head wider than
long; clypeal protuberance in profrle 0.6 width of
eye; galea in repose reaches anterior margin of
midcoxa; mandible with subapical tooth; flagellomere I equal to combined length of next 3 . B
flagellomeres, l.l7 as long as scape (excluding
basal bulb) and 2. 1 times as long as last flagellomere; flagellomeres 2-9 about equal in length and
about 0.87 times as long as wide; distance between posterior ocelli equal to I .22 ocellocular
distance, distance from median ocellus to posterior ocellus 0.52 ocellocular distance. Marginal
cell length equal to 0.78 distance from apex of cell
Tlpt
materiaL Holotype
female:
Malawi,
Rumpi, Northern Dist., 12 April 1967, C. D.
Michener [Lawrence].
Comments. This species is so strange that without the male I am uncertain as to what subgenus
it may be in. It possibly is an Aframegilla. It is
easily recognized by the strangely punctured face
and lack of a tooth on the mentum, which suggest
Anthophora, yet it lacks arolia and has a single
basistipital process, features that place it in Amegilla. The color and pattern of the pubescence
make it look very similar to Amegilla (Megamegilla)
albocaudala (Dours) and A. (M ) paradoxa n. sp.,
yet the structure of the labrum and all black face
easily separate it from those two.
Erymologlt This species is so named because it is
the only known species of Amegilla without an
anterior tooth on the basal third of the mentum.
v
B?
D
r-l
c/\
-/r\\
z
F
)
./
FIG. 52. New species of Amzgilla. A-C, A. (Asaropoda)houstoni,female holotype; head, frontal view;
apex of 56, side and ventral views. D, A. nonconforma,
female holotype, head, frontal view. E-G, A.
(Asaropoda)epaphrodita,female holotype; head, frontal view; apex of 56, side and ventral views. H, ,4.
(Megamegilk) paradoxa, female holotype, head, frontal view.
ery1-ladrec'esuep pu€ urelred r;3tËrtiil:l
-eroJ lurur Z' gT-+' ç1 qrSual :apaat'uotg(ttcsaq
'(E 'lZç 'sEt.D qlool pcrdeqns e]€ur8rerue
stl Jo asneceq (turrur"qc 'd1ano1''rC 'so4tpottldoda) dpunlq ql1,v\ 9g lrreq
Tretq Jo rlrled uerpaurocrde
'(Eopu(1g
perueu os sr sarcads s5{I
qll^Â
aln{/v\
!(rreq
ldacxa
9J
I[e gI-tJ
3o qcled
'(erprlsnv
urelseM
urog)
erlga lerelelocrde ll€rrrs qll^À tI
ldecxa) Trelq
ru|Fnoq 'V Pewlar l,lasolr eqt ruo{ (drolrrral ure
lrreq ("lpq reddn) atuero pue ("lpq
IIe SI-II
'V eteredesgrrnte8rel
-t{rro111
eq} ruo4) oTlpoa@eda
rarurol) {c€lq q}lû\ xeror{} lrreq (lpq raddn) a8uero
Ieruossleur eql uo urelled lueJseqnd aqr s€ IIa/v\ tqEI ol (J1eq rer*o1) âllqÀ\ r{}yv\ peoH 'stsou7otg
s€ ("2ç 'Stg) s8urlreur FIrBJ eql 'erlerlsnv ul
sarcads rrÀerr Drlporqdoda (opo{o.tosv) ùllfauv
rerllo t(ue ruo{ sarcadssIIl ateredes dpsea IIIÀ\ gS
uo qtool pcrdeqns aql qlyv\ Euop stsouîptq eql ur
'eete peddlcerdns pu" peddlc pesrporrr dle8uerls
uanrt uralled rolor e^rlJurlsp aqJ 'sruauwoC
'[ecuerr*e1
'(Eopu(19
eqr roJ perueu sr serceds sHI
'[errequeC gaqdrueC'C 'L'7,til drenrqe.{ I ]
'?JI{JV rtr Dtotnl of pe}€ler rurreJ
t]
?t
(apura; I) 'Zgil drenrqe.{ çI (saleure; g) >1aar3 -ng Jo snue8 peqrJf,sopun TJe ur osle ueas uaeq
s(Tcorfl 'aprsurng 'drolrrraa urerluoN ruor;
s?r{ uor}scgrporu e qrns 'plro^A plo eq} uI seeq
sepureJ rnod :saddtercd '[erraqueC] uoldn .S
eurroqdoqlue
Euoure punoJ uor]erglpour
qcns
'Il{ 'r 'N 'BL6I
'9,0çoZtI 'S,ZçoZI
rlrr?trAtr
6-9
'lllr{eC'}W'g
urT çI'drolrrra; urerluoN
'erprlsny
:add1o1oq eleruad 'îotrar)w ,d.*n*n
sqlpr/v\-eJnlJundg ot snontrluos dlreau sarnlcund
'durqs etre]
sarnlcund
'ilnp xeror{I Ieruosererq'snon8rluoJ
'sqlpr/v\-ernlcund
0'g-ç'0 Âq prrnt
-edas seJnlcund isnaddlc Jo {reur m.ogadusrperu
uo dlprcadsa 'durqs ered 'uour4eun1'?S pue gS
uo rreq elFl/r\ Jo segcled prawl v ro3 ldacxe Trelq
eurels l"ruosslerrr lrreq Tc€lq Jo qclsd esuap
II"
'uerpeurocrde
qll^Àosle çI '( [rpl^ ruru tz'g] eæq
?J Jo urtreur usrpeuocrde) srrerl Tr"p parelt"rs
pue rleq ellq,v\ pessardde qlp\ gI-?I
l(Â11era
-1e1ocrdeqcled elFlr\ passarddeqtrrnrtI) Tcelq tJ
-II 'rl3rl TrBp rlllÀ\ slqll Jo assJrnsrorrelu€ pu?
eJeJJnsJeuw lnq dporrelue
Jeuur pue JnrueJ
Jo
rlsr{ tuol tur,req elql} pue rnueJ 'ralueqcort
'(paxrru
'Bxor
rll1,v\
ia8uero
Ealaro;
JIEr{ reddn
-relur srrsrl er1lr'r ^ÀeJe rlll^À) TcBIq x€rorll Jo
'o}lq/v\
'a8uero prrtn-auo ot
JI"q re/v\oT
JI"t{ raddn
r{}l^ÀPser{Jo sPrFl}-o/vuo} JIsr{ râ/v\oT 'xBror{} Pu?
'u/v\orq qslPper
peeq uo asueQ 'aeuatsaqnd
?S-IS
pue EJorueJ 'srelueqcoJl 'esxoc 'eapt 'urnrqel
'sneddlr
Jo sTreur 1essq'ttd
1eretel
Jo urtrenr pcrde
rvrogel.sTr"ru FIoBJ 'ur*orq
uerperu :(tZE
qsrpper euuelue ]decxa æEIfl 'ulrtp.toîo7 '(8
'Eurlceford
'IZ,ç 'stt'D
dlrorralsod
qlool
lecrdeqns
'e8re1 qll^À gS 'âqol
'aleur8rerrre dpunlq
leuue^
Jo {}tua1 L+'0 aqol ptnf i.C+W Jo Bssrrsqe
puorâs sB 8uo1 sB serur] g6't IN ure^ i.C+tr\t
Jo essrrsqepuocesJo tp8ual o1 pnba tnoqz tuu*
pugJo ^-nr idrt tuuv\ ol IIarJo xade ruor; erue}srp
'ef,uelsrprel
06'0 o1 pnba qttuel IIao 1eurtre141
-nrolleco L+'0 ot pnba snllero rorrelsod or snlleoo
uerpetu ruo{ eoue}srp lacuelsrp relnoolleoo I'I
rllero rorrelsod uearrrlaqeruelslp !1 eratuoilateg
;o qfuel I L'0 o1 pnbe eretuogeteg rsel iqrtual
ur tursearcur dgenperE 6-g seretuogateg lg
aiauroilateg 5b qiÊ"ri o'0 oi pnba 7 ariurolfeseg
:(qpq Fseq turpnlcxa) adecs se tuol se 16'0
pue sereurogeteg I 'g ]xeu Jo tlltual pewqtuoo
ol pnbe I ereuoga8eg iqloot pcrdeqns qllÀ\
elqrpusur içpyrr ada 6ç'0 egord uI aJueraqnlord
padllc ituol uerll reprm p€erl iada Jo {}Eue1
IBtuorJSB'0 sadauee/vueqeouelsrplseuotls lanoqe
tur8ranlp stlqro reuul 'ruru Zf rnoqe qlEual Eurm
lsrg eql sr pue ( t rlqnl ul g elq€rre^ aas) sarcads
(otoq ot1\uoco4s(74J) o"tot1d oqtuv avros Jo tuec srur ruer
d
sr uorlergrporu lepr.g yo adft slql 'sapsrrq Trsp
'uoqs
qll^À
pexrtu
srreq
dllecrde
pa{ooq
lnots
'allr{a 'rr5 e^€q 'uorlrppe ur 'qclq^ ssa*re
1ee
-ddlcerdns pue peddlo peuoueg dlaEuerls aql l'q
sarcads oppauy rerllo ,lue pue lr ruor; paleredas
aq dpsee usc 1nq (srnoq) oppnntoq1o (q1tîauofla747)
'sruarazul7
oppawy
solquesar
sercads slrlJ
'[acuarurel] rauaqclry
'Cg 'C 'Lg6I
''lslc[ ureguoN 'rdun11
IIrdV U
'1rvrep141 :eprrral
add1o1o11 'lnuapw
ad[J
'snoJqu18
'Sururqs
arour
eurels
dllearnr
etra]
]nq
'rlur{ dq pernrsqo dlalalduror
}sorulg
I"uroseretr\l
'secedsrelq ur 8rl
ereJrns lnq Eururqs xeroql
-uF{s dpleervr lnq asBJ uo snon8r}uoc tsoullv 'uolt
-qeund 'elHS 'çS uo dlpralel srr€q /v\oJs pue
tS
-gS uo qcled 1erâtel âsuep e 'ZS uo dllerelelocrde
pue dgerparuocrde srr€rl ù\eJ e ' I S uo a8urr;
pcrdu uorls 'rrg e ]dacxe rrer{ TrBp r{}l^À Bura}s
Ietuosslaru ldlerpaurocrde u/v\oJq par tdeJxa Trep
IIB 9J ll,lprpeurocrde rl€r{ Jo rlr}Bd >1ce1qesuep s
pue dlprparuowq rlerl Tr"p 'tcalarruas 'parellers
uonlppe ul 'dgelpetu pesserdde 1nq dllerelel rleq
elFl/v\ lJere r{}y/t çI lsrreq TrcP lJârerures 'pera1
-]ers e\al e pu€ rlerl âllr1m pesserdde qll/n\
JIsq
'ereJrns
pcrde uo ?I l1ce1q ile SI-II
leseq relno
uo elrqû\ Jo qcled ll€urs q]lm erqr]prru lsnsrelero;
Pu? srqrleroJ Jo sereJrns ralno uo ellq/v\ qlyv\
pexru ]dacxe Toelq dpsour qrp\ stel lelrqrvr JFq
remol 'a8uero JIsr{ raddn uo xeror{J 'sJIBr{ u/v\orq
'palooq
dlpcrde 'arg qrl^À dlerpeuroffiq
lqtï
urruqsl lsepsrrq Trcp 'lq8rerls '1no1s 'poqs qll^À
snaddlc puu eete paddlcerdns uorllppe q lsecrde
'sleq otlq,v\ eug q}lû\
lB perlue^ poToor{ dpsoul
urnrqel pus sneddlc 'eete paddlcerdns lurtrenr
relnJosred tuop dy*orreu pue olqrpu"ur 'euat
'lndrcco
uo
elrqrvr arnd lnq rrer{ TcBIq pue e}5{1v\
qfyv\ P?âH 'aeuatsaqnd' u/v\org l{slppâr
Jo arnl{ul
€urals lBruosB]atr\tr 'TrEIq r{srpper s8al 'u/v\orq
qsrppar eept pue elqrpueur Jo dTr :(qZç '4g)
al${/v\ Lruearc sTreru 1epe1 F}uaun8alur
}decxa
'uoztnroloJ
l,lprpaur
leg dlelelduroc eete
TOBIS
'eqol
leaddlcerdns pue sneddl3
I"uuB^ ;o qfual
gt'g aqol le8nl i*C + tr\lJo Bssrrsqe puores se tuol
su serurt L't trAtrure^ inC + nt Jo essrrsqe puores
;o qf8uel I'1 Eurrrr pqq Jo ^-nf, urel ldrt 8ur/v\ ol
Nrrg-r-rnfl scNgrcs svsNvx .{o Àrrsus^rNf} sHJ
,çç
SvstB*rATIcs AND PuvrocENY
Arnegilla (Asaropoda) houstona'new species
Diagnosis: Similar to A. epaphroditabut facial
marks differ (FiS. 52a) and T2 and T3 with well
developed apical bands of appressedwhite hair;
vein M 4.67 times as long as second abscissaof
M + Cu; 56 with acute subapical tooth (Figs. 52b,
c).
