Scientific Papers

58 Pages·2008·4.3 MB·English

QL
568
.S7
M35
1999
Papers
Scientific
Natural Museum
History
The Kansas
University of
10 December 1999 Number 14:1-55
Phylogenetic Relationships and Classification of the
Major Lineages of Apoidea (Hymenoptera),
with Emphasis on the Crabronid Wasps1
By
Gabriel A. R. Melo2
Division ofEntomology, NaturalHistoryMuseum
The UniversityofKansas, Lawrence, Kansas 66045, USA
CONTENTS
ABSTRACT 2
INTRODUCTION
2
Classification andPhylogenetic RelationshipswithintheApoidea 2
ThePresentStudy 4
Acknowledgments 4
MATERIALAND METHODS 4
SelectionofRepresentativeTaxa 4
DissectionofAdultSpecimens 6
CharacterSelection and Delimitation 6
Terminology 8
Larval andBehavioralCharacters 8
DataAnalysis 8
CHARACTERSANDCODESFORTHEIRSTATES II
RESULTS 23
DISCUSSION 23
ChoiceofAnalyticalMethod 23
Apoideaanditsbasalclades 26
Heterogynaidae 34
Ampulicidae 34
'ContributionNumber3234fromtheSnowEntomologicalDivision.NaturalHistoryMuseum,and DepartmentofEntomology,TheUniversityof
Kansas.
-Presentaddress:DepartamentodeBiologia,FFCLRP.UniversidadedeSaoPaulo.Av.Bandeirantes3900.14040-901,RibeiraoPreto,SP,Brazil.
©NaturalHistoryMuseum,TheUniversityofKansas ISSNNo. 1094-0782

QL
568
.S7
M3 5
1999
Papers
Scientific
Natural Museum
History
The Kansas
University of
10 December 1999 Number 14:1-55
Phylogenetic Relationships and Classification of the
Major Lineages of Apoidea (Hymenoptera),
with Emphasis on the Crabronid Wasps1
By
Gabriel A. R. Melo2
Division ofEntomology, NaturalHistoryMuseum
The University ofKansas, Lawrence, Kansas 66045, USA
CONTENTS
ABSTRACT
2
INTRODUCTION 2
ClassificationandPhylogenetic RelationshipswithintheApoidea 2
ThePresentStudy 4
Acknowledgments 4
MATERIALAND METHODS 4
SelectionofRepresentativeTaxa 4
Dissection ofAdultSpecimens 6
CharacterSelection and Delimitation 6
Terminology...^ 8
LarvalandBehavioralCharacters 8
DataAnalysis 8
CHARACTERSANDCODESFORTHEIRSTATES 11
RESULTS 23
DISCUSSION 23
ChoiceofAnalyticalMethod 23
Apoideaanditsbasalclades 26
Heterogynaidae 34
Ampulicidae 34
'ContributionNumber3234fromtheSnowEntomologicalDivision.NaturalHistoryMuseum,and DepartmentofEntomology,TheUniversityof
Kansas.
;Presentaddress:DepartamentodeBiologia,FFCLRP.UniversidadedeSaoPaulo.Av. Bandeirantes3900.14040-901,RibeiraoPreto,SP,Brazil.
©NaturalHistoryMuseum,TheUniversityofKansas ISSNNo. 1094-0782
2 Scientific Papers, Natural History Museum, The University of Kansas
sphecidae (sensustr1cto) + [apidae (sensu lato) + crabronidae] 35
Sphecidae (sensustricto) 36
Apidae (sensulato) + Crabronidae 36
Apic/.e (sensu lato) 38
CRABRON.DAE 38
LITERATURECITED 43
APPENDIX(Figures 10-82) /ARD 46
ABSTRACT The superfamily Apoidea is one of the three major groups of Hymenoptera Aculeata,
beingcomposed ofthesphecoid wasps, thebees,and theHeterogynaidae, a small and poorlyknown
groupofwasps.Thephylogeneticrelationshipsamongthemajorlineagesofapoidswereinvestigated
using 130 charactersfrom the morphology ofthe adult insects, six from larval morphology, and three
characters fromadultbehavior. These 139 characterswereanalyzed underthreeparsimony methods:
equal weighting, implied weighting, and successive weighting. Different phylogenetic hypotheses
wereproduced by each method (55 treesunderequalweighting, four trees underimplied weighting,
and one tree under successive weighting, for analyses including all 54 exemplar taxa). The results
from implied weighting are favored over those of the other two methods and are used to propose a
higherlevelclassificationfortheApoidea.HeterogynaidaeandAmpulicidaeconstitutethemostbasal
apoid clades; however, the position ofHeterogynaidae remains ambiguous: in three implied weight-
ing-trees, it comes out as the sister group ofAmpulicidae and in the fourth as the sister group of the
remaining Apoidea, excluding Ampulicidae. The remaining families recognized and their relation-
ships are: [Sphecidae (sensu stricto) + [Apidae (sensu lato) + Crabronidae]]. Only five subfamilies of
Crabronidae are recognized: Astatinae, with the tribes Astatini, Eremiasphecini and Ammoplanini;
Bembicinae; Crabroninae (including the genera Dinettes, Laphyragogus, Mellinus and Xenosphex);
Pemphredoninae, with the tribes Psenini (including the genera Odontosphex and Entomosericus) and
Pemphredonini; and the Philanthinae.
