A cladistic analysis and classification of the subgenera and genera of the large carpenter bees, tribe Xylocopini (Hymenoptera: Apidae)

48 Pages·1998·3.5 MB·English

by  Minckley, Robert L

1
Papers
Scientific
Museum
Natural History
The University Kansas
of
07 August 1998 Number 9:1-47
A
Cladistic Analysis and Classification of the
Subgenera and Genera of the Large Carpenter Bees,
Tribe Xylocopini (Hymenoptera: Apidae)^
By
Robert L. Minckley^
Division ofEntomology. NaturalHistoryMuseum, andDepartmentofEntomology,
The UniversityofKansas, Lawrence, Kansas 66045-2109, USA
CONTENTS
ABSTRACT 2
INTRODUCTION
3
Acknowledgments 3
MATERIALS AND METHODS 3
Selection ofTaxa 3
Selection ofCharacters 5
CharactersListandCoding 7
PhylogeneticAnalyses 13
RESULTS 14
Relationships among Larger CladesoftheXylocopini 15
Relationships among Members ofthe Major Clades andTaxonomic Recommendations 21
KEYS TOTHE SUBGENERAOFXKLOCOPA 27
NewWorld Females 27
New World Males 28
Old World Females 28
OldWorld Males 29
DIAGNOSES AND DESCRIPTIONS OF SUBGENERAOFXYLOCOPA 3
SubgenusXylomelissa Hurd and Moure 31
SubgenusNodula Maa 31
'Contribution No. 3198 from the Department of Entomology and Snow Entomological Division, Natural History
Museum, The University of Kansas.
-Present address: Department of Entomology, 301 Funchess Hall, Auburn University, Auburn, Alabama 36849-5413, USA,
©NaturalHistoryMuseum,TheUniversityofKansas ISSNNo.1094-0782
Scientific Papers, Natural History Museum, The University of Kansas
QL Subgenus Maaiana new subgenus 32
Subgenus Bihinn Maa 32
^^ (X Subgenus Rhysoxylocopa Hurd and Moure 32
"-^ I Subgenus Pwxylocopa Hedicke, new status 33
A q- -2 Subgenus Nyctomelitta Cockerell 33
Subgenus Ctenoxylocopa Michener 33
V\ S'L:> Subgenus Xylocopa Latreille (sensu stricto) 34
Subgenus Xylocopoides Michener 34
I I ( Subgenus Calloxylocopn Hurd and Moure 34
Subgenus Copoxyla Maa 35
Subgenus Gnathoxylocopa Hurd and Moure 35
Subgenus testis Lepeletier and Serville, new status 35
Subgenus Notoxylocopa Hurd 36
Subgenus Schonnherria Lepeletier 36
Subgenus Dasyxylocopa Hurd and Moure 37
Subgenus Xylocopsis Hurd and Moure 37
Subgenus Nanoxyhcopm Hurd and Moure 37
Subgenus Cirroxylocopw Hurd and Moure 37
Subgenus Xylocopodn Hurd and Moure 38
Subgenus Stenoxylocopa Hurd and Moure 38
Subgenus Neoxylocopa Michener 38
Subgenus Prosopoxylocopa Hurd and Moure 39
Subgenus Zonohirsuta Maa 39
Subgenus Bomboixylocopa Maa 39
Subgenus Xenoxylocopn Hurd and Moure 40
Subgenus AUoxylocopa Maa 40
Subgenus Mesotricliia Westwood 40
Subgenus Koptortosoma Gribodo 41
LITERATURE CITED 42
APPENDICES 44
ABSTRACT Phylogenetic analyses of the genera and subgenera of the tribe Xylocopini (Large Carpenter
Bees) were madeusing morphologicalcharacterscodedfor51 speciesofthe ingroup.Thesespeciesrepresented
all butthreeofthepreviouslyrecognizedsubgenera.Twentyspeciesofbeesconstitutingtheoutgroupalsowere
examined toevaluate primitive and derivedcharacterstates within the Xylocopini. Fourdifferent analyses were
made because of uncertainty in hypotheses of character transformations and among the relationships of the
outgrouptaxa. Cladisticparsimonyanalysesshowedthatpreviousclassificationswerebasedoncombinationsof
charactersand not unique synapomoiphies. Three majorclades were supported in most analyses presented here;
twoarefromthe OldWorld, andone is fromthe New World.Abiogeographically mixed groupofsubgeneraare
not cleariy placed in this analysis; amongthis group are likely tobe found the basal members ofthe tribe.