Description. Female: Length 15 mm; forewing
length 12 mm. Inner orbits diverging above;
shortest distance between eyes 0.9 1 frontal length
of eye; head wider than long; clypeal protuberance in profile 0.63 eye width; mandible
with subapical tooth; flagellomere I equal to
combined length of next three flagellomeres and
0.BB as long as scape (excluding basal bulb);
second flagellomere equal to 0.84 length of flagellomere 3; flagellomeres 4-9 about equal in length
and each segment slightly longer than wide ; last
flagellomere equal to 0 .72 length of flagellomere
1 ; distance between posterior ocelli 1 . 1B ocellocular distance; distance from median ocellus to
posterior ocellus equal to 0. 52 ocellocular distance. Marginal cell length equal to 0.85 distance
from apex of cell to wingtip; cu-v of hind wing
slightly longer than second abscissa of M * Cu;
vein M 4.67 times as long as second abscissa of
M + Cu; jugal lobe about half as long as vannal
oF THE ANTHOPHORINN
BNBS
555
lobe. 56 with distinct acute, posteriorly projectirg, subapical tooth (Figs. 52b, c). Coloration.
Blâck except facial marks yellow (FiS. 52a); malar
space, narrow apical margin of clypeus and median area of mandible reddish brown. Pubescence.
Like ,4. epaphrodita except T1 with narrow apical
band of appressed white hair; T2 with distinct
apical white band about twice as wide as that of
T1; T3 with distinct apical white band about
twice as wide as that of T2; white, appressed,
band of hair on T4 departing from margin medially. Punctation. Like /. epaphrodita but apicomedian margin of T4 impunctate and about twice as
wide (0.39 mm).
female: Australia,
fypt
materiaL Holotype
Western Australia, top of Napier Range, Windjana Gorge, Kimberley Div., 10 April 1980, G:
Anderson, D. Symor/on Trichodzsma sp./WAM
87 -1303/#350 Amegilla (Amegilla?) sP. det l9B7 ,
female, by T. F. Houston [Perth].
Comments. This species is most closely related
and similar to A. epaphrodita from which it differs
by the characters given in the Diagnosis.
Etymologlt This species is dedicated to Terry F.
Houston for his meticulously illustrated and fine
work on Australian bees.
APPENDIX 2. LECTOTYPIC AND NEOTYPIC DESIGNATIONS
Each section (Lectotypes and Neotypes) of this Appendix is arranged alphabetically by
author. Each type designation is amanged in the following sequence:original combination,
type locality (TL), sex, label data (L) with each label separatedby a slash, type depository in
brackets according to the list given in Methods, and notes (if any). Each specimen bears a
red lectotype or neotype label.
LECTOTYPES
Lectotypes are designated for various anthophorine and habropodine species when syntypic series exist without published designations
of holotypes. Even authors who labelled a type, as
distinguished from paratypes or cotypes, often
failed to indicate which specimen was the type
(: holotype) in their publications. Because of
superficial similarity of species in different subgenera, type series are sometimes composite.
Designation of lectotypes is the simplest way to
clarify such problems. The genus and subgenus
to which each name belongs can be determined
through the index function of Appendix 3.
T. D. A. Cockerell
Podaliriuscleomtr(TL: New Mexico, Sante Fe
Co., Santa Fe. Male. L: Ckll. +452., Sta. Fe,
NM., Veri Boyle/Co-TypeNo. 3+04 U.S.N.M./
P. cleomisCkll. COTYPE) [Washington,D.C.].
Cockerell's "cotypes" are normally paratypes,
but no "tyP." (: holotype) can be found for this
and the next species.
Podalirius aallorum (TL: New Mexico, Dona
Ana Co., Las Cruces. Female. L: Ckll +795, or
Ipomoea/LasCruces NM/P. vallorum COTYPE)
[Washington,D.C.l.
J. Dours
Anthophorabadia (TL: México. Male. L: Mex
5B/+ males, 2 females/Anthophora badia n. sp./
Museum Paris, Coll. O. Sichel 1867) [Paris]. 19
"Museum
male paralectotypes all with the label
Paris, Coll. O. Sichel1867". In addition t had no
other label, 2 with "Mex 58" , 12 with "Oaxaca'', 2 with "Oax. Salle" and 2 with "Orizaba
66" [15 Paris, 4 Lawrence].
Anthophorabelieri (TL: Sicily. Female. L: Sicile/l53) [Paris].
Anthophoralaticincta(TL: France, Corsica. Female. L: Cors/Anthoph laticincta Sich n. s. 3
females/- nigro-cincta Lp. ,r. female aJurin ghli/
Drews,J.) [P"ris].
H. Friese
Anthop horafulu itarsi s murutica (TL : Afghan i stan,
Qpndahâr, Ma'rùt. Male. L: Caucasus, Murut)
ueeq seq puofi(xo 'V eureu ar{J '[srre4] ('drg
'^/6tl1 :T 'aptr\tr 'eru€rrepenC
/'srg euotdxo
ep €rrers 'uredg :TJ) puofi(xo o^tot1{otltuV
'uorldrrf,sep pur8rro aq]
r{}lÀ\ saarSe eddloeu eql 'rl€t{ par t{}!/v\ Pore^oc
dpsour stel eqt Jo IIe pue II uo rleq elFl/v\
snoncrdsuoc B rllr/v\ tnq >[r?lq II3
Jo pusq pc1d3
lsorule sr 'spue1s1âpre1 ade3 âql ur pazrutocar
dpsee 'sercadssHI '[ocsrcuerd ues] (pttU xg
'uorlcelloClsgreH 'd/eeg,'T 'B6BI-IIX 'ur
009-0
'nelocr1q 'S 'apren ode3 'sI :T 'elurrrad '(a1uec
-t^ ogs se/v\d1r1eco1ed& pur8rro) nelocr51 ogs
'spue1s1epre ede3 :TI)
ryu{opoî o,roqdotltuV
olIseu
sed1rp,,lio.lff"Lx
il*tuifl3#
'uortdrrJsap pur8rro aqt
qly!\ saar8e pue Iecrddlodot sr addtoau aql 'rlel{
ellqn\ ol snoeoerrlro Jo spueq l€cTde ]rurlsp lnq
asreds a^Br{ etret leurose}eru eq} puu snoeJeJrloo
sr xerorll eql lneq a8uero qll^À pare^or dllsour
ere stal aqJ 'r*ogad dpsour sr urnrqel eq] pue
snaddlc eqt uo Tretu padeqs-; paua^ur mo11ad
e seq epural ar{I '[srre4] (?n.tg slsreu^lnJ 'V
:T 'elerrrad 'erra8ly :TJ) slsrqlaln{ otoqdoqtuv
addroeu
searEe
*,,;îï'iii':iË.'ïf
t:îli;,TH
dlpcrdu uB su IIe/r\ sB 'qsnrq l€sr€]r]s1pplur e
Jo acuâsard eql pu€ sâqsTrrq lesr€}Is€qplur pu€ I€s
-Jslprtu Jo ef,uasqe oql Lq sereads reqlo tuo{ pelsr
-edas âq uer eletu âqI '[srre4] (sruge 'V/6IOI
pur8rro âql qll/u\ saer8e addloau eql 'uosrred
allneg ep 'f '11oC 'srre4 tunasntr^Ilç-11 'suo4
'erratly
'I :T 'eletr
-rrror roJ ouo{uozuou 'V pue oxopon( opfawy
sluffi ntoqdotltuy
:TI)
tr
'punoJ ueeq seq
aes 'rlzl{ at5l,v\ dpsour q}l/nl çJ pue +L pue
'ecuersaqnd a8uero
q}lû\ xeroq}
sarceds
{oBIq IIB SI-IJ'slreur
?ilrug EurrvrolloJ arll Jo Isueletu adft otr1
'V
VpeJ [s]ueurn8a]ure]Ir{^/v\-r{sr
JoJIstI raddn
?IINJfl
-mo11ed
s1r'(urur 6t rnoqe) azrs a8rel s1rdq pezru
-8ocar aq dpsee uer r4npnnzoqp'V '[srre4] (srnoq
'saddlolrel
roJ
se
peluaserd
ere
Bl€Cl
er"pnef,oq1eeroqdoqtuy' 2,06I_haWIuorlsur*Irr
'saddloeu Eurrnogo; eqt paleutrsep â^eq
-ap s,legceAlllOOl uossdnofl 'f 'oEo1,N enpoSo
I (xel
''cuurg o8uo3 :T 'elstrred '(eaurng se/v\l,1r1eco1 e^oq" aqr aas) serrotal?c rrroua8qns w rueql aculd
addr purtrro) U,ç2,o6 'N,lzog 'anpo8g 'ranr11 pue serueu EurrrrogoJ orlt ezrlrqs]s o] râpro uI
o,tot1(otltuV
o8o4,qq'uoqe11 :-II) oîopnugoqlv
sgdÀrogN
od& .srnoc leuolllpp" ,tue dn urnl Ji'iJ1n1i
'1aseg pue 'eneueg 'uol,1 le srunesnur agl lu,
s3 IIa/v\se 'urnesntr\trsrrsd erll le qcJsese^Ilsn€r{xe
uV 'I MM ur acuurd 'suarruy te quoq v dq
padoæsâpsem (aroqe pelsï leuelsur eddlolcel eql
ro; ldacxa) prreleru ed& (srnoq Jo IIe lsorulv
srnoq 'f
'tuo1 ruru gl-U sr pu" ecuoJseqnd
Lart seq aDllannbsal'FZ'alqlpueru eq] Jo lurodprur
oql 01 lelslp dpqtrls sI qloo] lecrdeqns oql pue
rrrruqel er{} uo (rrtoged I1e q }f) nrrr lecrdu >lrelq
ou sBtI alsur o.tafituupo'V 'alqrpusrrr âI{} Jo tIUg
pcrde eql uo sr qlool pcrdeqns eql pue urruqsl
'V'
eqt uo urlr pcrde 1celq B set{ e1etuaoya.ranbsa1
'e>l"lrâquurll o.ra7ruupa'pz q aopatanbsal
oroyfuqruy
otoqd
qtp1r'\pesnJuoc eq plnor lBI{r (o^rot1dor17uoîQ)
-oqruv PIroM /u\eNJo sarradsÂpo eql 'uol]sallor
uef,rreurV çro51 dra,ra ]sorule ur ruar{} roJ paTool
e^eq I q8noqtle perâ^orslp ueeq e^eg sarcads
saddloc oN '['C'A uo]EqqffiMl (ra>1re4
srrll
'g
'oruotuy uBS (*l
'qg Jo '
t'6I)
LL-I I-AI
'lur d ''oC
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ZI
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''oC orroc
oruoluv ues (*rt g'6I) 'g rtu
Z,I
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sntrzîopod
'>l-J'p 'q 'egl
'sEI.{w pe}Br}snp ere sorn}BalEurqsrn8ur}slpotIJ
'uossarC Duoluouo"totldotltuv
p urduouds rotunf e
sr:lnprut d l' D'CI 'uol8url{w6l (t1e* '1ptCInprer
eroqdoqtvv/Dtvlnu,asawoalJIo 'srg w/('11>1C)'I I
'se8el
wT ''oC
dpf 6'tr^{'51 'seta1 seT :T 'alBtrAI
'ocrxa;41/n\eN:TI)
snltllupod
lnprpe
Ian8ltr\trues
'v 'c
I
llara{coc
NIIsT-Infl
'tB6I ul ldrrcsnueru ur eddlot
-re[ sB pateu8rsapraTeg'g'61
]Br{} uaunceds
erues aqt sr s${I '[uopuol] (gOg-OO'ou uol]cel
-lor 'arlq6 uarred 'srtr\tr'uorlcalloC qlrus/re^l1
puourrlclg 'erlerlsny :T 'oleured 'g , ç g" t ç I
'se1e11 qrnos
'S,
're,u11 puotrtqrlll
tE oBZ
sya"tofiqwoqopo(o^tog
:TI)
/r\eN 'erprlsny
'
opclsopnas{
6Z6I'âTrlpap
'v
'[,rop
'v se poIueuar s?/v\D\nels
Jo eletu s(tltrus
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'V :T 'el"tuod l'lTtls :TJ) D1ntlso"rotldotltuV
qIFIS U
'[srre4] (apur rrpueruo1 :.I
'eletr\l 'uBrO 'erra8ly :TI) tîpumalr ntoqdoqruV
'[srre4] ('Ereg 'lS 'erpaurre]q 'V :T 'eletu
-ed 'surred 'acuerg
o^tot1(otltuV
;TJ) Dzpawnrur 'V
:T 'eletu
[srre4] (apure; Et"rlrrlg
-ad 'uBrO 'erratly :TI) otnlllelq oroqdotltuV
'[srre4] (a1eura;tunroeupfl :T 'eleured
'saauard4 'acuerg :TI) wnroaqoq n"totldoqtuy
'[srre4] (epnre; eue8rqp 'V :T
'aleruod 'uol1 'acuerg :TI) ouafuqlo
o,toq(otltuV
'S ep rarlaladal 'V
U
sualnolTfauv sr serresad& âg1," $lii,if;?;
-qrqsaq; '[u11rogl (oaowodyuo 'doo11-']]ng 'slllH
'ecce1e141 :T 'eptrAtr 'slllH Eurdrel '1era4
'dltl
'ersde1e141 :TI)
szsuageolnuD1nslîlla ntoqdoqruV
'adl.tolcel eqt qllÀ\ cgrcadsuoc
lou ere pue ?11rug slsrnîlaln{o,rotldoqtuv are_'aleureJ
'[urlrag]
'sarras addr âr{rJo
}ser aqJ
I pue saleru g
sCNsIcS SVSNVX .tIO ÀIISus^INn
sHJ
9çç
Svsrprr,tATlcs AND PrrvrocENY
applied to various species of Anthophora. I have
even seen a determination label of A. orcygonaon
an undescribed species of Pachymelus.From material in the Paris Museum and studying the original.description I am certain that A. ztcygo-na.isa
junior synonyin of Anthophora robusta (Khg),
a
well known Mediterranean
species. It is easily
recognized by its large size (17-20 -*),
upper
half of the thorax and T1 and T2 with fulvous
hair and the middistitarsus
without a lateral
fringe of black hair. It can be confused with
Anthophora fuliginosa Morawitz
which has a well
developed middistitarsal lateral fringe of black
hair and is broadly black along all of the lateral
margin of the clypeus (oxygona has only a small
laterobasal spot on its clypeus and is all yellow
below the anterior tentorial pir).