INTRODUCTION
The superfamily Apoidea is one of the three major since1974andnewstudies,inparticularCarpenter's(1986)
cladesoftheAculeata Hymenoptera (Brothers1975,Gauld investigation on the Chrysidoidea (= Brothers'
and Bolton1988, BrothersandCarpenter1993).Apeculiar Bethyloidea). Their results largely support the phyloge-
difference exhibited by aculeate females in relation to the netic patternsfound in thesetwo previous works, includ-
remaining Hymenoptera is their modified ovipositor, no ing the three major lineages of Brothers (1975). The now
longer used for laying eggs, but only as a sting to inject widely accepted superfamilial classification for the
venom into the host or prey, as well as into potential at- Aculeata proposed by Brothers (1975) isbased on therec-
tackers (defensive function). As in many groups of para- ognitionofthesethreelineages,i.e.Chrysidoidea,Apoidea
sitic Hymenoptera, females of most aculeate lineages be- and Vespoidea. Chrysidoidea, the basal clade of the
haveas idiobiontparasitoids, i.e., upon finding a suitable Aculeata, contains small wasps most ofwhich behave as
host, usually concealed in protected places, the female parasitoids or sometimes as cleptoparasites. Vespoidea is
waspparalyzesitwithitsvenomousstingand laysanegg alargeassemblageofverydistinctaculeatelineages;most
on the host surface (Gauld and Bolton 1988). However, are parasitoids, but well-known groups like ants and so-
several lineages of Aculeata departed from this ancestral rial paper wasps arealso included,
modeoflifeand haveevolved complexnestingand social
Classification andPhylogenetic Relationships
behaviorstoadegreenotparalleledbyanyothergroupof
within theApoidea
insects,excepttermites.Severalaspectsofthebiologyand
evolution of the aculeate wasps are presented and dis- The Apoidea is composed of the sphecoid wasps
cussed by Evans and West-Eberhard (1970), Iwata (1976), [Sphecidaesensu Bohartand Menke (1976)], thebeesand
Gauld and Bolton (1988) and Hanson and Gauld (1995). thegenusHeterogynaNagy(thegeneraDaycatincaandDaya
The phylogenyof the majoraculeate lineageswas re- are treated here as synonyms of Heterogyna; see below), a
centlyinvestigatedby BrothersandCarpenter(1993).This small and poorly known group ofwasps placed in a fam-
comprehensivestudymostlyreevaluated Brothers' (1975) ilyofitsown (Brothers and Carpenter 1993). Mostapoids
work, incorporating new characters systems proposed showderived lifehistory traitscompared to theancestral
Major Lineages ofApoidea
aculeate parasitoid behavior, with the females exhibiting for Brothers's Sphecoidea. Brothers (1975) also proposed
a high degree of parental care. Their host, or better, the an informal division of theApoidea into two groups, the
immature'sprovisionsarenowtransportedandconcealed Spheciformes (= Sphecidaesensu Bohartand Menke) and
ina pre-existingorespeciallybuiltcavity.Constructionof theApiformes (bees).
a nest before prey capture apparently evolved only once Heterogx/nawithitsreducedsizeandparticularlyitsvery
in theApoidea (Melo, in prep.). reduced forewingvenation remained an enigmaticgroup
The"Sphecidae" ofBohartand Menkeformsalargeand forarelativelylongtimesinceitsproposalbyNagy(1969).
diverse assemblage of predatory wasps, attacking most Thisauthorclearlyhadveryconfused ideasaboutitsrela-
insect orders, as well as spiders [see Iwata (1976) and tionships with otherAculeata lineages, since he placed it
Bohart and Menke (1976) for prey records]. In the monu- in a large, heterogeneous assemblage combining
mentalrevisionaryworkofBohartand Menke(1976), this 'Ampulicidae, Dryinidae and Cleptidae'. Brothers (1975)
group was divided into 11 subfamilies: Ampulicinae, placed Heterogyna in his plumariid group based on Nagy
Sphecinae, Pemphredoninae, Astatinae, Laphyragoginae (1969). Day (1984), upon gathering material of new spe-
(containing only the genus Laphyragogus), Larrinae, cies from Africa, provided convincing evidence that
Crabroninae, Entomosericinae (containing only Heterogyna belonged in the Sphecidae sensu Bohart and
Entomosericus), Xenosphecinae (containing only Menke, placing itina separatesubfamily. Day (1984) also
Xenosphex), Nyssoninae (= Bembicinae; see Menke 1997) described for the first time the females, which are brac-
and Philanthinae. Larrinae and Crabroninae have been hypterousand haveaveryunusualmorphologycompared
treated underonenameinthepast(e.g.,Evans 1964a)and to other sphecoid wasps. The phylogenetic analyses by
more recently, Lomholdt(1985) and Menke(1988),among Alexander(1992a)andespeciallybyBrothersandCarpen-
others,haveadvocatedsuchclassification.Alternativeclas- ter (1993) confirmed Day's placement ofHeterogyna, and
sifications, based on a division oftheaculeate wasps into in the latter work, the genus was assigned to a separate
several superfamilies, have recognized a superfamily family, the Heterogynaidae.