A new generic and subgeneric classification is proposed forthe Xylocopini. I conclude that recognition of
Pwxylocopa Maa and testis Lepeletier and Serville as genera creates a paraphyletic genus o'iXylocopa: these
shouldberecognizedassubgeneraofanexpandedXy/oco/ja.Anew subgenus.Maaiana. isproposedhere forX.
hentoniand related species formerly recognized as members ofthe subgenusNodiila. The following subgeneric
synonymies also are proposed: Xylomelissa Hurd & Moure isexpanded to include Epixylocopa Hurd & Moure,
AUoxylocopa Hurd& Moure,Dinoxylocopa Hurd&MoureandAcroxylocopa Hurd& Moure;Ancylocopa Maa
is synonymized with Proxylocopa Maa; Schonnherria Lepeletier is expanded to include loxylocopa Hurd &
Moure andXylocospila Hurd & Moure; Stenoxylocopa Hurd & Moure is expanded to include Xylocopina Hurd
& Moure; Neoxylocopa Michener is expanded to include Megaxylocopa Hurd & Moure; Bomboixylocopa Maa
isexpandedtoincludeMinioxylocopaHurd& Moure;MesotrichiaWestwoodisexpandedtoincludePlatynopoda
Westwood,Hoplitocopa Lieftinckand Hoploxylocopa Hurd & Moure, and; Koptortosoma Gribodo isexpanded
Subgenera and Genera of Carpenter Bees. Xylocopini
to include Ow.wlocopa Hurd & Moure, CyphoxYlocopa Hurd & Moure. Cycmcoderes Hurd & Mt)ure and
Afroxxlocopa Hurd & Moure. A key to identify the subgenera i,s provided.
Ke\i words: Insecta; Hvmenoptera; Apidae, Xylocopini; Cladistic analyses; Classification.
INTRODUCTION
Thelargecarpenterbees,tribeXylocopini,generallyare catory changes were provided. Nevertheless, it is worth
robust, fast-flving insects, some of which are among the noting that the relationships generated by Da Cunha's
largest of all the bees. As their name implies, most mem- study should be used with caution, because similarity
bers of this group place their nests in dead solid plant methodscan result in poorestimatesofphylogeny (Hillis
material,usuallyvarious kindsofwood,butsomeusethe etal., 1994;Michener, 1970),especiallywhenthedataused
hollow culms of bamboo (subgenus Biluna; Maeta et al., were intended to diagnose groups and not to infer phy-
1985),orthepithvstemsofAloe(subgenusGiuitlioxylocopa; logeny.
Watmough, 1974). Females oi Pwxylocopa construct their In thisstudy, I usean exemplarapproachinthecladis-
nests in the ground. Some species of Xylocopini are im- tic analysis (see Phylogenetic Analysis). I also propose
portant pollinators of particular plants (e.g., Corbet and some changes in the classification based on my phyloge-
Wilmer, 1980; Gottsberger et al., 1988) but none seems to netic analyses. Myapproachfortaxonomicdecisionsis to
restrictitspollencollectingtooneorafewplanttaxa.Other combineterminaltaxathataremonophyleticbasedonthe
well-studied features of the biology of the group include phylogenetic analyses and retain the names of the prob-
itssocialbehavior (reviewed inGerlingetal., 1988; Maeta lematic taxa as they were recognized before this study.
andSakagami, 1995;Michener, 1990),exocrineglandmor- Future analyses may result in the recognition of fewer
phology(Mincklev, 1994;Vinson, 1994)andproductchem- supraspecific groups than are accepted here.
istrv (Gerlingetal, 1988),and malereproductivebehavior
Acknowledgments
(Alcock, 1992; GerUng et al, 1988; Minckley, 1994).