Anthophora rufolanata (TL: South Africa, .Cape
Province, Kaffraria. Male. L: Capland, Willowmore, Arg. 25, 1918, Dt. Brauns) [Pretoria].
This species, endemic to the Cape Province of
South Africa, can be recognized and separated
from its close relative , Anthophora aestita Smith, by
the yellow to red hair (mixed with black) on the
thoracic and metasomal dorsum and the lack of a
middistitarsal lateral fringe of hair. I found 16
specimens at the British Museum (Natural History) determined
by Cockerell as Anthophora
rufolanala. His material agreed well with the original description and I have followed his interpretation of the species.
Anthophora tarsata (TL: México. Male. L: Mejico, Mus:
Drews, Tarsata Sichel, 02: sr.)
[Copenhagen] . A. tarsata, a junior synonym of
Anthophora californica Cresson, is easily recognized
in the western U. S. ând the northern half of
México by the white, apical, integumental bands
on the metasomal terga and the red legs. The
hind tibia of the male is greatly inflated with a
large anterior tooth. The topotypical neotype
agrees with the original description.
Anthophora zonata analis (TL:
Equatorial
Guinea, Rio Benito, 1"36'N, 9"37'8. Male. L:
San Benito, Guiral 1885, 5335-85) [Paris]. This is
a dark form of and conspecific with Amegilla
acraensis (Fabricius). It is almost all black with
scattered white hairs on the head, thoracic
scutum and anterior half of the thorax, and T4
and T5 have predominately
white hair. This
topotypical specimen agrees with the original
description.
W. Erichson & J. Kl,tg
India, Pondicherry
Megilta sesquici,ncta(TL:
PonState, Karikal. Male. L: SOUTH INDIA,
1964 (P. S.
April
dicherry
State, Karikal,
Nathan) ) [Lawrence]. The supposed type material of M. sesquicinctain Berlin, consisting of four
specimens, is not from the type locality nor of the
proper sex. There is no doubt of its identity from
Erichson's description and from where it occurs
in India. The head and thorax are almostly
completely covered with white hair, T1 and T2
OF THE ANTHOPHORINB
BNNS
557
are all black, T3 and T4 have medially interrupted apical bands of white hair and T5 has
white hair laterally. See Commenls under Dizonamegilla for further discussion.
E. Germar
Megilla subterranea (TL:
France. Male.
L:
II-23-1837) [Paris]. There is almost no recognizable remnant of Germar's material in the museums
at
Berlin,
Hamburg,
Eberswalde,
Frankfurt or Munich. In Paris I found material
which is topotypical, agrees well with the original
description and may even be type material of
Germar's. This large species is common in the
Mediterranean region. The head has mostly long
pale hair, scutum has long black hair, scutellum,
metanotum, propodeal area and T1 with long
white hair, T2 and T3 with mostly dark hair, and
the rest of the terga has mostly white hair except
for the median apical patch of dark hair on T5.
J. Giraud
Anthophora obesa (TL:
Italy,
Piemonte,
Fenestrelle. Female. L: 34-72) [Genoa]. This
specimen was collected on the 34th expedition on
the 26-29 July, lB72 (according to the field notes
in the Genoa Museum which were brought to my
attention by R. Poggi). I could not find type or
topotypical
material of Anthophora obesa in the
Paris Museum.
Subsequent searching in other
museums uncovered this specimen in Genoa. ,4.
obesa, a junior synonym of Anthophora balneorum
Lepeletier, is recogni zed by its almost all black
head, pale to yellow thorær and almost all black
metasoma dorsally, except Tl and T5 apicomedially have pale to yellow hair.
J. Klug
Megilla farinosa (TL: Egypt. Male. L:
Aegypten, Schinker/Anthophorafarinosa Klg.
determination
label]/Anthophora
[Alfken's
farinosa Klug [Lieftinck's determination label])
"
l+67 lTyp.
[Berlin]. The female that is labeled
label]/farinosa
Klug*
[orange
[yellow label]/tabsachlich nicht farinosa [Lieftinck's handwriting]/
This specimen is not the type, cf. with Priesner
1957, p. 91 [D. Baker's handwriting]"
is not M.
farinosa. From the Klug material in Berlin I have
selected as neotype the above male which agrees
with Priesner's (1957, see pp. 14, 19 and 90 for
separation from other species and a description)
interpretation
of Amegilla farinosa.
A. Lepeletier de S. F.
Anthophora atrociicta (TL: Senegal. Male. L:
Althoygh.molt
of Lepeletier's
l-pe9up.) [Paris].
cies are represented in the Paris Museum collection, I was unable to find suitable material for
lectotypic designation . A. atrocincta is one of the
most distinctive African species. The upper half
'op1e6-ep"etrAIaopuoln'uV:
Gr-gtv 'ggv-Tgt'Bvr)
['dss oelqnuse] ?16I
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tuol sr snldlsouot eql pue (rrrern FsroP tuor; e^F^
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sruad B
Jo
pue eleut8:Brrre dlese_q sr atprrq
Jo qlpy*) peorq 'eltxocouoE
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eq]
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Jo
ur8reru lerel€l eq] uo pâPunor dlqloours sr ]nq
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pue lroqs sr snldlsouot eql pue (rnn'arnlesrop ruor;
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SYstnuATIcs
AND PuvrocENY
OF THE
Friese,1922
n. syn., dark form; :An. nigriuentras
:An. cyanicollisFriese,
las analisssp.l n. syn. i
1922 las nubicassp.] n. syn., dark form; :An.
Cockerell, 1935 las bipartitassp.] n. syn.l
comperei
=An. lounidgeiCockerell, 19+6 lasf,aaicol/rsssp.] n.
syn., semi-darkform) (438,505-507r 557)
acutilabrisMorawitz, lBB0 An, PO (496)
adamsella(Rayment), l9++ Am (An), A, publication
date, not 19+2 (512)
adnmsorunBrooks n. sp., An, P (438, 462, 544-546)
afulaidae(Cockerell), 1905 Am (At), A (: An. nnesti
Rayment, 79++ las adelaidacssp.] n. syn., publication date, not 19+2, see Text Fig. IX, figt. 3b, 4c
and described as new 1947) (511)
adaenaSmith, seeacraensr(Fabricius)
Cockerell, see albocaudala(Dours)
aduenifurme's
adoenula(Cockerell), 1930 Am (An), E (515)
(Dalla Torre Friese), 1895 An (Megilla), n.
aegyptiaca
name for An. senilisWalker, l97l, not Eversmann,
1846 (460)
aegltptorunPriesner, see atricilla Eversmann
Hedicke, 1931 An, P (4841 486)
aeneiuentras
(Vachal), 1903 Am (An), E (506' 548,558)
aerizusa
(Smith), 1854 Am (An), A (438, 5ll, 512)
aeruginosa
aestiualisPanzer, seeretusa(Linnaeus) (462)
(Friese)
aethiopica(Friese), seeabessinicus
Cameron, see acraensis(Fabricius)
aethiopicu.ç
ajfab;lisCresson, 1B7BAn, N (=An. fulaicollis
Timberlake, 1951 n. syn.) (466)
ûnit Brullé, 1832 An, P (:An. liturata Lepeletier,
1841 n. syn. i :An. asiaticaMorawitz, 1880i :An.
lusitanicaFriese, 1919 las asiaticassP.] n. syn.)
(464-466, 556)
aj,mbriata LaBerge & Michener, see badia (Dours)
af.abellataGribodo , 1926 An, P (466)
afra Priesner, see albigena(Lepeletier)
africana(Friese), 1905 Am (An), E (:An. pseudococcina
Meade-Waldo, l9l+, n. syn.) :An. odontura
Cockerell, 1920, n. syn.) (506' 546, 547)
africanaBenoist, see balneorumLepeletier
aganlaRadoszkowsky, 1869 An, PO (:An. kessleri
Fedtschenko,1875, n. syn.) (464-466)
agamoidesStrand, seedalmaticaPérez
ogràt Smith, see badia (Dours)
alaica Hedicke, see retusa(Linnaeus)
Cockerell, see urbanaCresson
alamosanals
alashanicaGussakovsky, seepekinensisCockerell
albataCresson, 1876 An, N (494)
albellaGussakovsky, 1935 An, P (491)
Dours, seequadrfasciata(de Villers)
albescens
albiceps(Rayment), 1951 Am (Asaropodo),A (515)
albicepsFriese, 1916 An, T, not Friese, 1922 (468)
Brèthes
albicepsFriese, seearequipensis
albicillaPérez,1895 An, P (464,486)
albida Dours, seealbigena(Lepeletier)
Alfken, 1936 An, O (484, 486)
albifascies
albifronellaBrooks An, P, n. name for Heliophila
albifronsWu, 1985, List of Insects of Mt. Tumur,
Tienshan, in Biology of Tienshan, Beijing, p.
