Table Of Content,
A Review of Perdita, Subgenus Macrotera
(Hymenoptera: Andrenidae)
Roy R. Snelling1 and Bryan H. Danforth2
ABSTRACT. The taxonomy of the species comprising Perdita subgenus Macrotera is reviewed; these
beesare oligolegeson Opuntia (Cactaceae).A keyto thespecies isgiven; two newspeciesare described
from Mexico: P. pipiyolirt fromJalisco and P. nahua from Michoacan; new synonymy is proposed: P.
secundaCockerell,1904,andP.texanaablusaTimberlake,1958=P.texana(Cresson,1878).Appropriate
morphological features are illustrated forall species and the known distributions are mapped.
INTRODUCTION MATERIALS AND METHODS
Macrotera althoughoriginallydescribedasagenus INSTITUTIONAL COLLECTIONS
,
by F. Smith (1853), has long been treated as a sub- Duringthecourseofthisreviewwehavestudiedmaterial
genus ofPerdita F. Smith (1853) (e.g., Timberlake, fromthe followingcollections: Bee Biologyand System-
1954). The few species occur in Mexico and in atics Laboratory, USDA, Logan, Utah (BBSL); Natural
Texas and New Mexico in the United States. All HistoryMuseum,London(BMNH);CaliforniaAcademy
are presumed oligoleges on Opuntia and possibly ofSciences(CAS);CentralTexas MelittologicalInstitute,
otherCactaceae. The specieswere treated byTim- Austin, Texas (CTMI); Natural History Museum of Los
berlake (1954, 1958). With the discovery of two AngelesCounty(LACM);UnitedStatesNationalMuseum
additionalMexicanspecies,wedeemitappropriate of Natural History (USNM); Universidad Autonoma de
to review the previously described species in ad- Mexico(UNAM);UniversityofCalifornia,Berkeley(UCB)
dition to describing the new ones. and University ofCalifornia, Riverside (UCR); University
ofKansas (UKAN).
Species ofMacroteraare relativelylarge forPer-
dita (body length up to 10 mm); males have heads
thatare conspicuously broaderthan long, with the TERMINOLOGY
inner eye margins somewhat divergent below. Al-
though the head is broader than long in females, aInmognengerable,etshyestetmeartmiisntso.loFgoyreampdleosycreidpthieorneoifs psutnacntduarred
thedisparityislessextremethaninthemales.Other sizes and spacing, see Snelling (1985). The following ab-
featurescharacteristicofMacroterainclude:glossa breviationsare used in the descriptions.
of both sexes 2-3 times length ofprementum and HL—
extending between metacoxae; second segment of headlength:infrontalview,themidlinedistance
betweenthelowerclypeal marginand thesummitofthe
maxillary palpus longer than first or any of the
following segments; both sexes with well devel- verHtexW.—head width: in frontal view, the maximum dis-
oped facial foveae; marginal cell offorewinglarge, tance ac—ross the eyes.
atleastsubequaltofirstsubmarginal;stigmaslender IOD interocellardistance:the minimumdistancebe-
andtapering;metatibiaand basitarsusofmalewith tween —the posteriorocelli.
long,flattened,scale-likehairs; both sexeswith ba- LID lowerinteroculardistance:thedistancebetween
sitibial plate on metatibia; tibial spurs stout and the inner eye margins at the level of the lateral angle of
aobfrmupatlleyfhloatotkeeneddaatnadpberxoiandebrotthhasnexmees;somseotmaas.oma theOcDly—peuosc.ellus diameter: the transverse diameterofthe
anteriorocellus.
OOD—ocelloculardistance:theminimumdistancebe-
1. Entomology Section, Natural History Museum of tween eitherposteriorocellusand the adjacent innereye
gLeolsesA,nCgaelliefsorCnoiuan9t0y0,079.00 ExpositionBoulevard,LosAn- marOgiVnD.—ocellovertexal distance: with head in frontal
2. Department of Entomology, Smithsonian Institu- view, the minimum distance between either posterior
tion NHB 105, Washington, D.C. 20560. ocellus and the dorsal margin of the vertex.