Sphecoidea, with the subfamilies of Bohart and Menke Alexander (1992a) was the first to investigate the rela-
treatedasfamilies(e.g.,Krombein 1979).Othershaveused tionships among the major lineages of the apoids using
only one superfamily for bees and sphecoid wasps, but modern phylogenetic methods. His study combined two
raised all sphecoid subfamilies to family level (e.g., majorsetsofmorphologicalcharactersusedpreviouslyin
Finnamoreand Michener 1993). Bohartand Menke (1976) determiningrelationshipsamongsphecoid wasps: Evans'
revised allgeneraofsphecoid waspsthenknown,provid- larvalcharacters [seereviewsinEvans(1959a, 1964a)]and
ing subfamilial, tribal and generic identification keys, as Bohart and Menke's (1976) adult characters. One of the
well a summary of the known aspects of the biology for majorproblems ofEvans' and Bohartand Menke's works
each genus. istheirassumption thatthemajorlineagesof"Sphecidae"
Becauseofthedistinctfeeding habits ofbees compared couldbeproperlyclassifiedwithoutincludingbeesamong
to other aculeates, including sphecoid wasps, the older them,evenafteradmittingthatsomelineagesofsphecoid
Linnaean classifications fortheAculeata always had bees waspsseemedmorecloselyrelated tobeesthantotherest
and sphecoid wasps in separate higher categories. The of "Sphecidae". Before Alexander's (1992a) study,
sphecoid waspswereusuallyamonga largegroupoffos- Lomholdt (1982) had already proposed dividing the
sorial wasps (e.g., Shuckard 1837) and the bees, like the "Sphecidae" into two, according to him, monophyletic
ants, were not recognized as having any clear links to a groups,oneunitingSphecinae+Ampulicinaeandtheother
particular group. Despite relatively earlier recognition of containingalltheremainingsphecid subfamilies,forming
the close relationship between bees and sphecoid wasps his Larridae, which he considered the sister group of the
(Miiller 1872), a formal classification placing these two bees.Alexander'sstudyalsoprovidedampleevidencefor
groups into one superfamily was proposed much later the paraphyletic nature of Sphecidae sensu Bohart and
(Handlirsch 1907). Such a classification received strong Menke; however, none of his analyses specifically sup-
supportfromMichener's(1944)studyontherelationships ported Lomholdt's phylogeny. The overall results of
among bees. Brothers' (1975) study on the phylogenetic Alexander's analysesareinconclusive regarding therela-
relationships within the Aculeata provided reliable evi- tionships among the major groups ofApoidea, especially
dence, in terms of shared derived features, for the close because numerous conflicting relations are supported by
proximitybetweenbeesandsphecoidwasps. Basedonthe oneormoreofhisanalyses. Hewaswellawareofthepre-
phylogenetictreeobtainedinhisstudy,heplacedbeesand liminarystatusofhisworkandconcludedthatmuchmore
sphecoid wasps in hissuperfamilySphecoidea; Michener remained tobe done.
(1986)hasshown,however,thatApoideaisthevalidname
Scientific Papers, Natural History Museum, The University of Kansas
The PresentStudy
sas. He was an outstanding teacher, and I profited im-
The present study developed from an investigation of mensely from being his student. I also thankall members
the relationships among the genera of the tribe of my dissertation committee, in particular Charles
PemphredoninisensuBohartandMenke(1976). Earlyinto Michener, Steven Ashe, Deborah Smith and Bryan
that study I realized that their Pemphredonini seemed to Danforth for their suggestions to the dissertation manu-
bediphyletic,butproperevaluationofthisquestionwould script. Michener was very kind for taking me as his stu-
requireabroaderinvestigationoftherelationshipsamong dent after the sudden death ofAlexander, and I am very
the different sphecoid lineages. Because of the poor reso- grateful forhishelpand support.Specimensforthiswork
lution obtained byAlexander (1992a) when using mainly were kindly provided by several institutions. I thank in
character systems from previous authors, I decided to re- particular Wojciech Pulawski of the California Academy
peat his studybutmostlyusing original characters and a ofSciences forproviding material ofvery important taxa,
representation of taxa not requiring hypotheses ofmono- Robert Brooksforallowingmetodissectnumerousspeci-
phylyabove thelevel ofgenus (Alexanderused thetribes mens from the Entomology Division of the Natural His-
recognized by Bohart and Menke). For obvious reasons, tory Museum of The University of Kansas, and Michael
the Pemphredoninae received closer attention and better Prentice for allowing me to use his unpublished data on
representation. Despite this bias, I am confident that re- the biology and larval morphology of Odontosphex
sults obtained here represent a fair investigation into the paradoxus, as well as for calling my attention to the close
phylogeny ofthe majorapoid lineages. relationship of Odontosphex and Entomosericus to
The preferred phylogenetic hypothesis found by this Pemphredoninae. Ialsothank BruceCutlerforallhishelp
study is used to propose a higher level classification for withtheworkIcarriedoutinhislabandAntonioMarques
theApoidea. From now on, I will be using the classifica- formakingavailablehisMacintoshcomputer. Duringmost
tion proposed here, and reference to higher taxa whose of my stay at The University of Kansas, I was financially
previous definitions conflict with theones proposed here supported by a scholarship from the Brazilian Conselho
willbe marked as such. Nacional de Desenvolvimento Cientifico e Tecnologico
(CNPq) (200233/92-0).ThankstoStevenAsheand Robert
Acknowledgments
Brooks,Iwassupportedbyacuratorialassistantshipfrom
The present work is part of my Ph. D. disserta- the Natural History Museum during my last semester. I
tion, and as such, I would like to thank posthumously am also grateful to the Department of Entomology and
Byron Alexander, first, for agreeing to be my Ph. D. ad- the PanoramaSociety forproviding fundsforsomeofmy
viserand alsoforallhissupportand guidanceduringmy research activities.