148-149(in Chinese, English summary on P. 150)
not An. albifronsEversmann, 1852 or Smith, 1854
(4e 1)
Brèthes
albifronsSmith, seearequipensis
ANTTOPHORINN
BEBS
559
albifrons Eversmann, see bimaculata(Panzer)
albifronsWu, seealb'fronella Brooks
albigenaLepeletier, 1841 Am (At), P (: Megilla
lllige4 1806 n. syn. , noflrennudum;
leucomelaena
=Megilla moribundallliger, 1806 n. syn. , nomen
nudum; :Antofhora (sic) binotataLepeletier, 1841 n.
syn. ) :An. albidaDours, 1869 [as albigenassp.] n.
syn.; :An nanaRadoszkowsky, 1869 fas qwdrifasciatassp.l n. syn. i : Po. nigrithoraxDalla Torre,
lB77 fas albigenassp.]; - Po. pyramidalrsKirby,
1900 n. syn.) :An. afraPriesner, 1957 las albigena
r.tp.l n. syn.) (438, 499, 502-504, 556)
albigenaRayment, see albigenellaMichener
albigenellaMichener, 1965 Am, n. name for Asaropoda
albigenaRayment, 1931, not Lepeletier, 1841 (515)
albipesFriese, seeplumipes(Pallas)
Hedicke, 1936 An, P (462)
albobarbata
(Dours), 1869 Am (An), E (:An. adueniftr'
albocaudata
mis Cockerell, 1946 n. syn.) (506' 553, 5541556)
Saunders, see humilis (Spinola)
albocinerea
albofurugineusFriese, seefuruginea Lepeletier
Radoszkowsky, see dubia Eversmann
albomaculata
Timberlake, seecaliftrnica Cresson
albomarginata
albopictaCockerell, l9l7 An, P (491)
albopictulaCockerell, see oldi Meade-Waldo
llliger, see aiolacea(Lepeletier)
albopunctatu
(Friese), 1922 fas circulatussp.] A- (At)'
alboscopacea
E (504)
(Friese), 1896, An (Po), P (:An. fastuosa
albosignatu
Gribodo, 1924, n' syn' ) :An' tenuicilialaAlfken,
1926) (460, 464)
Gribodo
alj,nii Alfken, seecaelaDs
alfkenellaPriesner, 1957 An, P (491)
aliceaeCockerell, seepraecoxFriese
Saunders,
allwudi Pérez, 1895 An, P (: Po' canariensls
Lieftinck, 1958 las allwudi
1903i :An. fuerteaenturae
ssp.l n. syn.) (460' 462)
alpatouiKuznetzov-Ugamsky, 1927 Un (An), P (450)
alpha (Cockerell), 1904 Am (Saropoda),A (515)
altaicaRadoszkowsky, 1BB2An, P (:Megilla tersa
Erichson, 1849 n. syn.) (460, 462')
altunans(Klrrg), 1845 An (Megillo), P (: Po. wegeneri
Friese, 1899, n. syn.) (474, 476)
alticolaHedicke, see quadrimaculata(Panzer)
ambigw Pérez, seeriuolettiPérez
ambitiosaAlfken, 1935 fasfemoralassp.l At, P (474,
476)
(Lepeletier)
amolitaCockerell, see subcoerulea
Cockerell, seeniaeala(Friese)
amphigltnops
amplicepsFriese, see aestitaSmith
amseliHedicke, 1936 An, P (464, 466)
amtmone(Bingham), 1896 Am (An), O (513)
analisDours, seeacraensr(Fabricius) (557)
andnlusicaPérez, 1902 An, P (464)
andsrsoniCockerell, see mimadaena(Cockerell)
andicolaSchrottky, 1911 An, T (468)
andinensisRuiz, 1940 (Po) nomennudum
andresi(Friese), lgt+ Am (An), P ( :An. glauca
Alfken, 1926) (510)
andrewsi(Cockerell), 1910 fas zonatassp.l A* (An), O
( 5 0 9 ,5 1 1 )
(Dalla Torre), 1896An (Po), E, r.
angolensis
name for
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a
I
I
I
AND PuvrocENy
beqwerti(Cockerell), 1930 Am (An), E (5O4)
berylae
(Rayment), 1947 Am (Ar), A (511)
bettoniCockerell, see conspicuus
Smith
bicalcarataHedicke, see disparilis Friese
biciliataLepeletier, 1841 An, P (:An. caucasica
Radoszkowsky, lB74; : An. mucidaGribodo, lB73;
: An. rnorawitzrAlfken & Blùthg€n, 1937 I: ,tntilabrisof Morawitz, 1895, not Lepeletier, 18411)
(464-466, 556' )
(Erichson & Klug)
bicinctaFabricius, see sesquicincta
(505)
bicolor(Saussure), 1890 Pachltmelus
(Hu) n. comb., E
bicolorSichel, see balnelrumLepeletier
bicornisZetterstedt, 1B3B(A") : Osmia(Megachilidae)
bidentataProvancher, see terminalisCresson
bifasciataFedtschenko, 1875 An, P (465)
bihamataPanzer, 1809 Un (Apis), P (450)
bimaculata(Panzer), l79B An (Apis), P (: Apit
rotundataPanzer, l79B; : 4n. saropodaLamarck,
lBlT; :An. squalidaLepeletier, 1841 n. syn.;
--An. albifronsEversmann, 1852i :An. cognata
Popov, 1952
Smith, 1854; :' Heliofhila mesasiatica
[as bimaculatassp.l nomennudum)(446, 449, 453,
489-491, 493, 499)
(Smith)
bimaculiftraWalker, see niaeocincla
(Smith)
bimaculosruDalla Torre, seeniaeocincta
binghami(Schulz), 1906 Am (Po), P (r. name for An.
Bingham, 1897, not Krg, 1845) (511)
crocea
binotataLepeletier, see albigena(Lepeletier)
bipartita Smith, seeacraensei
(Fabricius)
birkmanna(Cockerell), 1905 Ha (Emphoropsis)n.
comb., N
bispinosaCockerell, 1949 An, T (494)
bistrigosaIlliger, 1806 (Megilk) nomennudum
bisulcaPérez, 1895 An, P (496)
blandaPérez, 1895 An, P (476)
bogdanowiFedtschenko, 1875 An, P (466)
bogutensis(Marikovskaya), 1976 An (Paramegilla), P
(476)
bohartiBrooks n. sp. An, P (438, 466, 468, 532-534,
536)
Brèthes
boliaiensisFriese, seearequipensis
(Smith), 1854 Am (Saropodo),A (45O, 496,
bombiftrmes
513-515, 556)
bombiomorphaWu, seep seudobomboidzs(Meade-Waldo)
bomboiùsKirby, 1837 An, N (--An. canadensis
Cresson, 1869; :An. sodalisCresson, 1879 n.
syn.; --An. insularisSmith, 1879 n. syn. i :An.
solitariaRitsema, 1BB0[n. name for An. insularis
Smith, 1879, not Smith, 18571n. syn.) :An.
ssp.] n.
neomexicana
Cockerell, 1900 las bomboides
syn.; --An. stanfordianaCockerell, 1904 n. syn.;
--An.
Swenk, 1909 n. syn. i :An.
fumipennes
scutellarisSwenk, 1909 n. syn. i --An. willingi
ssp.l; :An. raui
Cockerell, 1911 las bomboides
Rohwer, 1923 n. syn. i :An. aterrimaCockerell,
192+ n. syn.) (448, 478)
bombyformes
Smith, see badia (Dours)
bombylansMocsâry seeganula (Rossi)
boopsAlfken, see aidua (Kl"S)
borealisMorawitz, 1864 An, P (466, 468, 478, 489,
534)
Priesner, see zanoni Gribodo
borgensis
oF THE ANrHopHoRINe
Bens
s61
(Cockerell), 1910 fas zonatassp.l A* (4.),
borneensis
o (511)
bothai(Friese), 1911 Am (An), E (506)
bouwmani(Lieftinck), l9++ Am (An), O (513)
braunsiana
Friese, 1905 An, E (492)
brmùftrmrs Dalla Torre, see badia (Dours)
Illiger, 1806 (MegiUQ nomennudum
breaiuscula
(Bingham), 1890 Am (An), O (513)
brookiae
bucconis
(Friese), 1910 Ha (An), O
bucharica
(Gussakovsky),1935 Am (An), P (515)
buruensis
(Cockerell), 1911 fas zonatassp.] Am (An), O
(511)
byssina(Kltg), 1845 Am (Saropoda),P (: Po. schmiedtknechtiFriese, 1899 n. syn.i :An. dusmetiGuiglia,
Priesner, 1957 n.
1933 n. syn.; --An. helouanemas
syn.) (510)
futssinaIlliger, 1806 (Megilla) nomennudum
Benoist, seepuluerosaSmith
byssinoidzs
caelebs
Gribodo , 1924 An, P (-- A". alfinii Alfken,
1942 n. syn.) (464, 466)
(Cockerell), 1919 Am (An), E (506, 558)
caelestina
caaulea(Friese), 1905 Am (An), E (:An. ella Strand,
l9l4 fas coerulea
(sic) ssp.l n. syn.) (506, 548)
cafra (Friese), 1905 Am (An), E (515)
cala Strand, see calens(Lepeletier)
calcarataLepeletier, 1841 An, P (455, 486)
calceifua(Cockerell), 1911 Am (An), O (:An. tainaru
ssp.l n. syn.) (510, 511)
Strand, 1913 las calce'ifera
calceina(Cockerell), see calens(Lepeletier)
caldwelli(Cockerell), 1911 Am (An), O (515)
calens(Lepeletier), 1841 Am (An), E (:An. tonida
Smith, lB79; :An. cala Strand, 1914 [as tonida
Cockerell, 1936 [as
ssp.l n. syn.) :An. calceina
tonida ssp.l n. syn.) (504)
califurnicaCresson, 1869 An, N (:An. tarsataDours,
Cockerell, 1904 [as
1869 n. syn.) =An. subtarsata
tarsatassp.] n. syn.i :An. texanaCresson, lB72;
- Po. doursiiDalla Torre, 1896
[n. name for An.
tarsataDours, 1869, not TetraloniatarsataSpinola,
Provancher,
1B3B] n. syn. ) : An. quinquefasciata
1895 l,: S-fasciataProvancher, 18951i : An. nysimi
Cockerell, 1937 fas califtrnicassp.] n. syn. ) :An.
Timberlake, 1937 las califurnicassp.l
albomnrginata
n. syn.) (448, 45O, 454, 478-481, 557)
califurnieases
Michener, see ursina Cresson
caligataGerstaecker, see circulata (F abricius)
caliginosaKhg see robuila (Klug)
calua(Rayment), 1935 Am (At), A (515)
Smith
camboueiSaussure, see micrelephas
(Cockerell), 1911 Am (An), P, n. name for
cam.elorum
An. ruj,cornisFedtschenko, 1875, not Dours, 1869
(515)
cana(Walker), lBTl Am (An), P (:An. niuosella
Priesner, 1957, female, not male, n. syn.) (515)
Kirby
canadrnsisCresson, see bomboidzs
canarienszs
Saunders, see alluaudi Pérez
(Pérez), 1879 Am (An), q n. name for An.
candens
candidaPérez, t879, not Smith, lB79 (515)
candidaPérez, seecandens(Pérez)
candidn(Smith), 1879 Am (An), PO (515)
candidataGribodo, see hispanica(Fabricius)
candiùlla (Priesner), 1957 Am (An), P (515)
candidifronsCockerell, 1946 An, E (492)
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BIBI'uossarp olo"trs1{oc
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(tvg
'IuV 'ds 'u sToorg nsuapuaa(oc
'0Ig 'gç7)
d
,'gig
(srnoq) naysa{ eos 'uoretu"C nsua(oc
(tVç 'g7g 'VOg) ('ur(s 'u Z,T,6I
'1are>1Jo3
'asarrg s.saepîou'uV: !'ur(s 'u
1,061
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Gvç
olltuvlto 'uV:) a '(.tV) -V tçBI '(ql1urs) suo.trtuoc
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'ggf)
d
(t"fX) DîDuolees 'srnoe suetsauot
@Vg'0Ig
ssNslcs svsNv) do Àrlsus^rNn
sHJ
z,gç
SvsrnuATrcs
AND PHvrocENy
johnsoniCockerell, 1905 [as
lB73 (: Emphoropsis
mucidassp.li :An. aentralisMichener, 1936 [as
mucidassp.l)
crinipesSmith, seesalaiae(Panzer) (457, 460)
croceaBingham, see binghami(Schulz)
crocea(Kl.rg), 1845 Arn (Megilla), EP (504)
croceipes
Morawitz, see laraata Giraud
croceitarsas
Gussakovsky, 1935 An, P (496)
crotchiiCresson, 1B7BAn, N (=An. washingtoni
Cockerell, 1905) (445, 460-462)
crlsocnemrs
Morawitz, IBTB An, P ( = Po. chrysocnemis
(sic) Dalla Torre, 1896, and Friese, 1897) (482)
cuneataCockerell, see mimadaenaCockerell
cuniculanaFriese, 1914 An, E (4S6)
curta Provancher, see squamrnulosa
Dours (489)
curaicornrs
Hedicke, 1940 An, P (476)
custos(Dalla Torre), 1896 Am (Po), O, n. name for
An. uigilansSmith, lB7B, not Smith, 1860 (:An.
mcnamarae
Cockerell, 1929 las uigilazs ssp.] n. syn.)