Contributionsin Science,Number436, pp. 1-12
Natural History Museum of Los Angeles County, 1992
Figures 1-5. Frontalviewsofmale heads;(1)P. texana,(2)P. bicolor,(3)P. nahua,(4) P. pipiyolin,(5)P. sinaloana.
TL—total length: the sum of HL + length of meso- as key characters, although they are illustrated for each
soma (from anterior margin of mesoscutum to a point species (Figs. 6-11).
directlyabove metasomalattachment) + lengthofmeta-
soma(frompointofattachmentofmesosomatofarthest KEY TO SPECIES OF MACROTERA
extremity).
—
UID upperinteroculardistance:with head in frontal la. Male; antenna with 13 segments 2
view, the minimum distance between the inner eye mar- b. Female; antenna with 12 segments 7
ginsW.L—winglength: minimum distance from margin of 2a. Lowerfacewithclypeusandlateralfacemarks
tegula to extremity ofwing tip. yellowish (Figs. 2-5) 3
The last visible metasomal sternum of the females is b. Lower face concolorous with remainder of
characteristicallydifferentforeachspecies.Inordertobe face, distinctly not yellowish (Fig. 1) .... 6
properly examined, it must first be removed and cleared 3a. Facial fovea long and narrow (but broader
in KOFI. For this reason, we have not used these sterna above), about parallel with inner eye margin
2 Contributions in Science, Number 436 Sneliing and Danforth: Perdita, Subgenus Macrotera
Figures 6-11. Ventral views offemale S6; (6) P. texana, (7) P. bicolor, (8) P. nahua, (9) P. crassa, (10) P. pipiyolirt,
(11) P. sinaloana. Scale bar = 0.5 mm.
and upper end separated from eye margin by scutellum mostly separated by apuncture di-
0.25 timeslength offoveaorless;mesoscutal ameter or more bicolor (F. Smith)
interspaces dull and conspicuously tessellate b. Supraclypeal area subcontiguously punctate
4 and without impunctate median zone; punc-
b. Facial fovea short and relatively broad, turesofmesoscutumandscutellumuniformly
strongly oblique to inner eye margin, upper subcontiguous 5
endseparatedfrom eyemargin byabout0.50 5a. Extreme base of propodeum roughened but
times length of fovea (Fig. 5); mesoscutal in- without short longitudinal rugulae; aedeagus
terspacesdistinctlyshinyandveryweaklytes- without ventral hook-like process (Fig. 21)
sellate over most of disc pipiyolin, new species
sinaloana Timberlake b. Extreme base of propodeum roughened and
4a. Supraclypeal area with distinct impunctate withshortlongitudinalrugulae;aedeaguswith
zone along midline; punctures in postero- ventral hook-like process (Fig. 22)
medianareaofmesoscutumandofmiddle of nahua, new species
Contributions in Science,Number436 Snellingand Danforth: Perdita, SubgenusMacrotera 3
Figures 12-17. Ventral views ofmale S7and S8; (12) P. texana, (13) P. crassa, (14) P. pipiyolin,(15) P. bicolor, (16)
P. sinaloana, (17) P. nahua. Scale bar = 0.5 mm.
6a. Front of head, including clypeus and para- clypeus; subgenital plate narrow, apex nar-
oculararea, with conspicuous hairs, many of rowly rounded (Fig. 12) . . texana (Cresson)
them distinctly barbulate; subgenital plate 7a. Scopalhairsflattenedandconspicuouslywavy
(metasomal sternum 7) broad, apex subtrun- or crinkled 8
cate (Fig. 13) crassa Timberlake b. Scopal hairs simple, i.e., long, somewhat flat-
b. Frons with short, simple hairs, clypeus and tened and evenly tapering to acute, often
paraocular area virtually without hairs other curved apices 9
than some exceedinglyshort, simple hairs on 8a. Extreme base of propodeum roughened but
4 Contributions in Science,Number 436 Snellingand Danforth: Perdita,Subgenus Macrotera
Figures 18-20. Dorsal,ventral,lateralviewsofmalegenitalcapsuleanddorsalandlateralviewsoftheaedeagus;(18)