initialyearsofgraduatestudiesatTheUniversityofKan-
MATERIALAND METHODS
SelectionofRepresentativeTaxa genera, i.e. genera whose taxonomic positions in Bohart
and Menke's classification were not supported by
iewanegcreleRsuedtopeafrdkeeasicnneunilttnehataetotpiearvsceecsasoseonuoftnutaslttlwgumrhdaoyej,unoparsstelawlixeenalc.eltaiTganewsgsootofhflseiAomspimpoteeiicnidbgfaeisafcaalwcttaelxoriarne:s- Aov1,felremAsxaipattoenyirdodiefaerlK'aasondf(se1asp9seo9v,s2eiawrt)eaelrsdteouitadnhlyes.trohBeetexasixainamsd,eiecsnitnetdchpo.ealrlMtteaoicxcrtauielolanirrsetolsefedpvteaihcnneitTmUatenbanilxs-ae
availability ofspecimens (adult insects) forcomplete dis- are listed below.
section and of published information on larval morphol-
ogy. However, a few important taxa whose larvae areun- Material of the following additional hymenopteran
known were still included; also, Laphyragogus and taxa not included in the analyses were also completely
Xenosphexwereincluded,despitelackofmaterialforcom- dissected and exa—mined:
plete dissections (only mouthparts, including oral plate, Crabronidae. Astatinae:AmmoplanopscockerelliPate
andexternalgenitaliaweredissected).Insomecases,Ialso (female), AmmopIaneUus wnatilla Pate (female),
chose specific genera that were considered previously to Ammoplanellus sp. (female), Dryudella sp. (female);
havea relativelybasalpositionwithintheirrespectivelin- Bembicinae: Alysson melleus Say (female), Argogorytes sp.
eages; this is the case for the exemplar taxa of bees, of (male); Crabroninae: Bothynostethus sp. (male), Ectemnius
Sphecidae(s.str.),and formostoftheCrabroninaeand the stirpicola (Packard) (female), Entomognathus texanus
Bembicinae.Table1 liststheexemplartaxaincludedinthe (Cresson) (male), Oxybelus emarginatum Say (female and
formal parsimony analyses. Among the Crabronidae, I male),Tn/poxylonfrigidumSmith(male);Pemphredoninae:
tried to include representatives of all subfamilies recog- Araucastigrnus masneri Finnamore (female), Arpactophilus
nized by Bohart and Menke (1976), as well as "problem" sp. (female),Carinostigmussp. (female),Diodontusatratulus
Major Lineages ofApoidea
ingTatbolteh1e.clLaissstioffictaatxiaonusperdopaosseexdemhpelrea.rsF;=infgermoaluep;tMaxa=amrarlaen.gedaccord- Table 1. Continued
Philanthinae:
M
48.AphilanthopsfrigidusSmith,
OUTGROUP 49.Pliilanthusgibbosus(Fabricius),F,M
Bethvlidae: Heterogvnaidae:
M M
1.Epyrissp.(fromBrazil),F, 50. HeterogynafantsilotraDay,
Pompilidae: Sphecidae(sensustricto):
M
2.Notocyphussp. (fromCostaRica), 51.Chlorionaerarium Patton,F M
Rhopalosomatidae: 52.Palmodesrufiventris(Cresson),M
3. RhopalosomanearticumBrues,F 53.Podaloniacommunis(Cresson),
Sapygidae: 54. Stangeellacyaniventris (Guerin-Meneville),F
4. Eusapygaproximo (Cresson),F
Scolebythidae:
5. ClystopsenellalongiventrisKieffer,F Taschenberg(male),MicrostigmusnigrophthalmusMelo(fe-
Sierolomorphidae: M male), Passaloecus cuspidatus Smith (female), Pemphredon
6.SierolomorphacanadensisProvancher, lethifer (Shuckard) (female), Polemistus braunsii (Kohl)
INGROUP (male), Polemistus dickboharti Menke (female), Spilomena
Ampulicidae: subterraneaMcCorquodale&Naumann(female),Spilomena
789...AADpomlhpieucllhoeutxroumssap.srpu(.ffi(rvfoermnotrCmiosCsoTtsuatrnRaiecRrai,)c,aM)F,F,M sKpo.h(lfe(mfaelmea)l,e)S;tiPghmiulsafnutlhviinpaees:FoCxer(cemrailse)r,uSfotpiigcmtuasStmeimtpohra(lfies-
Apidae(sensulato): male).