(511)
cyaneotinctaCockerell, see qundratu(Cockerell)
cyanicollisFriese, seeacraemr (Fabricius)
cyanipennes
(Saussure), 1890 Am (An), E (506)
cygni(Rayment), 1931 Am (Ar), A (515)
cypriacaMavromoustakis, seeplumipes(Pallas)
cyrenaicaGribodo, see hispanica(Fabricius)
cyrtandrae
(Lieftinck), 1944 Am (An), O (513)
dalmaticaPérez,1902 An, P (--An. agamoides
Strand,
1915 las agamassp.] n. syn.) (462)
dalmatiensas
Strand, 1916 Un (Ar), P, n. name lor An.
subterranr.a
Gistel, 1857, not An. subtenaneaGerrnar,
1826 (450)
damrnersiTimberlake, 1937 An, N (464-466)
dammersi(Timberlake), 1937 Ha (Emphoropsis)n.
comb., N
darwini Cockerell, see utalkeri(Cockerell)
dawsoni(Rayment), 1951 Am (An), A (513, 515)
deceptrix(Priesner), 1957 Am (An), P (492')
decrepitallliger, seesubterraneaGermar
deiopea(Cameron), lB97 Ha (An), O
delicataCockerell, see mucorea(Kl"S)
deltoides
(Buysson), lB97 Am (An), P (515)
dentataRayment, see rubricata(Rayment)
dentilabrzsMorawitz, lB94 An, P (496)
dentipesllliger, seepodagraLepeletier
dentiuentras
(Rayment), 1951 Am (Asaropoda),A (515)
dentricrusMorawitz, see crassipesLepeletier
dtpressaFowler, 1899 Ha, N
deserticola
Morawitz, lB73 An, P (474-476)
desatorumGussakovsky, 1935, An, P (496)
dtsertorumPriesner, see arida Brooks
determinatuFriese, see reichardtiStadlemann
diloloensis(Cockerell), 1932 Am (An), E (504)
dimidiata Alfken, seeplurnipes(Pùlas)
dimidiozonataDours, see sichelii Radoszkowsky
disconotaLieftinck , 1974 Ha, O
disparLepeletier, 1841 An, P (: An. specæsa
Friese,
1919 fas disparssp.] n. syn.l :An. niaeohirta
Friese, 1922 [as disparssp.] n. syn.) (462, 464,
465)
disparilis Friese, 1922 An, P (: An. bicalcarataHedicke,
1936 n. syn.) (464-466')
disrupta(Cockerell), 1920 Am (An), E (504)
oF THE ANrnopHoRINn
Bnns
s63
distinguendaSpinola, lB51 (An) :Diadasra (Anthophoridae; Emphorini)
diunsipesFriese, 1922 An, E (: An. fulleri Brauns,
1929 n. syn.) (462)
diaesDours, see dufourii Lepeletier
dohntyi(Gribodo), 1894 Am (Ar), O (510, 511)
domicola(Cockerell), see atrocincta(Lepeletier)
domingensas
Lepeletier, see maculicornis(Lepeletier)
dorsalisVachal, 1909 An, T (:An. ecuadoria
Friese,
1922 n. syn. ) :An. trichromaFriese, 1925 n. syn.;
=An. incertaJanvier,1955 n. syn.) (468)
dorsimaculaDufour, see qwdricolor (Erichson)
doursiana(Friese), 1897 An (Po) (:An. uentilabrisof
Dours, 1869, not Lepeletier, 1841) (496)
doursii Dalla Torre, seecaffirnica Cresson
doaeriCockerell, see zonata(Linnaeus)
drewseniiSichel, see rufolanataDours
dubiaEversmann, 1852 An, P (:An. saussurei
Fedtschenko, lB75 ; -- An. semperiFedtschenko,
1875i : An. faddei Radoszkowsky, 1BB2i : An.
albomaculata
Radoszkowsky, I 874; -- An. carbonaria
Morawitz, 1876 n. syn.; :An. ctrbnusFriese, 1919
las semperissp.] n. syn.) (473, 476)
dufouriiLepeletier, 1841 An, P (: An. tectaSmith,
1854, type locality of tuctuis not the New World as
in description but probably southern France from
color form of type; --An. diuesDours, 1869; :An.
nigropygaStrand, l92l las dufoura(sic) ssp.J n.
syn.i =An. fasciculipesAlfken, 1935 n. syn.) (438,
4 7 O ,4 7 l , 4 g g )
dulciJna(Cockerell), 1926 Am (Ar), O, (:An. senahai
Yasumatsu,1935 [as cingulata
ssp.l;:An. subYasumatsu, 1965 las senahafssp.] n. syn.)
flaaescens
(511)
durnetorumPanzer, seefurcata (Panzer)
duplicicornesllliger, 1806 (MegiUQ nomennudum
dusmetiGuiglia, see byssina(Klug)
eburnea
Radoszkowsky, 1876 An, P (472, 476>
ectndoriaFriese, see dorsalrsVachal
edwardsiiCresson, 1B7BAn, N (:An. gohrmanae
Cockerell, 1903 n. syn.):An.
coloradmsis
Michener, 1936 fas gohrmanaessp.] n. syn.)
(460-462, 488)
ffissa Heyden, 1862 Un (An), P (fossil; it is
impossible to be sure that this is a speciesof An.
[Zeuner and Manning, 1976]) (524)
ekuiaensis
Cockerell, 1908 An, E (492')
elbanaPriesner, 1957 An, P (492)
elefas(Friese), 1916 Deltoptila(An), T
elegans
(Smith), 1859 Am (An), O (511)
(Lieftinck), l9++ Am (An), O (513)
elephas
elgonica(Cockerell), 1946 Am (An), E (504)
elimataCockerell, 1933 An, E (:An. infracana
Cockerell, 1933 n. syn.) (492)
ella Strand, see caeruleaFriese
elseiBrooks n. sp. Am, E (438, 506, 548, 549)
emarginalaTimberlake, pee rhodothoraxMichener
emmdataSmith, seecingulata(Fabricius)
Dours
ensenadençes
Cockerell, seesquarnmulosa
Brooks n. sp., Am, A (438, 515, 553-555')
epaphrodita
epeolinallliger, 1806 (A") : Ammobatesuinctus
Gerstaecker, 1869, according to Gerstaecker, 1869
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SysreuATlcs
AND PHyrocENy
footei Crawford, l9l4 An, T (466, 468)
forbesi Cockerell, t907 An, N (460, 462)
franciscanaCockerell, 1949 An, T (496)
fratucula Gribodo , 192+ An, P (466)
fratnna Bingham , 1897 An, O (490-492)
freimuthi Fedtschenko, 1875 An, P (--A". oschanini
Fedtschenko, 1875; : An. gertaliRadoszkowsky,
nomennudumby Friese, 1909) (496)
froggatti Cockerell, seepreissi(Cockerell)
frontata Say, 1837 Un (At), N (450)
Lieftinck, see alluaudi Pérez
fuerteuenturae
fuliginosa Illiger, see qrudrifasciata(de Villers)
fuliginosaMorawitz, lB94 An, P (466, 557)
fullai Brauns, see diuersipesFriese
fulaa (Smith), 1879 Deltoptila(Ha), T
fulua Eversmann, 1852 (Saropoda): E4eoloidescoecutiens
(Fabricius), 1775 (Anthophoridae; Epeoloidini)
fuluata Fabricius, 1804 (Megilk) : Nomia (Halictidae)
fuluicaudaTimberlake, 1937 An, N (450, 476, 479)
fuluicollis Timberlake, seeafabilasCresson
fuluifrons Smith, see taurea(Suy)
fuluipes Cameron, see khasianaSchulz
fuluipesEversmann, 1846 An, P (476)
fulaiscopaAlfken, see semirufaFriese
fuluitarsis Brullé, lB32 Att, R (: Megilla personata
Illiger, 1806, nomennudum; -*An. nnsutaLepeletier,
l$+l ) : An. arietinaDours, 1869; : An. euris
Dours, 1869 fas personatassp.l) (457, 460, 556,
558)
fuluitecta Kirby, see inclfia Walker
fuluiuentris Zetterstedt, 1B3B(A") :Megachile (Mega'
chilidae)
fulaocinereaDours, seeplagiatu (Illiger)
fuluodimidiataDours, 1869 An, P (491, 492)
fuluohirta Meade-Waldo, see insularis(Smith)
fumipennesAifken, see aidua (KlnS)
Kirby
fumipenmsSwenk, see bomboides
furcata (Panzer), l79B An (Apis), P (:Apis dumetorum
Panzer, l79B; :Megilla noraegicaNylander, 1852
- Po. caucasicus
Friese,
lasfurcala ssp.] n. syn.l
lB97 lasfurcalzs ssp.] n. syn.l - Po. montislinguarum
Schulz, 1906 lasfurcalru ssp. and n. name for Po.
Friese, 1897, not An. caucasica
caucasicu.r
Radoszkowsky, lB74l n. syn. ) : An. atrata
Hedicke, 1929 lasfurcalassp.] n. syn.l :An.
Hedicke, 1929 [asfurcata ssp.l n. syn.;
caucasicola
: An. obscurella
Alfken, original description not
found) (487 -4Bg)
furcotibialesWu, 1985 An, O (476)
fuscipennisSmith, 1854 (An), N, :Melissodes bimaculata
(Lepeletier), 1825 n. syn. (Anthophoridae; Eucerini)
Priesner, 1957 An, P (492')
galalensis
gallica Dalla Torre & Friese, see quadricolor(Erichson)
garrula(Rossi), 1790 Am (Apis), P (:An. bomfutlans
Mocsâry, 1879) (5O4, 505)
galti Spinola, 1851 (A") : Eucerinodagayi (Spinola)
(Anthophoridae; Eucerinodini)
gedzeniRadoszkowsky, seeglasunoaiMorawitz
gemellaMorawitz, 1B7BAn, E (492)
germabica
Radoszkowsky, 1893 Un (An), P (450)
gntali Radoszkowsky, seefreimuthi F edtschenko
ghigii Gribodo , 192+ An, P (464)
giganteaFriese, seehirtiaentrisFriese
oF THE ANtHoPHoRrNn
BuBs
565
gigas(Friese), 1922 Am (An), O (513)
gilberti Cockerell, see cingulata(Fabricius)
glasunouiMorawitz, lB94 An (:An. gedzeni
Radoszkowsky, nomennudumby Friese, 1909) (476)
glaucaAlfken, seeandresi(Friese)
glaucopisFriese, 1905 An, E (476>
globosaFabricius, 1793 (Apis), T, : Exomalopsis(Atthophoridae; Exomalopsini)
glycyrrhizaa
(Gussakovsky), 1935 Am (An), P (511)
godofredi(Dours), 1869 Am (An), E (:An. modesta
Smith, lB79 n. syn.) (508, 5lO, 55tr 556)
Cockerell, see edwardsiiCresson
gohrmanaa
gracilipes
Morawitz, lB73 An, P (452, 473-476' 489)
graeca(Alfken) , l9+2 Am (An), P (504)
grandicep.r
(Friese), 1905 Am (An), E (:An. leucophora
Cockerell,
Cockerell, 1910 n. syn. i : An. semigrisea
1930 n. syn.) (502, 504)
grandidieraCockerell, see heydeniiSaussure
grandisLepeletier, see hispanicaFabricius
grayella(Rayment), 1944 Am (At), A, publication
date is not 1942 (512\
grelaeCockerell, see uestitaSmith
(Fabricius)
griseaChrist, seepubescens
griseaSchletterer, 1890 (A") -Suastridesmelanura
(Spinola), 1851 (Anthophoridae; Eucerini)
griseaFabricius, 1804 (4") : Halictus (Halictidae), not
Apit griseaFabricius, 179+
(Friese)
grisella(Cockerell), see obscuriceps
griseotecta
(Cockerell), 1946 Am (At), E (508)
griseouestitaCockerell, see uestitaSmith
grisescens
(Rayment), 1931 Am (An), A (515)
Ducke, rcA7 (Po) : Melitoma (An'
grisescens
thophoridae; Emphorini)
Cameron, seepilifrons Packard
guachalae
guigliaeDusmet, see riaolleti Pérez
guinea(Strand), 1912 [as aiuidassp.l Am (An), E
(506, 558)
(Popov), 1946 Am (An), E, n. name for
gussakoasÈyi
An. pygmaeaGussakovsky,1935, not Dours, 1869
(515)
(Dours)
Dalla Torre, see aurulentocaudatu
habropodoides
hackeri(Rayment), I 947 Am (At), A (511)
haef,iguiFriese, 1905 (A") : Eucara (Anthophoridae;
Eucerini)
hanitschi(Meade-Waldo), 1914 Am (At), O (513)
hanseni
Morawitz, 1BB3An, P (484, 486)
harmalaeMorawitz, LBTBAn, (476)
harttigi (Alfken) , 1926 Am (An), P (504)
hastulaVachal, 1909 Un (An), E (450)
haworthanaKirby, seeretusa(Linnaeus)
hediniAlfken, 1936 An, O (: Heliophila latitarsalisWu,
1985 n. syn.) (470\
Priesner, 1957 fas concinnassp.l At, P (492)
hegasica
Fedtschenko, 1875 An, P (:An. hilgendori'
heinemanni
Cockerell, 1911 n. syn.) (466)
Pérez, 1910 An, P (496)
heliopolitensis
Priesner, see byssina(Klug)
helouanenses
Sichel, see acraensis(Fabricius)
hemithoracica
herbstiFriese, see ruf,caudisCameron
Cockerell, see xanthostomaCockerell
heteropoda
(Hu) n. comb., E
hefi"enii(Saussure), 1890 Pachymelus
Cockerell, 1916 n. syn.)