This paper is partofa dissertationpresented as partial
The Xylocopini is one of four tribes in—the subfamily requirement forthedegree ofPh. D. to the Departmentof
Xylocopinae. The other—three tribes Allodapini, Entomology,TheUniversityofKansas.Thecuratorsofthe
Ceratinini, and Manueliini are mostly smaller than the institutions listed in the section on Selection of Taxa (be-
Xylocopini and build theirnests inbroken twigs or stems low) are gratefully acknowledged for the loan of the ma-
(Daly et al., 1987; Michener, 1974, 1990). terialused inthisstudy. Dr.CharlesD.Michenerprovided
The purpose of this study is a cladistic analysis of the generoussupportand guidancein thisproject fromitsin-
phylogeneticrelationshipsamongthegeneraandsubgen- ception. Others who wereinfluential in thecompletion of
era ofthe large carpenterbees, tribe Xylocopini. Aworld- thisprojectincludeDrs. B.Alexander,J.S.Ashe,WilliamJ.
wide revision of this group was last undertaken in 1963 Bell, E. O. Wiley, S. G. Reyes, R. A. B. Leschen, A. Roig-
bvHurdandMoure.Theyrecognized53supraspecifictaxa Alsina,D.YanegaandM. D.Greenfield. BruceCutler,who
inthetribe:Threegenera,testis,Pwxylocopa,and Xi/locopa; allowed me access to his laboratory and gave consider-
48subgenerainXylocopaandtwosubgenerainProxylocopa. ableguidance in the analysis ofsome ofthe internal mor-
Although their work is the most comprehensive study of phological structures used in the study, deserves special
thegroup todate,both withrespecttothenumberoftaxa commendation. I dedicate this paper to the late Byron
examined and the number ofcharacters considered, they Alexander; he substantially improved the quality of this
did not provide anexplicit hypothesisofphylogeneticre- study and piqued my appreciation for the importance of
lationships for themembers ofthe tribe. Da Cunha (1992) systematics in evolutionarybiology. This workwas facili-
did a quantitative similarity analysis of the subgenera of tated by grants from the American Museum of Natural
Xylocopa using the characters of Hurd and Moure (1963). History (Theodore Roosevelt Fund), DepartmentofEnto-
This study was largely concerned with phenetic method- mology at The University of Kansas (Hungerford Fund),
ologyandwasnota taxonomicrevisionoftheXylocopini, and by the National Science Foundation grant DEB-
so no discussion of phylogenetic relationships or classifi- 9302186 (C. D. Michener, principal investigator).
MATERIALSANDMETHODS
Selection ofTaxa
In order to examine ingroup relationships, I used ex- Xylocopa were not included in the analyses: Euxylocopa,
emplartaxa ofallsupraspecific taxa oftheXylocopinirec- Lieftinckella, loxylocopa, Moiwxylocopa, Diaxylocopa and
ognizedbyHurd and Moure (1963) for which I could ob- Dinoxylocopa. Euxylocopa waserected by Hurd and Moure
tain both sexes. This meant the following subgenera of (1963)fora speciesX. (Euxylocopa)fraiidulentaknownonly
—
!