—
1110..AConntahnotphhaloincdtausalnbiagtraic(aTnismbTeirmlbaekrel)a,keF,F Apidae (s.l.). CalliopsisandreniformisSmith(female),
12.Ctenocolletesstnaragdinus(Smith),F Callomelittaantip—odes(Smith) (female),Hylaeussp.(female).
13.HesperapiscarinataStevens,F Mutillidae. Myrmosa unicolorSay (male).
14. Lonchopriazonalis (Reed),F —
Crabronidae: Pompilidae. Apori—nellusfasciatus (Smith) (female).
A1s5t.atAisntaaet:a nevadicaCresson,F Sierolomorphidae. Sierolomorpha nigrescens Evans
16.Eremiaspheciumsahelense(Simon-Thomas),F (male). —
1178..APumlmvoeprrloanmeusscaclfre.roapPaacthee,PF,atMe,F Vespidae. —Eumenesfratemus Say (female).
19. TimberlakenayucaipaPate,F Braconidae—. unidentified species (female).
B2e0m.bEiecmibneacei:nusquinquespinosusM(Say),F Evaniidae. u—nidentified species (male).
21.Didineistexana(Cresson), Trigonalidae. —unidentified species (male).
2223..HHoeplliioscoaiudseussslpairlroopitdeersus(S(pHianonldal)i,rsFch),F Xiphydrii—dae. Xiphydria sp. (female).
24. Nysson rusticusCresson,F Xyelidae. Macroxyelaferruginea (Say) (male).
C2r5a.bOrcohnlienwapet;erabipunctata(Say),F Undissected or only partially dissected (e.g.,
26.Dinetuspictus(Fabricius),M terminalia) specimens of the following taxa were also
27.LaphyragoguspictusKohl examined:
M —
2289..MXeelnloisnpuhsexalpteismtbreirslaCkaemieWriolnl,iams Ampulicidae. Aphelotoma fuscata Riek (female and
30.AnacrabroocellatusPackard,F male), Dolichurus corniculus (Spinola) (female),
31.Lindeniuscolumbianus (Kohl),F Paradolichiirus boharti Kimsey (female and male),
32. Lyrodasubita (Say),F Paradolichurusobidensis(Ducke)(femaleandmale),Trirogma
33.NitelaamazonicaDucke,F
M caerulea Westwood (female).
34.Palarus latifronsKohl,M —
35.PlenoculusdavisiFox, Crabronidae. Clypeadonlaticinctus(Cresson)(male),
Pemphredoninae: M EntomosericuskaufmanniRadoszkowski(femaleandmale),
36.OdontosphexparadoxusMenke,
37.EntomosericusconcinnusDahlbom,F Eremiasphecium budrysi (Kazenas) (female), E. longiceps
38.MimesacressoniiPackard,F,MM (Gussakovskij) (female and male), Laphyragogus ajjer
39.Plutominutus(Malloch),F, Beaumont(femaleand male),Mellinusarvensis (Linnaeus)
40.PsenulusmayorumBohart&Grissell,F (femaleand male),Mellinus bimaculatus Packard (female),
4412..ADripoadcotnotpuhsilruusgossteuisndFaocxh,neF,riMKohl,F Odontosphex damara Pulawski (female and male), Palarus
43.Parastigmushuecuvus FinnamoreM,F variegatus (Fabricius) (female and male), Paracrabro
44.PassaloecusareolatusVincent,F, froggratti Turner (female), Tiguipa cfr. fiebrigi (Brethes)
45.Pemphredon inornataSay,F (male), Timberlakena cahuilla Pate (female), Xenosphex
4467..SSptiilgommuesnatecmpaotraamlairscaKoAhnlt,roFpov,F xerophilus Williams (female).
Scientific Papers, Natural History Museum, The University of Kansas
Heterogynaidae—HeterogynaproteaNagy(femaleand CharacterSelection andDelimitation
male) and males of all African Heterogyna species, except Most of the characters used in the present study are
forH. ravenala Day. derived from the morphology of the exoskeleton of the
DissectionofAdultSpecimens adultinsects,includinginternalprocesses (e.g.,furca,2nd
phragma).Selectionofcharacterswasbasedondry,pinned
in TAatblleeas1to(nexecaedputltLsappehcyirmaegnogfursomaenadchXoefntohsepthaexxa)liwstaesd supseicnigmeansst,earseowseclolpiacs omnicdriosssceoctpeedOslpyecmipmuesnsSiZn6g0ly(cueprint,o
processed as followsbefore examination: 126X) and incident and transmitted light. The remaining
(1) Soaking in 10% KOH solution overnight; characters were taken from the morphology of immature
(2)Clearingin3% hydrogenperoxideforspecieswitha stages (larva) and from thebehavior ofadult females.