grandidiere
(: Pachlmelus
(558)
hilaris Smith, lB79 An, T (468, 469)
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AND PsvlocENy
lanata(Kl.rg), 1845 An (Megilla), P (: An. canescens
Dours, 1869 las senesrrnJ
ssp.] n. syn.l :An.
aennabilisCockerell, 1911 n. syn.) (457 r 460)
lanataFabricius, 1775 (Apit) : Megachile (Megachilidae)
langi (Cockerell), 1935 Am (An), E (502)
laraataGiraud, 1863 An, P (:An. croceipes
Morawitz,
lB76 n. syn.) (474, 476)
lata (Cameron), 1905 Pachymelus
(Hu) n. comb., E
laticepsFriese, 1905 (A") : Eucara(Anthophoridae;
Eucerini)
laticepsFriese, see aestitaSmith
laticinctaDours, 1869, seesubtenaneaGermar (555)
latigenaMorawitz, 1886 An, PO ( -- An. reinigi
Hedicke, 1931) (482)
latipesFriese, seeatrocinctaLepeletier
latitarsalrsWu, see hedini Alfken
latizona(Spinola), 1B3B Arn (Saropoda),P (510)
leoninallliger, 1806 (MegiWq)nomennudum
leonisCockerell, 1933 An, E (476)
leopoldiCockerell, seefelina (Friese)
lepidaEversmann, see areos(Pallas)
Dours, 1869 Un (An), P (450)
lepidodea
Lieftinck, see malaccensas
leptocoma
Friese
lesquerellaa
(Cockerell), lB97 An (Po), N (460, 462,
556)
leucocephala
Friese, see annls (Vachal)
Torre, 1896 Un (Po), P, n. name for
IeutomclaenaDalla
An. melaleuca
Walker, lB7l, not Lepeletier, 1841
(450)
leucomelaena
Illiger, see albigena(Lepeletier)
Pérez, lB79 An, P (464)
leucophaea
leucophora
Cockerell, seegrandicepsFriese
leucopyga
Friese, 1911 An, E (476)
leucorhina
Cockerell, l9l7 An, P (460, 464)
leucospilaCockerell, see epicharifurmisGribodo
Hedicke, l9+0 Un (An), P (450)
leucosticta
leucostomella
Cockerell, see urbanaCresson
libuica (Cockerell), 1930 Am (An), E (515)
librata llliger, 1806 (MegiUa) nrmennudum
libyphaenicaGfibodo, 1893 An, P (462)
lieftinckiBrooks n. sp. Am, O (438, 512, 513,
550-552)
lilacineCockerell, see cingulata(Fabricius)
limassolicaMavromoustakis, An, P, original description not found (464)
(Hu) n. comb., E
limbatus(Saussure), 1890 Pachymelus
linsleyiTimberlake, l9+l An, N (479,481)
liriope (Bingham), see mucorea(Klug)
lisalanaCockerell, see cincta(Fabricius)
lisbonensas
Cockerell, seeplumipes(Pallas)
litoranaPriesner, see quadrifasciata(de Villers)
liturata Lepeletier, see ûnit Brullé
liuingstoneiCockerell, see tetradontaCockerell
loczyiMorawitz, lB92 An, O (496)
loewi Fedtschenko, 1875 An, P (492)
lomamica(Cockerell), 1936 Am (An), E (504)
longipesMorawitz, lBB4 An, P (492)
longmani(Rayment), 1947 Am (Ar), A (511)
Iongula(Rayment), 1947 Am (An), A (511)
(Fabricius)
loueridgeiCockerell, see acraensis
lucknoa
iensesRadoszkowsky, see subcoerulea(Lepeletier)
luluana Cockerell, see triangulifera(Cockerell)
oF THE ANtHopHoRrNe
Bnns
567
lumbwanaCockerell, 1946 An, E (492)
lusitanicaFriese, seeffinis Brullé
lusoriaCockerell, see oldi Meade-Waldo
luteolaRayment, see chlorocyanea
(Cockerell)
luteodimidialaDours, 1869 Un (An), T (450)
lutescens
Walker, lB7 | Un (An),P (450)
lutulenta(Klug), 1845 Am (Saropodo),P (515)
luzonica
(Cockerell), 1914 Am (Ar),O (513)
maclachlani
(Fedtschenko), 1875 Am (An), P ( --An.
meraensis
Cockerell, 1933) (508, 510)
macrognathaGerstaecker, lB7 | (Eucna) -- Eucara (Anthophoridae; Eucerini)
mncroleuca
(Cockerell), 1922 Am (An), O (515)
maculicornes
(Lepeletier), 1841 Am (An), E (:An.
Lepeletier, 1841) (515)
domingemes
maculifrons
Cresson, 1879 An, N (496)
maculigera
Priesner, 1957 An, P (474, 476)
madecassa
(Saussure), 1890 Am (An), E (504)
madnaeSichel, see qundrifasciata(de Villers)
Fedtschenko, seesaaignyi(Lepeletier)
magnilabras
magrettii(Bingha*), 1897 Elaphropoda(Hu), O (- Ho.
fletcheriCockerell, 1920)
maiellaLieftinck, see tainanicola(Strand)
malabaricaGmelin, 1790 (Apit), E, : Nomia curuipes
(Fabricius) I 78l (Halictidae)
malaccensas
(Friese), 1918 [as aillosulassp.] Am
(Antophora[sic]), O (: Am. leptocoma
Lieftinck, 1956
n . s y n . ) ( 5 1 2 , 5 1 3 15 5 6 )
malelanaCockerell, see circulata(Fabricius)
malenominalruSchulz, seefestiaus(Dours)
mnngkamensis
Wu, 1982 An, P (478)
nnnni Mocsâry, see quadricolor(Erichson)
marginataSmith, 1854 An, NT (:An. histrio Dours,
1869; - Po. cleomisCockerell, 1896; : An. popenoei
Cockerell, l9+9 n. syn. i : An. peleniCockerell,
l9+9 n. syn.) (479,481)
marqueti(Pérez), 1895 Am (An), P (515)
marsupoda
Christ, l79l Un (Apis), P (450)
martensiFedtschenko, 1875 An, P (475, 476)
matopoenses
Cockerell, 1933 An, E (492)
mnuritanica(Benoist), 1950 las altnnazr ssp.l A* (A.),
P (505)
mcnamarae
Cockerell, see custos(Dalla Torre)
mtdicorumCockerell, seepunctifrons(Walker)
mediocindruCockerell, l9l7 Pachymelus,E
mediorufa(Cockerell), 1920 Am (An), E (504)
mediozonataLaboulbene,1870 Un (An), P (450)
meditnraneaAlfken, see qwdrifasciata (de Villers)
meganhinaCockerell, 1910 An, O (: An. soluta
Cockerell, 1910 [as meganhizassp.] n. syn.) (438,
482-484)
megasoma
Cockerell, see armntaFriese
melaleucaWalker, see leucomelaena
(Dalla Torre)
melaleuca
Lepeletier, 1841 Un (Ar), T (450)
melanocephala
Morawitz, 1894 An, P (466)
melanocera
(Cockerell), 1946 Am (An), E (504)
melanodonta
(Cockerell), 1933 Am, E (504)
melanognatha
Cockerell, 1911 An, O (457, 4591 460)
melanopoda
llliger, 1806 (MegiUQ nonun nudum
Dours
melanopsCockerell, see squammulosa
Fedtschenko, 1875 An, P (476)
melanopyga
Dours, 1869 Un (An), T transferred to
melanopyrrha
Deltoptilaby LaBerge & Michener, 1963, but
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AND PHYTocENY
nigriuentrisZetterstedt, 1B3B(A") - Osmia(Megachilidae)
Friese, see acraensis(Fabricius)
nigriaentres
Dours, 1869 Un (An), Unknown origin
nigroaeruginosa
(450)
nigrociliataPérez, 1895 An, P (486)
nigrocinctaLepeletier, seesubtenaneaGermar
nigrocinctaProvancher, seeflaaocinctaHuard
nigrocinctulaDours, see aentilabrisLepeletier
Friese, 1909 Am (An), E (506)
nigroclypeata
nigrofuluaLepeletier, seeplumipes(Pallas)
Lucas, see robuila (Klug)
nigromaculata
nigromixtaCockerell, see mimadaena(Cockerell)
nigropilosa(Friese), 1896 Am (Po), P (515)
nigropygaStrand, see dufourii Lepeletier
Friese, see pedataEversmann
nigroscopacea
nigrotarsaWr, 1979 Elaphropoda,P
nigroaittalaDours, see balneorumLepeletier
nitidula Dours, 1869 (A") : Exomalopsis(4.thophoridae; Exomalopsini) according to Timberlake, 1980
niuea(Lepeletier), 1841 Am (An), E (515)
niaeata(Friese), 1905 Am (An), E (:An. amphigltnops
Cockerell,1933 fas niuealassp.] n. syn.) (438,
508-5 10)
Brèthes
Friese, seearequipensis
niaeiceps
Hedicke, l9+0 An, P (496)
niueifacies
Friese, 1919 [as robustassp.] An, P (466)
niaeiaentras
(Smith), 1854 Am (An), P (:An. biniueocinda
maculifna Walker, l9Tli :Po. bimnculoszsDalla
Torre, 1896, n. name for bimaculifua Walker
female, no. 284; - Po.f,auicepsFriese, 1899 [as
ssp.] n. syn.) (498, 5O2r 509-511)
albigenus
niueohirtaFriese, see dispar Lepeletier
niuescens
(Cockerell), 1936 Am (An), E (:An.
microleuca
Cockerell, 1937 n. syn.) (510)
niuosellaPriesner, see cana(Walker)
Brooks n. sp., A*, E (438, 444, 496,
nonconforma
515, 552, 553, 556)
lrlylander, seefurcata (Panzer)
noruegica
nubana(Cockerell), 1946 Am (An), E (504)
nubica(Lepeletier), 1841 Am (An), EP (:An.