P. texana, (19) P. crassa, (20) P. bicolor. Scale bar = 0.25 mm.
withoutshort longitudinalrugae; side ofpro- and inconspicuous hairs near lateral margin
mm
podeal discwith some hairs at least0.07 11
long pipiyolin, new species 10a. Punctures ofclypeal disc subcontiguous, me-
b. Extreme base of propodeum roughened and dian impunctate line absent or narrow and
with several short longitudinal rugae on each incomplete;mesoscutumdullandsharplytes-
sideofmiddle;hairsonsideofpropodealdisc sellate between punctures
mm
uniformlyveryshort,noneover0.05 long bicolor (F. Smith)
nahua, new species b. Punctures of clypeal disc very irregularly
9a. Propodeal disc with abundant long, conspic- spaced,medianimpunctatelinebroadandof-
uously plumose hairs 10 ten poorly defined, disc sometimes with only
b. Propodealdiscbareorwithsparse,veryshort scattered punctures; mesoscutum shiny and
Contributions in Science,Number436 Snelling and Danforth: Perdita, Subgenus Macrotera 5
:
Figures21-23. Dorsal,ventral,lateralviewsofmalegenitalcapsuleanddorsalandlateralviewsoftheaedeagus;(21)
P. pipiyolin, (22) P. nahua, (23) P. sinaloana. Scale bar = 0.25 mm.
weakly tessellate between punctures minute to fine punctures that are mostly sep-
crassa Timberlake arated by about puncture diameter or less;
11a. Face between lateral ocelli and upper end of scutellum slightly shiny between mostly sub-
facial foveae slightly shiny, distinctly tessel- contiguous punctures
late between variably spaced, minute punc- sinaloana Timberlake
tures,someinterspacesseveraltimespuncture
diameter; middle of scutellum shiny, punc- Perdita Macrotera bicolor (F. Smith)
ttuurreesdoifatmeentesreparated by sevteeraxlantaim(eCsrepsusnocn-) Figu(res 2, 7, 15,)20, 24, 31, 35
b. Face between lateral ocelli and upper end of Macrotera bicolor F. Smith, 1853:130; 9.
facialfoveaemoderatelyshinybetweenmixed Perdita bicolor Cockerell, 1905:321.
6 Contributions in Science, Number 436 Snellingand Danforth: Perdita, Subgenus Macrotera
28 29
. .
Figures 24-29. Dorsal views ofmale T7; (24) P. bicolor, (25) P. sinaloana, (26) P. pipiyolin, (27) P. texana, (28, 29)
P. nahua (showingextent ofintra-specific variation).
Perdita Macrotera bicolor: Timberlake, 1954:352, Perdita Macrotera crassa Timberlake
( ) ( )
353; $ <5. Timberlake, 1958:375, 376; 5 2. Figures 9, 13, 19, 35
yelTlhoewmfaalcee omfarPk.sb.icItoldoirffiesrosnferoofmfPo.ursisnpeaclioeasnwaibtyh Per3d7i6t-a37{M7a;cr2o6t.era) crassa Timberlake, 1958:375,
thelong,narrowfacialfoveaethatareparallelwith
the inner eye margins (Fig. 2). From P. pipiyolin Males of P. crassa are easily recognized by the
and P. nahua, it differs by the distinctly sparser lack ofyellowish face marks, the denselypunctate
punctation ofthe supraclypeal area and the center supraclypeal area, and the broad, apically subtrun-
of the mesoscutal disc. Both P. crassa and P. tex- cate subgenital plate (Fig. 13). The female shares
ana lack yellow face marks. with P. bicolorthepresenceofnumerouslongplu-
The presence of numerous long, plumose hairs mose hairs on the propodeal disc, but differs from
onthepropodealdiscwillseparatethefemalefrom thatspeciesintheveryirregularspacingoftheclyp-
all except P. crassa. From that species P. bicolor eal punctures and the smooth, shiny interspaces
differs in the evenly spaced clypeal punctation, but between mesoscutal punctures.
withadistinctimpunctatemedianline,andthedull, Although presently known only from Texas, P.
densely tessellate mesoscutal interspaces. crassaprobablywill be found to occurin Mexico,
Althoughmostfemaleshavemetasomalterga2- at least in the states ofNuevo Leon and Coahuila.