dark integument; In order to confirm the glandular nature of two
(3) Transfer to 50-70% ethanol, then boiling for a few characters (82 and 83), female specimens of Ammoplanus
minutes, followed by slow cooling; cfr. apache, Passaloecus areolatus, and Stigmus americanus
(4)Transfertowaterand thenslowadditionofglycerin; preserved inKahle'sfixativewereembedded inLRWhite
(5)Transfer toand storage in pure glycerin. resinfollowingtheproceduresdescribedbyLindley(1992)
Inordertoavoidexcessiveclearingofmouthpartsand and sectioned with a Sorvall Ultra Microtome (MT 5000);
terminalia, these parts were dissected before transferring thesectionswereslide-mountedusingEuparal.Theslides
toperoxide.Boilinginethanolisimportanttoremove,from wereobservedandphotographedunderaOlympusBH-2
inside the specimen, especially from the head, bubbles microscopewith differential interference contrast optics.
produced by the peroxide. Partial dismembering of the It is difficult to explain or justify the process of
specimen was carried out before transferring to pure characterdiscovery and subsequentdelimitation ofthose
glycerin,sinceatthisstagetheintegumentisstillrelatively characters into states. For complex characters, i.e.
KOH
malleable from the treatment. characters that show a great amount of apparently
For species in Table 1 with only one sex listed, the importantcladistirinformationbutarenotreadilydivisible
terminaliaoftheoppositesexwereremovedandsubmitted intodiscretestatesurexpressibleinaquantitativemanner
tothesameproceduredescribedabove,sothatsexspecific (e.g., Characters 58 or 68), I tried to provide detailed
characters from this part of the body could be examined; descriptions and illustrations, so that the states here
forthreegenera,onlymaterialofothercongenericspecies recognized can be apprehended and more properly
was available: a male of Eusapyga sp., a female ofAphelo- evaluated by other people. But the problem of making
toma nigricula RiekandafemaleofHeterogynaprotectNagy. explicit the decision processes followed when delimiting
The heads of specimens preserved in fixative (or thestatesforthesecomplexcharactersstillremains.Inmost
sometimes in alcohol) of the following species were such cases, I included a larger number of states to match
dissected for examination of the morphology of the more closely the condition present in the different taxa.
pharynx (see characters 13 and 14): However, if taken to an extreme and if the states are
Ampulicidae.——Dolichurus sp. (Costa Rica, male). nboynaadsdsiitginvien,gtahidsifpfreorceendtusrteatceafnomraekacehatnayxcohna.racteruseless
Apidae (s.l.). —Augochlora pura (Say) (female). In some cases, to preserve the informational content
Crabronidae. Alysson melleus Say (Bembicinae, of a complex character, I divided it into two characters,
female), Crossocerus sp. (from USA, Crabroninae, male), oneofthemrepresentingpresenceorabsenceofastructure
Didineis texana (Cresson) (Bembicinae, female), Diodontus or of a particular condition and the second character
flavitarsis Fox (Pemphredoninae, female), Hoplisoides sp. representingitsdifferentstates(e.g.,Characterpairs70and
(fromCostaRica,Bembicinae,female),Nyssonspp. (female 71, or 77and 78). Taxa inwhich thestructureorcondition
fromUSA,malefromCostaRica,Bembicinae),Ochleroptera is absent are assigned a question mark for the second
bipunctata (Say) (Bembicinae, female), Philanthus gibbosus character; this corresponds to treating inapplicable
(Fabricius)(Philanthinae,female),Psenulussp.(fromCosta charactersasmissingdata.Thiscanbeproblematicunder
Rica,Pemphredoninae,female),Spheciusspeciosus(Drury) certaincircumstances Maddison1993),butthecurrent
(e.g.,
(Bembicinae, male), Spilomena alini Antropov computer algorithms are not able to handle inapplicable
(Pemphredoninae, female), Stigmus americanus Packard charactersdifferentially.Onealternativewouldbetohave
(Pemphredoninae, fem—ale). only one character, but this is exactly what was being
Sphecidae (s.str.). Isodontia sp. (from USA, female), avoided in the first place.
Sphex ichneumoneus (Linnaeus) (male). Another issue that should be mentioned is the
—
Sapygidae. Sapyga sp. (from USA, male). treatment given to morphometric characters (approxi-
Major Lineages ofApoidea
mately 25% of the characters used here, not considering Thisassumption isbased on presenceamongmembersof
meristiccharacters,e.g.,Character4or73). Inmostcases, thistribeofadisproportionalelongate1stsubmarginaland
a quantitative description was adopted to express a therelativelywideseparationbetweenlm-cuand thevein
qualitative nature not easily captured as such; most here interpreted as 2rs-m, especially in genera like
commonly, this qualitative nature involved shape of Diodontus and Pemph—redon.
structures. For example, the labrum (Character 1) in Ammoplanini. The presence of two or only one
Pemphredonini and most Ammoplanini is relatively submarginalcell in thisgroupseemsatfirstmoredifficult
similarinoverallshapeand ina fewotherattributes,butI toexplain,becausemembersofthebasal lineage, Pulverro
tried to express this similarity mostly by the proportions (andAmmoplanops),haveaveryreducedvenationpattern.
