, 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