5 or 3-5 largely reddish, we have seen a few with
the entire metasomal dorsum blackish brown. Fe- Perdita (Macrotera) nahua, new species
males of P. bicolor have the inner margins of the Figures 3, 8, 17, 22, 28, 29, 33, 35
forecoxae (in ventral view) straight and the inner DIAGNOSIS
apical angle abrupt (Fig. 31).
Thisspeciesisknownonlyfromtheeasternhigh- FEMALE. SeparablefromallMacroteraspecies
lands ofMexicowhere it has been collected in the exceptP. pipiyolin by the distinctlywavy or“crin-
States of Hidalgo and Puebla. kled” scopal hairs; differs from P. pipiyolin by the
Contributionsin Science,Number436 Snelling and Danforth: Perdita, Subgenus Macrotera 7
pro-pleuron
pro-sternum
31.
Figures30-34. 30-32. Ventralviews offemale forecoxae; (30) P. pipiyolin, (31) P. bicolor, (32) P. texana. Scale bar
= 0.5 mm. 33, 34. Forewingvenation; (33) P. nahua, (34) P. texana.
distinctlyrugulosepropodealbaseandtheveryshort view. Facial fovea linear, parallel to innereye mar-
hairs laterad on the propodeal disc; additionally, gin and distinctly broader above. IOD about 1.6
the metasoma is reddish to largely brown. times OD; OOD about 2.9 times OD; OVD about
MALE. Separablefrom P. crassaand P. texana equal to OD. Antennal scape extending almost to
by the yellow clypeus and lateral face marks; sep- levelofposteriorocelli;firstflagellarsegmentabout
arable from P. sinaloana by the long facial fovea as long as broad and no longer than second seg-
that parallels the inner eye margin; separable from ment. Clypeus shiny, with or without narrow im-
P. bicolorbythedenselypunctatesupraclypealarea punctate median line, but most of disc with sub-
and the shiny, subcontiguously punctate mesoscu- contiguous moderate punctures. Supraclypeal area
tal disc; separable from P. pipiyolin by the sharply duller,puncturesfineand contiguoustosubcontig-
rugulosepropodealbaseandtheprominentventral uous, sometimes sparse along midline; lowerpara-
process of the penis valve (Fig. 22). ocular area shiny between sparse punctures finer
DESCRIPTION than on clypeus, becoming duller above between
minute subcontiguous punctures. Frons dull and
HW
FEMALE. Measurements (mm): Holotype sharply tessellate between minute subcontiguous
2.42; HL 1.84; WL 5.2; TL 8.8. Paratypes: HW punctures. Vertex dull and tessellate, ocellocular
2.23-2.54;HL 1.77-2.00;WL4.8-5.5;TL8.1-9.2. areawithminutesubcontiguouspuncturesthatgrade
Headabout 1.3 timesas broadaslong; innereye into coarser, more separatedpunctureson vertexal
margins slightly divergent below, LID about 1.03 summit.
timesUID;vertexalmarginevenlyconvexinfrontal Mesoscutum shiny over most ofdisc, becoming
8 Contributions in Science,Number 436 Snelling and Danforth: Perdita, Subgenus Macrotcra
Figure 35. Distribution map of P. Macrotera species.
( )
lightlytessellatetoward margins,puncturesminute cephalad; lateral margins of propodeal disc with
mm
to fine, dense, even along mid-line. Scutellum sim- inconspicuous hairs less than 0.05 long.
ilar, punctures not noticeably sparser in middle. Colorhead and most ofmesosoma dark brown-
Metanotum dull, densely tessellate, punctures mi- ish, mesoscutum darker, antenna and legs paler;
nuteandsubcontiguoustocontiguous.Mesepister- metasoma reddish to largely reddish brown, seg-
num slightly shiny and roughened between dense ment margins distinctly reddish. Tegula yellowish.
fine punctures. Metepisternum shinier, with scat- Wings clearlight brownish, veins mostlyyellowish
teredfinepunctures.Propodeummoderatelyshiny, brown, subcosta and stigma darker.