ofthelabrum(state1-1);characters19and20areadditional However, I am postulating that the ancestral lineage for
examples. thistribehad twosubmarginalcellsasseeninsomespecies
Different procedures have been developed to divide of Timberlakenn (and also in Pwtostigmus). This two-celled
M
morphometriccharactersintomoreorlessobjectivestates conditionwascreatedbylossof3rs-mand distalto2rs-
m
taking into consideration intra- and intertaxon variation (character 89). These postulated vein losses, instead of
[see reviews by Stevens (1991) and Thiele (1993)], as well loss of the segment of Rs separating the 1st and 2nd
astoproducecharactersrepresentingshape(e.g.,Zelditch submarginalcellsassuggestedforthePemphredonini,are
et al. 1995). I did not employ any of these procedures for inferred from the close proximitybetween lm-cu and the
the quantitative characters used here. The characters veinhereinterpreted asthissegmentofRsinPulverroand
selected are believed to show very little intraspecific Timberlakenn(seeFigs.20and21).Also,thevenationpattern
variation, since they were chosen exactly for being stable of the Ammoplanini can be easily derived by patterns
acrossatleasttworepresentativetaxa. However,thelimits similar to that ofsome species of Eremiasphechtm (see Fig.
for the different states were usually arbitrary (see e.g., 22) assuming—the changes postulated here.
Characters 8, 9 or 29) and were defined to circumscribe Dinetus. Thetwo-celled condition in this genus can
two or more taxa thought to form a natural group. This be derived easily from a shortening of the marginal cell
approach has been criticized for its "otential to bias the accompaniedbylossof3rs-mand thesegmentofMdistal
phylogenetic analyses in favor of pre-conceived ideas of to 2rs-m. The genus Gastrosericus (Crabroninae, Larrini)
rSeulbamtiitotnisnhigptsh(ees.eg.c,hSatreavcteenrss1t9o91t,heGipfrtocaenddurSetsevmeennsti1o99n7e)d. hasaNistiemlial,arLcmonddeintiiouns,bauntdclAenaarlcyrianbdwe.p—enTdehnetlpyreasceqnuicreedo.f
above could diminish these potential biases, but I think it only one submarginal cell in these three genera can be
would represent little improvement for the quantitative derived from a pattern with two submarginal cells (in
ccahlcaacrsoasrcidtfieinregsdlyuastlelodt1hh3eei9rrecn[hastaeeurraaeclst(oesreFesarTruaisbsle(ed1949)i0.n)]T.htihIsneasanonymaeclwayshseae,tsI wsahengidmclehonststhooeff2tRhnsedsseceepglalmreaistnitpneogtfito2hlreast1-esm)tanbonytdlf2ousnssdeodsfuwMbimtahdriRgssti.anlIanltdoeceetldhl,es
cwrhuodseescolratdiisntgiccainnfboremautsieodnatlociodnetnetnitfyshtohuolsde bcehavraicetweerds iannEdnicnopsoognmaetlsuptesci[esseeoFfiNgi.te1l1a6,AaicnleBaorhainrdtiacnatdiMonen(okreth(e19v7e6i)]n
with more caution. remainsinthecaseofNitela)ofapetiolate2ndsubmarginal
In the case of absence of oneor more of the veins cell canbe seen.
—
delimiting theforewingsubmarginalcells (seeCharacters Hesperapis. The two-celled condition in this genus
87-89), I used relational information to infer the putative is assumed to have occurred by loss of 2rs-m. This
bloesesnes.ciTrhicsumpvreonbtleedmboyfsciomnisliadrietryinagssoenslsymetnhteunsuuamlbleyrhaosf assusbummaprtgiionnaliscbeallse(di.eo.n2pnrdesaenncde3ofrda rfeulsaetdi)velaynldonlgm-2cndu
submarginal cells present [e.g., Alexander's (1992a) connected to the segment ofM delimiting the2nd cell. In
Character 59, or Alexander and Michener's (1995) bees, lm-cu apparently almost always connects to M
Character84]. However, I think this approach can lead to between the segment of Rs separating the 1st and 2nd
loss of information, and therefore I tried to introduce a submarginal cells and 2rs-m.
mlIiopnrreoaegveipsdreoefcmiCsorearbearsodsneeitsadsialmeeehdnatjv,uesteliosfspitecscaitoaimolenlsyoffboterhcetasheuesveesiinmssie.lvaHererirately veryHerteerdougcyenda.—wiTnhgisgveennautsihoans.asIoumseewhhaetreuntuhseuaslaamned
assessments made in each case: interpretation given by Day (1985) for H. protea: first
Pemphredonini.—The presence of only two submarginal cell complete, second cell petiolate and
submarginal cells in this group is considered here to be distallyopen(2rs-mpresent,butnotreachingspectral M).
derived from loss of the segment ofRs separating the 1st I made no especial effort to include known
and 2nd submarginal cells, i.e. fusion of these two cells. synapomorphic characters or to look for new putative
8 Scientific Papers, Natural History Museum, The University of Kansas
synapormorphiesforthefollowingtaxa previouslyfound by Evans and Lin (1956b), and Ctenocolletes, from the
tobe monophyletic: Sphecidae (sensu stricto) (Alexander descriptionofthelarva ofStenotrituspubescens (Smith) by
1992a), Apidae (sensu lato) (Alexander 1992a, Alexander Houston (1975); these pairs of genera have very similar
andMichener1995)and thePhilanthinaesensuAlexander adult morphology and nesting behavior, respectively.