Friese, 1909 fas nubicassp.] n. syn.)
congoensis
(506, 5O7)
(Cockerell), 1930 Habrophorula(Ha), O
nubilipennas
nubiterraeViereck, see tuminalis Cresson
nudn(Radoszkowsky), 1BB2Elaphropoda(An) n.
comb., O, ( : Ha. moelleriBingham, lB97 n. syn.)
nudataProvancher, see terminalisCresson
nurranaCockerell, 1931 An, E (496)
obesaGiraud, see balneorumLepeletier (557)
obscurellaAlfken, seefurcata (Panzer)
(Friese), 1905 las circulalassp.] A* (An), E
obscuriceps
(:An. grisellaCockerell, 1932 n. syn.) (504)
obscurusFriese, seehoaa(Saussure)
obscurusFriese, see retusa(Linnaeus)
(Illiger), 1806 (A") nomennudum
obsoleta
obtusispinaWu,1982 An, P (478)
Cresson, 1869 An, N (45O, 478)
occidentalis
occipiteMorawitz, seepedataEversmann
occultaHedicke, 1938 An, P (496)
(Pérez), lB79 Am (An), P (504, 5O5)
ochroleuca
Friese, see ocularas(Saussure)
octomaculatus
oF THE ANTHOPHORINN
BBBS
s69
(Ha) n. comb., E
ocularis(Saussure), 1890 Pachymelru
(= PachtmelusnigerFriese, 1922 n. syn.;
: Pachymelus
Friese, 1922 las B-macuoctomaculatus
ciliatus Friese, 1922 n.
latusl n. syn. ; : Pachymelus
syn. ; - Pachymelus
flauithoraxBenoist, 1962 n. syn.)
odonturaCockerell, see africana(Friese)
ogiluiei(Cockerell), 1932 Am (An), E (504)
oldi Meade-Waldo, l9l+ An, E (-- A". lusoriaCockerell, l92l las oldi ssp.l n. syn.) :An. albopictula
Cockerell, 1933 n. syn.) (492)
olgaeFedtschenko, lB75 An, P (474-476)
Wu, 1979 Ha, O
omeiensis
omissa(Priesner), 1957 Am (An), P (504)
Morawitz, 1876 An, P (474, 476')
onosmannæ
optimaCockerell, see armataFriese
(Lepeletier), 1841 Ha (An), P
oraniensis
orbifronsLieftinck, 197+ Ha, O
orientalisMorawitz, 1B7BAn, P (: Po.f,aaiuentris
ssp.D e62, 464,489)
Friese, 1896 fas orientalrs
ornataGistel, 1857 Un (An), P (450)
orophilaCockerell, 1910 An, P (: An. pseudorophila
Wu, l9B2 n. syn.) (484)
(Saunders), 1904 An (Po), P (484, 486)
orotaaae
oschaniniFedtschenko, seefreimuthi F edtschenko
lxrgnnaDours, seerobuila(Klug) (556' 557)
pachyodonta
Cockerell, 1923 An, N (494, 496)
pachypo
da C ockerell, see quadrimaculata(Panzer)
pacijla Cresson, 1B7BAn, N (--An. carbonaria
Cresson, lB79 [dark form] i : An. ignauaCresson,
lB79; - Po. infunalis Dalla Torre 1896 [n. name
Cresson, 1879, not Morawitz, 1876]
for carbonaria
Cockerell, 1905 n. syn.
n. syn. l :An. coraicolor
[dark form] , :An. subignauaCockerell, 1929 n.
syn. | :An. pernicisTimberlake, 1951 n. syn.)
(450, 466>
padiolaVachal, 1909 Un (An), E (450)
pagdeniLieftinck, 1956 Am, O (513)
pahangenças
Cockerell, seependleburyi(Cockerell)
pahangemas
Meade-Waldo, see himalajensis
(Radoszkowsky)
palestinenças
Hedicke, seeplumipes(Pallas)
pallescens
Morawitz, 1895 Un (An), P (450)
pallida (Timberlake), 1937 Ha (Emphoropsis)n. comb.,
N
pallidicinctus Cameron, see aestitaSmith
palmipesRossi, seeplumipes(Pallas)
pamirica Hedicke, see robusta(Kl"S)
pamiricolaHedicke, seeplagiata (Illiger)
paradoxaBrooks n. sp., Am, E (438' 506' 516,
552-554, 556)
paranensai
Holmberg, 1903 An, T (:An. paranensis
Holmberg, 1887, nomennudum; :An. saltensis
hirpexVachal,
Holmberg, 1903 n. syn.i:An.
1 9 0 4 )( 4 6 6 , 4 6 S )
parapulchra(Rayment), l9+7 Am (An), A (511)
parhypateLieftinck, 1975 Am, O (511)
parietinaFabricius, seeplagiata(Illiger) (478r 489)
pascoensis
Cockerell, see morrisoni(Cresson)
passaini Sichel, see tarsala(Spinola)
patruelisCockerell, 1931 An, O (460)
pauperataWalker, lBTl Un (An), P (450)
pedataEversmann, 1852 An, P (:An. occipiteMorawitz, 1B7Blas pedatassp.l n. syn.) :An.
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SySTeuATICS AND PgyTOGENY OF THE ANTuoPHoRINB
Michener,
Michener, 1936 n. syn. i :An. utahensis
1936 fas chloroprssp.] n. syn.) (438, 463-466)
posticaVachal, 1910 An, E (496)
potanini(Morawitz), lB90 Am (Ar),P (515)
praecox
Friese, 1909 las krebsissp.] At, E (-Ar.
praecoxFriese, 1911 las wartmannissp.]; Friese
(1911) said he originally intended to describe
praecoxas ssp. of wartmanni and not krebsi; : An.
aliceaeCockerell, 1932 n. syn.) (489' 492)
preissi(Cockerell), 1910 Am (An), A (:An. froggatti
Cockerell, 1914 [as preississp.] n. syn.) (515)
pretiosaFriese, 1919 lasfuluitarsisssp.] An, P (466)
priesneriAlfken, 1932 An, P (486)
proboscifua
Lieftinck, 1956 A-, O (513)
proceraCosta, seesubturaneaGetmar
proserpinaGribodo, see himalajemas
(Radoszkowsky)
proximaMorawitz, 1894 An, P (496)
prshewalskiiMorawitz, 1BB0An, O (:An. pilosa
Morawitz, lBB0) (47O, 474, 476)
pruinosaSmith, 1854 An, P (--An. stefaniiPérez, 1902
n. syn.) (464)
pseudobasalis
Strand, seearmatuFriese
pseudobomboides
(Meade-Waldo), 1914 Am (An), O
Wu, 1983 n. syn.) (512' 513)
bombiomorpha
(:A*.
p seudo
coccinaMeade-Waldo, see africana (Friese)
pseudorophila
Wu, see orophilaCockerell
pseudosicula
Hedicke, seequadricolor(Erichson) (556)
pubescens
(Fabricius), lTBl An (Apis), P (: Apit grisea
Christ, l79l) : An. f,abelliferaLepeletier, 1841;
--An. labronigroLebedev, 1932
ssp.]
fas pubescans
numennudum; :An. flabellipesLichenstein, 1871)
(470,489)
pulcherrimaBingham, seeplagiata (Illiger)
pulcherrinmY,{u, 1985 Elaphropoda,O
pulchra(Smith), 1854 Am (An), A (:An. townleyella
Rayment, l9+7 las pulchlassp.] n. syn.) In the
British Museum the female type of An. pulchra
Smith, B. M. Typ. Hy-. 178666a, is not the type
according to D. Baker. The type was apparently
repinned and lost and the specimen in its place is
(Smith) (511).
Am. niaeocinda
pulsellaDours, 1869 Un (An), T (450)
puluerea(Walker), lB7 | Am (An), P, see note under
salaiae(Morawitz) (5 15)
Benoist,
pulaerosa
Smith, 1854 An, P (:An. futssinoides
1950 n. syn.) (492)
punctata(Rayment), 1931 Am (Asaropoda),A (515)
punctifrons(Walker), lB7 | Am (An), P ( : An. medicnrumCockerell, 1910 n. syn.i :An. politifrons
Cockerell, 1946 n. syn.) (504)
punctilabrisPérez, lB79 An, P (464)
puttalama(Strand), 1913 las zonatassp.l A- (An), O
(510,511)
kouskyi(Popov)
pygmaeaGussakovsky, seeg'Ltr.ça
pygmaeaDours, 1869 Un (An), T (450)
pltgmaeaMeade-Waldo, 1914 An, E (: An. torridella
Meade-Waldo, l9l4 n. syn. i : An. natensisCockerell, 1935 n. syn.) (492)
pyralitarszs
Dours, see ursinaCressson
pyramidalrsKirby, seealbigena(Lepeletier)
f u r o f u g aD o u r s , 1 8 6 9U n ( A n ) , E ( 4 5 0 )
pltrozonala
Dours, 1869 Un (An), T (450)
quadrata(Cockerell), 1911 [as albigezassp.] A- (At)'
BnnS
57r
Cockerell, 1920 n. syn.) (497 ,
O (: An. cyaneotincta
502, 504,509)
quadricinctaFabricius, l79B (Apit) - Halictus (Halictidae)
(Dalla Torre &
quadricinctaEversmann, seeeaersmannii
Friese)
(Erichson), 1840 An (Megilla), P (-An.
quadricolor
atrifronsSmith, 1854 n. syn.) :An. rectangula
Costa, 1863i :An- smithiiDours, 1869 n. syn';
:An dorsimacula
Dufo:.r, nomennudumby Dours,
-1869; : An. manni Mocsâry, 1BB3; Megilla gallica
Dalla Torre & Friese, 1895 [n. name for An. smithii
Dours, 1869, not Cresson, 1869] n. syn. i :An.
pseudosicula
Hedicke, 1929 [n. name for sicula
Smith, 1854, male only] n. syn.) (476)
(de Villers), l7B9 Am (Apis), P (:Megilla
quadrifasciata
scalarisllliger, 1806, nlmen nudum,n. syn.;
- Megilla fuliginosa llliger, 1806, nomennudunt, tr.
Sichel, 1867 [red
syn. [red form] i = An. maderae
Dours, 1869 n. syn. i :An.
form] i :An. albescens
Radoszkowsky, 1893; : An. farinosa Almeruensis
Alfken, 1927
fken, 1926 n. syn. ', : An' mediterranea
Cockerell,
n. syn. [red form] i : An. tenerifensas
ssp.] n. syn. [dark form];
1930 fas quadrifasciata
= An. litoranaPriesner, 1957 n. syn. i : An. klugi
Priesner, 1957 [n. name for An. farinosaAlfken,
1926,not Klug, 1845] n. syn.) (449-453, 456,
503-505)
(Panzer), l79B An (Apis). P ( -Apis
quadrimaculata
uulpinaPanzer, 1798, not Christ, l79l; -APis
Kirby, 1802; --An.mixta Lepeletier, 1841
subglobosa
-n. syn. ; An. uaraLepeletier, 1B4l n. syn. ', : An.
Gribodo , 1873 n. syn. i :An. pachypoda
segusina
Cockerell, 192+ fas uulpinassp.] n. syn.l :An.
alticolaHedicke, 1931 [as uulpinassp.] n. syn.)
(438, 485, 486)
quadristrigataDours, seecrassipesLepeletier
ciata Provancher, see califurnicaC resson
quinquefas
raddeiMorawitz, 1876 An, P (476,489)
(Dalla Torre), 1896 Ha (Po), O, r. name
radoszkowskii
for Ha. montanaRadoszkowsky, 1BB2, not An.
montanaCresson, 1869
Fedtschenko, 1875 An, P (466r 486)
radoszkowsfuii
(Ha) n. comb., E
(Saussure), 1890 Pachymelus
radouae
rapida(Smith), lB79 Am (An), E (515)
KirbY
raui Rohwer, see bomboides
Costa, seequadricolor(Erichson)
rectangula
regalis(Cockerell), 19+6 Am (An), E (506)
E (-An.
reichardtiStadelmann, 1B9BPachymelus,
Friese, 1905 n. sYn.)
determinala
reinigi Hedicke, see latigenaMorawitz
repletaDours, 1869 Un (An)' P (450)
retusa(T,innaeus), 1758 An (Apis). P ( -Apis pennipes
Forster, 1770, nomennudum; :APis aestiualisPantzer,
-1801; - Afis haworthanaKirby, 1802; An. intermediaLepeletier, 1841; -Megilla monachaBrichson,
Morawitz, lB71 n'
1849 n. syn. i :An. ruthenica
syn. ) :An. meridionalisPérez, lB79 las retusassp.]
Friese, 1896 las retusu'rssp'];
n. syn. i : Po. obscurus
- Po. sareptanus
Friese, 1896 las retususssp.]; :An.