HW
basal margin narrowly roughened and with short MALE. Measurements (mm): 2.48-2.97;
longitudinal rugulae; lateral portions of disc with HL 1.44-2.06; WL 5.0-5.6; TL 7.3-8.4.
finepiligerouspunctures. Innermargins ofprocox- Head(Fig. 3) broaderthanmesosoma,about 1.5
ae nearly straight, innerapical angle produced and times as broad as long; inner eye margins weakly
acute (similarto Fig. 30). Tibial spurs stout, apices divergent below, LID about 1.05 times UID; ver-
abruptlyhooked;hairsofmetatibialscopalongand texal margin nearly flat across middle one-half in
distinctly wavy or “crinkled”. anteriorview. Facial foveae about as in female but
Metasoma moderately shiny, terga transversely marginslessdefined.IODabout2.0timesOD;OOD
lineolate, sculpture becoming coarser on succeed- 4.8-6.0 times OD; OVD 2.5-2.7times OD. Anten-
ing segments, translucent apical margins broad on nalscapeextendingtoaboutlevelofposteriorocel-
terga2-4; puncturesultraminuteto minute, denser li; first flagellar segment about as long as wide and
onsuccessivesegmentsandlaterad on each. Pygid- slightly longer than second segment. Facial sculp-
ial plate broadly rounded or subtruncate at apex, tureaboutasinfemalebutclypealpuncturesslight-
margins slightly reflexed, disc granulopunctate. ly finer, mostly subcontiguous and median im-
Sternasimilartotergabutdiscsmoredenselypunc- punctate line very narrow or absent; supraclypeal
tate; S6 as in Figure 8. areawithmedianimpunctatelinenarroworabsent.
Pilosity normal for Macrotera mesoscutal hairs Mesosoma as described for female, but punc-
;
suberect to erect, mostly very short but with scat- tures of disc of mesoscutellum uniformly subcon-
tered longer hairs; hairs across middle one-half of tiguous in center. Femora, especially metafemur,
metanotum very short, subappressed, and directed robust; tibial spurs stout, apices abruptly hooked.
Contributions in Science,Number436 Snelling and Danforth: Perdita, Subgenus Macrotera 9
,
Metasoma, exceptusual sexual differences, as in Perdita (Macrotera) pipiyolin,
female;pygidialplate(Fig.28,29)broad,withslightly new species
reflexed margins, apex subtruncate, disc distinctly
Figures 4, 10, 14, 21, 26, 30, 35
coarsely rugose. Subgenital plate and genitalia as
illustrated (Figs. 17, 22).
DIAGNOSIS
Pilosityabout as in female exceptscopa lacking,
meso-andmetatibiaewithwidelyspaced,long,flat, FEMALE. Separable from all other species of
scale-like hairs. Macrotera except P. nahua by the wavy or “crin-
Coloras infemale except: mandible,exceptred- kled” hairs ofthe metatibialscopa; separablefrom
dish apex; labrum; clypeus, except small to large P. nahua bytheweaklyroughenedpropodeal base
mediobasal brown spot; lower paraocular area, and the longerhairs laterad on thepropodeal disc;
sometimes including stripe along inner margin of P. pipiyolin and P. nahua are both additionally
eye nearly to summit, all pale yellowish. Protibia separable from all other species except P. texana
yellowish red, metasoma bright reddish. by the shiny, denselypunctate mesoscutal disc and
from P. texana by the presence of conspicuous,
TYPE MATERIAL though short, hairs laterad on the disc of the pro-
podeum.