4 SciENTinc Papers, Natural History Museum, The University of Kansas
Table 1. List oftaxa examined organized by thesubgenera recognized Table1continued
herein.Anasterisk(*)indicatestypespecies;(!)indicatestheexemplars
usedinthephylogeneticanalysis.Theacronyms,asexplainedinMateri-
alsandMethods,showthesourcesofthespecimens.Subgenericassign- Koptortosoma(continued)
msehnotwsnbiynpparreevnitohuesseasu.thors, if different than those proposed here, are XX..tairpiniciainlaisanStmai)tiah Ritsema ((KKSSEEMM))
X.caerulea (Fabricius) (Cyaneoderes) (KSEM)
!
NCainroroxxty/llooccooppnn XX..dvUesattitaa BHuurrmdeiastnedrM*!oure*! ((UKSSNEMM)) XXX...cknaiufghfrnriaita(FLr(iFianebsnreaie!cuis(u)Csy)!p*h!ox(yAlforcooxpyal)ocopa) (((KKRSSNEEHMM)))
Xylocopoda X.elegnns HurdandMoure*! (KSEM) X.ocularis Perez ! (Cyphoxylocopa) (RNH)
Schonnherria X.bambusae Schrottky! (Xylocospila) (KSEM) X.parva Enderlein ! (Oxyxylocopa) (BM)
X.loripes Smith (KSEM) X.pubescens Spinola! (TAV)
X.macrops Lepeletier (KSEM) X.scioensis Gribodo (KSEM)
!
XX..ivinriicdain<s^asLterpaelLeetpieelret!ier! ((KKSSEEMM)) XX..ssienneinosrisVaScmhiatlh ! ((KKSSEEMM))
X.vindis Smith (KSEM) X.varipes Smith ! (Oxyxylocopa) (BM)
X.clm/sopoda Schrottky! Uoxylocopa) (USNM) testis X.aeratus Smith ! (KSEM)
Dasyxylocopa X. bumculata Friese*! ' (KSEM) X.bombylans (Fabricius) (KSEM)
Xylocopsis X.fimestci Maidl*! (KSEM) Proxylocopa X.olivieri (Lepeletier)* (TAV)
Monoxylocopa X.abbreviata HurdandMoure* (USNM) X.nifa (Friese) ! (TAV)
Diaxylocopa X.truxali HurdandMoure* (LACM) X.altaica (Popov) (Ancylocopa) (ZIA)
Notoxylocopa X.guatenmlensis Cockerell (KSEM) X.andarabana (Hedicke) (Ancylocopa) (ZIA)
X.tabaniformisnndroleuca Michener* (KSEM) X. nitidiventris (Smith) (Ancylocopa) (ZIA)
X. t.nztecnCresson (KSEM) X. nix (Maa) (Ancylocopa) (ZIA)
X. t. tabaniformis Smith ! (KSEM) X.parviceps (Morawitz) (Ancylocopa) (ZIA)
Xylocopoides X.californicaarizonensis Cresson (KSEM) X.pavlovskyi (Popov) (Ancylocopa) (ZIA)
X.cyanea Smith ! (KSEM) X.przcioalskyi(Morawitz) (Ancylocopa) (ZIA)
X.pallidiscopa Hu!rd (KSEM)
X.virginica Linnaeus* (KSEM)
Calloxi/locopa X. tenuata Smith*! (KSEM) fromfemalesand latersynonymizedbyMaa(1970)under
Neoxyhcopa X.aeneipennis (Degeer) (KSEM)
X. b.brasilianonim (Linnaeus)* (KSEM) X. (Perixylocopa)en/thrina. Iwasnotabletoexamineamale
X.dnrivim Cockerell (KSEM) of Lieftinckella. The remaining four monotypic subgenera
XX..ff.imfbrronitaatlaisFa(Oblriivciieurs)!((MMeeggaaxxyyllooccooppaa)) ((KKSSEEMM)) were examined, but are known from only one sex
X.grisescens Lepeletier (KSEM) loxylocopa and Dinoxylocopa from malesandMonoxylocopa
X.mexicanorum Cockerell (KSEM)
X.varipuncta Patton (KSEM) and Diaxylocopafromfemales.Forsomelargeorseemingly
!