(1992b). Obviously, I paid close attention to any evidence Informationforoutgrouptaxawastakenatthefamilylevel
that could contradict these previous studies. Characters from Evans et al. (1987), except for Scolebythidae and
found by Brothers and Carpenter (1993) to support the Sierolomorphidae, for which there is no available
monophyly of Apoidea were included (however, several information.
of them were circumscribed differently), as well as These six characters were selected from a list of 10
additional relevant characters providing resolution for characters considered of phylogenetic significance by
relationships among the outgroup taxa. Evans(1959a).Twoofthe10wereomittedbecauseoftheir
The sampleofcharactersused here is not intended to apparentcomplexity(larvalbodyshapeand mandibles); I
representtheresultofanexhaustivesearchforinformative am not comfortable using them without examining the
characters, but only as one of the many samples of specimens. The two others (parietal bands and opening
charactersthatcouldbeextractedfromtheseinsects.Some between atrium and sub-atrium of spiracles) were given
areasofthebodythatseemtocontainimportantcharacters less importance by Evans. Alexander's (1992a) study can
werecompletelyignored. Forexample,theinternalridges beusedtoevaluatethesignificanceofthesefourcharacters
andlamellaeassociatedwiththehypostomalbridgeinthe ina cladistic context.
head exhibit a large amount of variation among the taxa Three behavioral characters were incorporated in the
sampled,butIwassimplyunabletoorganizethisvariation formal analyses: type of larval food, food relocation and
inanymeaningfulway.Thisregionofthebodyand many nest construction. All prey records for ampulicids,
othersareworthexploring in future studies. crabronids, and sphecids were taken from Bohart and
Terminology Menke (1976), except for Eremiasphecium taken from
Kazenas (1991; prey records for £. budrysi Kazenas),
The morphological terms adopted here were mostly Arpactophilus from Matthews and Naumann (1989; prey
taken from Bohart and Menke (1976), Michener (1944) or record forA. mimiNaumann),Ammoplanus from Maneval
Snodgrass(1942, 1993) anddefinitionsforthesetermscan (1939; prey record forA. perrisi Giraud) and from Ahrens
be found in those works. However, terminology for wing (1948; prey record for A. handlirschi Gussakovskij),
characters(seealsoFig. 14)wastakenfromDay(1988);for Laphyragogus from Kazenas (1985; prey record for L.
the thoracic pleuron, from Gibson (1993); for external turanicus Gussakovskij), Entomosericus from Kazenas and
genitalia, from Smith (1970). Sources fora few additional Alexander (1993; prey record for E. kaufmani
morphological terms are given directly in the list of Radoszkowski) and for Odontosphex paradoxus from M.
characters. Forindicationofdirection forstructuresinthe Prentice (pers. comm.). Biological information for the
head,Iusedtheconventionofaninsectwithaprognathous outgroup taxa was taken from Hanson and Gauld (1995).
head, so that the frons is in a dorsal position and the
occiput,ventral. Referenceismade tometasomalsclerites DataAnalysis
(Michener1944),insteadofabdominalsclerites,exceptfor Methods incorporating parsimony have won
gonocoxites and gonapophyses offemale's sting. widespread acceptance among systematists interested in
LarvalandBehavioralCharacters producingphylogenetichypothesesforthevariousgroups
of organisms, especially because parsimony has been
Six characters (131-136) derived from the external considered the only criterion that implements Hennig's
morphologyofthelarvawereused. Thedata foralllarval auxiliary principle [e.g., Hennig (1966:121)] that the most
characters weretaken from theliterature (Table2), except preferable tree topology is the one that minimizes the
for Spilomena; I also examined larvae of a few additional number of ad hoc hypotheses of homoplasy (e.g., Wiley
taxa, like Pemphredon, Psenulus,and Stigmus, as well as of 1981, Farris 1983). Parsimony is usually associated only
sometaxanotincludedinthepresentstudy,likeSceliphron, with methods that do not assign different weights to the
Cerceris,and Megaehile. Thereisnopublished information charactersbeingused,i.e. methodsinwhichallcharacters
on the larvae of nine genera of the ingroup: Aphelotoma, are treated equally in terms of cladistic information they
Ctenocolletes, Didineis, Eremiasphecium, Laphyragogus, provide. However,parsimonycanalsobeapplied undera
Xenosphex,Timberlakena,Parastigmus,andHeterogyna.These weightingfunction.Herein,Iemploytwodistinctmethods
taxa have missing entries for larval characters in the (impliedandsuccessiveweighting)thatassigndifferential
charactermatrix,exceptDidineis,forwhichthestateswere weights to the characters based on their degree of
taken from the description of the larva ofAtysson melleus homoplasy.