Holotypefemale:Ziculcan,Michoacan,MEXICO, MALE. SeparablefromP. crassaand P. texana
6p86a,Jruasltayypm1ee9s8di8ant(LaGA.aCsRMoh,dolroUitgNyupAeezM.,GH.)oU.lSoPtNaryMap.teypiens:U5NA9$M,;4 abryabtlheefyreollmowP.csliypneaulsoaannadblyatterhaellfoancge fmaacrikals;fosveepa-
that parallels the inner eye margin; separable from
ETYMOLOGY P. bicolorbythedenselypunctatesupraclypealarea
and the shiny, subcontiguously punctate mesoscu-
This species is named for the Nahua people who tal disc; separable from P. nahua by the weakly
inhabited central Mexico prior to the Spanish in- roughenedpropodealbaseandthelackofaventral
vasion. The name is a noun in apposition. hook-like process on the aedeagus (Fig. 21).
DISCUSSION DESCRIPTION
In most respects both sexes of P. nahua are very FEMALE. Measurements(mm):Holotype,HW
similartoP.pipiyolin.Theprominent,acuteventral 2.55; HL 2.10; WL 5.9; TL 10.1. Paratypes: HW
processofthemaleaedeagus,however,willpermit 2.39-2.65; HL 1.94-2.13; WL 52-6.0; TL 9.3-
readyseparationofthisspeciesfromallotherMac- 10.2.
rotera males. Head about 1.2 times as wide as long; inner eye
The females ofP. nahua togetherwith those of margins slightly divergent below, LID about 1.04
P.pipiyolin,haveaverycharacteristicscopainwhich timesUID;vertexalmarginevenlyconvexinfrontal
the scopal hairs have very attenuated apices that view. Facial fovea linear, parallel to innereye mar-
are distinctly wavy or crinkled; in all other species gin,distinctlybroaderat upperend. IODabout 1.6
the scopal hairs, although often attenuated, are times OD; OOD about 2.9 times OD; OVD about
curled apically. These two species are also char- equal to OD. Antennal scape extending to level of
acterized by their relatively densely punctate clyp- anterior ocellus; first flagellar segment distinctly
eus and mesoscutum. The presence of numerous longerthanbroadandabouttwiceaslongassecond
short, longitudinal to oblique rugules across the segment. Clypeus shiny, with narrow impunctate
base of the propodeum will separate females from median line, but most of disc with subcontiguous
P. nahua from those of P. pipiyolin, as will the moderate punctures. Supraclypeal area duller,
reduced and very short pilosity at the side of the punctures fine and contiguous to subcontiguous,
propodeal disc. sometimes sparse along mid-line; lowerparaocular
Females ofthis species and ofP. pipiyolin often shiny between sparse punctures finerthan on clyp-
M
havevein Im-cu oftheforewingjoining beyond eus, becoming duller above between minute sub-
Rs(Fig.33);inthefourremainingspeciesofMacro- contiguous punctures. Frons dull and sharply tes-
tera lm-cu and Rs are interstitial (Fig. 34). Little sellate between minute subcontiguous punctures.
reliancecanbeplacedonthischaracterofvenation Vertex dull and tessellate, ocellocular area with
because some specimens of P. pipiyolin have the minute subcontiguous punctures that grade into
two veins interstitial. coarser,more separatedpunctures ofvertexalsum-
Of the paratype females of P. nahua, one has mit.
the entire metasoma bright reddish. In two speci- Mesoscutum shiny over most of disc, becoming
mens the terga are largely reddish medially but ex- lightlytessellate toward margins,puncturesminute
tensivelybrownish at the sides. The two remaining to fine, dense, even along mid-line. Scutellum sim-
females have the metasomal terga mostly dark but ilar, but punctures noticeably sparser in middle.
with dull reddish margins on segments 1-5. Metanotumdull,denselytessellate,puncturesmin-
Although no floral data are provided with these ute and subcontiguous to contiguous. Mesepister-
specimens, the pollen in the scopae appears to be num slightly shiny and rougher between dense fine
that of a cactus species, probably Opuntia. punctures. Metepisternum shinier, with scattered
10 Contributions in Science, Number436 Snelling and Danforth: Perdita, Subgenus Macrotera