Stenoxylocopa X.artifex Smith»! (KSEM) variable subgenera, additional species were included.
X.m. micheneri Hurd (KSEM) Character analysis was restricted to these exemplars. The
Ctenoxylocopa XX..freunfersctolrlaitsaH(uFrabdriacniuds)Mo»!ure! (Xy"locopina) ((FKMSNEHM)) twosubgenera ofProxylocopa differonly in one character,
Copoxyla XX..isruiskat(i(pfhersisMt)a*!a ((KKSSEEMM)) the size of the ocelli, so 1 included only Proxylocopm riifa
Rhyso'xylocopa X.cantabrita Lepeletier*! (KSEM) after initial examinations. All the taxa examined and the
Gnathoxylocopa XX.,kucolbersoiwcinsiBsraPuenrsez ((UUSSNNMM)) respective repositories ofspecimens that I used are listed
X.sicheli Vachal*! (USNM) in Table 1. Because the taxonomic categories recognized
Xylocopa X.valga Gerstaecker! (KSEM) by Hurd and Moure (1963) were narrowly defined, these
X.violacea (Christ)*! (KSEM)
Biluna X.auripennis Lepeletier! (KSEM) exemplars are likely to provide a good representation of
Bomboixylocopa XX..cltirianneqnuiesbaFrroireusme!(Swederus) ((UKSSNEMM)) themorphologicaldiversityandareasonablefirstestimate
X. rufipes Smith ! (Mimoxylocopa) (KSEM) ofthe phylogenetic relationships in the group.
NyctomeUita X. myops Ritsema (KSEM)
X. tranquebarica (Fabricius)*! (KSEM) Specimens used in this study were from the following
Nodula X.a.amethystina (Fabricius) (KSEM) institutions (acronyms in parentheses): Field Museum of
XX..abmeenttohhyistiCnoacke(rFealbrlicius)*! ((KKSSEEMM)) NaturalHistory,Chicago,A. Newton(FMNH);LosAnge-
ZProonsoohpiorxsyultoacopa XX..dmeijreaabniiliisLeHpuerledtiaenrd!!Moure*! ((AKMSNEHM)) les County Museum, Los Angeles, R. Snelling (LACM);
Xylomelissa XX..aalbbsiufrrdoinpsesLeEpnedleetrileeri!n ! {Dinoxy'locopa) ((UUSSNNMM)) NInasttiiotnutael,CSooluletchtiAofnriocfa,InCs.ectEsa,rdPllaenyt(PPrPolt)e;ctTihoen NRaetsieoanraclh
XX..ccaarpiintaattanSSmmiitthh!*(Acroxylocopa) ((UKSSNEMM)) Museum of Natural History, Washington, R. McGinley
X.erythrina Gribodo! (Perixylocopa) (PPl) (USNM); The Natural History Museum, London, G. Else
X.lugubris Gerstaecker (Apoxylocopa) (PPl)
Alloxylocopa XX..arupfpietnadrisicsulLaetpaecliertciwenrv!ol!{aEnpsixSyfmoictohpa*)! ((KKSSEEMM)) v(BaMn);AcRhitjekrsbemrugse(uRmNHv)a;n NSantouuwrlEinjtkoemHoilsotgoirciea,lLMeuidseenu,mC,.
Xenoxylocopa X.chiyakensis (Cockerell)*! (KSEM)
X.inconstans Smith (KSEM) University ofKansas, Lawrence, C. D. Michener (KSEM);
Mesotrichia X.acutipennis Smith! {Hoploxylocopa) (BM) Tel-Aviv University Tel-Aviv, D. Gerling (TAU); Zoologi-
XX..afslsaivmoirluifsa R(iDtesGeemear!)(Hoplitocopa) ((KRSNEHM)) cal Institute,Academy ofSciences, Leningrad, Y. Pesenko
X.latipes (Drury) ! (Platynomda) (KSEM) (ZIA).