Cockerell, 1911 n. syn. i :An. fasciata
retusifurmr^r
Alfken, 1913 las retusassp.] n. syn.; :An.
ssp.] n. syn';
Hedicke, 1929 las aestiaales
baicalensis
(3çù a '@tt@aW)un 9081 ,retr11 sqruas
(sneeuurl) osnTa^t
aes ,z1rrvrurotr
tr Drzuau?ru
,rV
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SysrpuATrcs
AND PHyrocENY
senilisEversmann, 1846 An, P (:An. aetulaEversmann, I 852; -- An. eaerstrunnr
Radoszkowsky, I 869)
(462)
senilisWalker, see aegyptiaca(Dalla Torre & Friese)
sngia (Nurse), 1904 An (Po) , O (464)
sesquicinda(Erichson & Kltg), lB+2 Am (Megilla), O
(-- Apis bicinctaFabricius, 1793, not Schrank, lTBl;
--An. indicaRadoszkowsky, 1BB2n. syn.) (438,
499, 503, 505, 557' )
shafryella(Rayment), 1947 Arn (At),A (511)
Priesner, 1957 An, P (492)
shagrensis
shestakoar
Gussakovsky, 1935 An, P (496)
sibiricusGussakovsky, see terminalasCresson
sicheliiRadoszkowsky, 1868 An, P (:An. pnsica
Radoszkowsky, 1876; : An. dimidiozonataDours,
Nurse, 1904 n. syn.) (460,
1869; - Po. cnnnetcus
464)
sichuanensrs
(Wu), 1987 An (AnthomegillQ,O (484)
Wu, l9B7 Ha, O
sichu,anemes
sicula Smith, seeplumipes(Pallas) (556)
siewersiMorawitz, lB76 An, P (486, 489)
s i g n a tB
a r o o k sn . s p . A n , N ( 4 3 8 , 4 4 8 , 4 7 8 , 4 7 9 ,
4gl, 499, 519, 535-539)
simbana(Cockerell), 1936 Am (At), E (515)
simia Dours, 1869 Un (An), origin unknown (450)
similisFedtschenko, 1875 An, P (486)
simillima Cresson, see ursina Cresson
simplexMorawitz, seeplagiata (Illiger)
simplicipesMorawitz, seeplagiata (Illiger)
Alfken, 1937 Ha, O
sinensis
sinensisWu, see wuu Brooks
sinensis(Wr), 1982 An Clisodon,P (489)
sintntitarsasFriese, see uestitaSmith
(Friese), 1909 Am (An), E (--An. sjostedti
sjoestedti
Friese, 1909)(515)
smirnouiKuznetzov-Ugamsky, seeplagiata (Illiger)
smithii Cresson, see montanaCresson
smithiùDours, seequadricolor(Erichson)
socia(Klug), 1845 Arn (Meg;llo), P (505)
Kirby
sodalisCresson, see bomboides
soikaiBenoist, 1961 An, E (492')
solitaria Rado szkowsky, seefemorata (Olivier)
Kirby
solitariaRitsema, see bomboides
solskliFedtschenko,1875 An, P (476)
somalica(Magretti), 1B9BAm (Po), E (:An. unicincta
Friese, 1922 n. syn.) (5O2,505)
sordida(Rayrnent), 1931 Am (Att), A (515)
sordidula(Rayment), 1931 Am (An), A (515)
sornrMocsâry see micrelephasSmith
sororPérez, seeplumipes(Pallas)
snrorPérez, see imitatrix Cockerell
speciosa
Friese, seedispar Lepeletier (465)
sperryiCockerell, see terminalisCresson
spilostoma(Camero.), 1905 Am (Podalarius[sic]), E
4(504)
spinicaudaCockerell, see rufoaestitaCockerell
spinacoxa
Brooks n. sp., Ar, P (438, 470, 540, 541)
spinipes(Friese), 1899 An (Po), P (476)
spinitarsisWu, 1982 An, P (484)
spinolanaPriesner, 1957 An, P (460, 462)
spodiaDours, 1869 Un (An), T (450)
sponsaSmith, see abrupta Say
squalensDours, seeplumipes (Pallas)
oF THE ANtHoPHoRINn
Bnns
573
squalidaLepeletier, see bimaculata(Panzer)
Dours, 1869 An, NT (:An. curta
sqturmmulosa
CockProvancher, 1895 n. syn.) :An. melanops
erell, 1926 fas curta ssp.l n. syn. | =An. ensenadmsis
Cockerell, 1941 las curtassp.l n. syn.) (489' 49O,
494, 496)
stanfordianaCockerell, see bomboidesKirby
stantoni(Cockerell), 1911 fas zonatassp.l Am (An), O
(511)
stefaniiPérez, seepruinosaSmith
strauchiFedtschenko, 1875 An, P (462)
Fedtschenko, see cinnascemLepeletier
stschurooskyi
(Kohl), 1905 An (Po), P (496)
subaequ"a
subcarinalaBenoist, see balneorumLepeletier
(Lepeletier), 1B4l Am (An), O (=An.
subcoerulea
Radoszkowsky, 1BB2n. syn.i :An.
lucknoaiensrs
rothneyiCameron, 1897 n. syn.; --An.amolita
Cockerell, 1911 n. syn.) (502, 5O4)
Yasumatsu, seedulcifna (Cockerell)
subflaaescens
Kirby, see quadrimaculata(Panzer)
subglobosa
Provancher, seeterminalisCresson
subglobulosa
subignauaCockerell, seepacificaCresson
subinsularùStrand [as insularisssp.] Am (An), listed by
Cockerell, 1919, but original description not found
(497, 5t3)
submicans
Gussakovsky, 1935 An, P (496)
(Cockerell), 1925 Am (An), E (513)
subrussata
subsaltni(Rayment), 1947 Am (At), A (511)
Morawitz, 1894 An, P (496)
subserricornis
subtarsataCockerell, see califtrnica Cresson
subtnraneaGermar, 7826 An, P (: Megilla chrysopoda
Illiger, 1806 n. syn., nomennudum; :Megilla
decrepitallliger, 1806 n. syn. , nomennudum; :An.
laticincta
nigrocinctaLepeletier,1841 n. syn.', :At
Dours, 1869 n. syn.) :An. procuaCosta, 1BB3n.
syn. i :An. flauescemGribodo, 1893 fas nigrocincta
ssp.] n. syn.) (457, 460, 462, 557)
(Strand)
subtenanrzGistel, see dalmatiensæ
subtonida(Cockerell), 19+6 Am (An), E (515)
sumatranaLieftinck, 1956 Am, O (513)
supnansWalker, lBTl An, P (472, 474, 476)
supnbaFriese, seeepichariftrmisGribodo
suprafmea(Cockerell), 1946 Am (A.), E (504)
sutepensis
Cockerell, 1929 Ha, O
suuorzeaiMorawitz, IBBB An, P (462)
sybilae(Rayment), 1944 Am (At), A, publication date
is not l9+2 (512)
syriacaFriese, 1922 An, P (476)
syringaeCockerell, see taminalis Cresson
tadzhicaPopov, 1948 Ha, P
tainanaStrand, seecalceifera(Cockerell)
tainanicola(Strand), 1913 Ha (At), O (: Ha. maiella
Lieftinck, 197+ las tainanicolassp.] n. syn.)
talaris(Pérez), 1895 Am (An), P (504)
tarsalisPriesner, 1957 An, P (492)
tarsata(Spinola), 1B3BHa (Tbtralonia),P (: Habrophora
:An. passuini Sichel,
fsic! ezonatuSmith, 1854;
1856) (447)
tarsataDours, seecalifurnicaCresson (557)
Fedtschenko, lB75 An, P (476)
tarsidens
taureaSay, 1837 (An) :Melitoma (Anthophoridae;
Emphorini) ( =An. fulaifrons Smith, 1854)
tauricaFriese, seepodagraLepeletier
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AND PHvrocENY
aestitellaBrooks An, R n. name for An. aestita
Morawitz, IBBB, Horae Soc. Entomol. Ross.
22:232, not Smith, 1854 (474, 476)
uestitula(Cockerell), 1936 Am (An), E (:An. uestibula
[sic] Cockerell, 1946) (515)
aetulaEversmann, see senilisEversmann
aetulaKrg, 1845 Un (Megilla), P (450)
aictoriensrs
(Rayment), 1951 Arn (Asaropoda),A (515)
uidua(Khg), 1845 An (Megilla), P (-An. boops
Alfken, 1926, [dark form]i :An. fumipennr Alfken, 1926 n. syn. [dark form]) (473, 474, 476)
aierecki(Cockerell), 1909 Ha (Emphoropsis)n. comb.,
N
uiereckiCockerell, see centrifurmrsCresson
aigilans(Smith), 1861 Am (An), O (513)
uigilansSmith, seecustos(Dalla Torre)
aillosaHerrich-Schaeffer, seeplagiata (Illiger)
aillosulaSmith, seeplumipes(PaLlas)(460)
uiolacea
(Lepeletier), 1841 Am (An), O (:Megilla
llliger, 1806 nomennudum, n. syn.;
albopunctata
:An. anthracinaGribodo, 1893 [as uiolacea
ssp.] n.
syn.) (450, 512, 513')
uirgoGribodo, 1893 (A") -- Centrismuralis Burmeister,
lB76 (Anthophoridae; Centridini)
uiaida(Smith), lB79 Am (An), E (506)
aiuidulaStrand, seecincta(Fabricius)
Dours, 1869 Un (An), T (450)
uolucellaeforzes
aulpina Panzer, seequadrimnculata(Panzer) (486' 489)
wadicolaAlfken, 1935 An, P (486)
walkeri(Cockerell), 1905 Am (An), O (--An. darwini
Cockerell, 1910 n. syn.) (510, 511)
wallacei(Cockerell), 1907 Am (An), A (511)
wallaciella Rayment, see perpulchra(Rayment)
utalshiiCresson, 1869 An, N (466, 468, 469)
waltoni Cockerell, 1910 las aulpiza ssp.] Att, O (486)
wartmanmFriese, 1905 An, E (492)
washingtoniCockerell, seecrotcàiiCresson
watsoniCockerell, seeporteraeCockerell
Ja
'+,,
OF THE ANTHOPHORINB
BBBS
575
wegeliniFriese, l9l+ An, P (486,489)
wegeneriFriese, see alternazs(Klug)
whiteheadi
(Cockerell), 1910 las zonatassp.] Am (An),
o (511)
(Rayment), 1947 Am (An), A (511)
whiteleyella
wickwari Bingham, 1908 (Po) : Eucara(Atthophoridae; Eucerini) t. comb.
Kirby
willingi Cockerell, see bomboides
wtureBrooks An, P, n. name for Anthomegillasinensis
Alfken, 1937, EntoWu, 1982, not Ha. sinensas
mology and Phytopathology 5:404-405 or Clisodon
Wu, 1982, Insects of Xizang, vol. II:419
sinensis
(482)
xanthochlora
Cockerell, 1923 An, T (496)
Cockerell, 1932 An, E (: Ar. uernali
xanthostoma
Cockerell, 1935 n. syn. The female holotype of
uernayi(Transvaal) has the metasoma of a speciesof
Tbtraloniaglued onto it; the same metasoma
described in the original descriptioni : An. heterossp.] n. syn.) (492)
PodoCockerell, 19+6 las aernaye
ssp.] Am
xerophila(Cockerell), 1911 [as quadrifasciata
(Ar), o (505)
(Wu), 1985 An (Clisodon),P (489)
xinjiangensas
xizangensls
Wu, 1979 Ha, P
Wu, 1983 Ha, P
)unnanenses
Wu, 1983 Am, P (513)
lunnanemes
zamoranellaCockerell, l9+9 An, T (496)
Priesner,
zanoniGribodo , 1924 An, P (= An. borgensis
1957 fas zanonessp.] n. syn.) (486)
Wu, see imitatrix Lieftinck
zhe;jiangenses
zimini Gussakovsky, 1935 An, P (496)
zinniae(Cockerell), 1932 Am (An), E (504)
zombanaCockerell, 1910 An, E (492)
zonata(Linnaeus), 1758 Arn (Apis), O (:An. nigritarsis
Dover, 192+ n. syn. i :An. doaeriCockerell, 1931,
n. name for An. nigritarsisDover, 192+, not Friese,
1 9 0 5n . s y n . ) ( 5 0 9 - 5 1 1 )
zonatulaSmith, 1854 Ha (Habrophora[sic]), P
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