X. tenuiscapa Westwood (Piatynopoda) (KSEM)
X. torrida (Westwood)*! " (KSEM) Phylogenetic relationships of the large carpenterbees
Koptortosoma X.aestuans (Linnaeus) (KSEM) with other tribes of Xylocopinae were analyzed by
Subgenera and Genera of Carpenter Bees, Xylocopini
TABLE2. Listofoutgrouptaxaexamined. stated, I use the term outgroup to refer to a hypothetical
ancestor that represents the inferred ground-plan state of
Apidae:Xylocopinae all characters. Ground-plan character states for the
Ceratinini Xylocopiniwerehypothesizedusingtheoptimizationtech-
CC.em(tEiucncara(tCienraa)ticnyaa)nceuacuKribribtvina(Rossi) nique of Maddison et al. (1984). This hypothetical ances-
C.(Lioceratinn)fliwolatcralhCockerell torhasall characterscodedaszero,witha fewexceptions
C.(Certinidia)japonicaCockerell (Table3).
C.(Calloceratina)kelaSpinola
C. (CreiivHa)rupcstris SelectionofCharacters
C. (Pithitis)smaragdula(Fabricius) I used all characters that fulfilled the conditions ofho-
C. (Ceratiiiula)zetekiCockerell mology (Remane, 1956), includingsomecharacters ofthe
MCctgeiniocccerraattiinnaastcaunlgpatnuyriacteanc(iSsmi(tSht)rand) malesthatarehomoplasiousinotherbees.Apriori,1could
Allodapini think of no reasonable criterion that would allow me to
Compsomelissazaxantha (Cockerell) decide whether one character or character system (e.g.,
Exonueridiahakariensis male genitalia vs. external sexual characteristics of the
ExonuerabicolorSmith
male) was phylogenetically more informative than any
Halterapisnigrinen'is(Cameron)
Manueliini other. This approach relies on the congruence with other
Manueliagayatinn(Spinola) characters to determine the general "goodness of fit" of
M.gayi(Spinola) anysinglecharacter(Kluge, 1989). Furthermore,ina large
M.postica (Spinola)
Megachilidae group, such as the Xylocopini, it seems reasonable to ex-
Pararhophitini pect that some characters homoplasious within some
PannhophitesorobinusMorawitz clades will serve as synapomorphies ofotherclades.
Fideliini
ParafideliafrieseiBrauns Homologyassessmentwasproblematicforsomechar-
Lithurgiini acters in Proxylocopa females ofwhich have a well-devel-
,
LithurgeapicalisCresson oped basitibial plate at the base of the tibia and a large
conspicuous pygidial plate. These conditions are typical
Sakagami and Michener (1987) and Roig-Alsina and for most other ground-nesting bees, but in the rest of the
Michener (1993). Both studies supported Ceratinini and largecarpenterbeesthepygidialplateconsistsofonlyone
Allodapini as sister groups and the terminal members of orseveralspines,andthebasitibialplateisreduced (some-
the subfamily. The sister group to the Ceratinini + timesonlyrepresentedby its apex) and positioned on the
Allodapini clade is less certain; Sakagami and Michener apicaltwo-thirdsofthetibia.Theplacementanddevelop-
(1987) concluded that this group was the Xylocopini, but mentofthesestructuresinProxylocopahaveledsomework-
the analysis of Roig-Alsina and Michener (1993), which ers to consider it as the most primitive taxon in the sub-
used more characters, concluded that it could be either family (Xylocopinae sensii Hurd, 1958; Malyshev, 1913).
XylocopiniorManueliini.Roig-AlsinaandMichener(1993) Others have argued that thesestructures collectively rep-
favored theplacementoftheManueliinias themostbasal resentanadaptivecomplexassociatedwithnestingin the
taxoninthesubfamilybecauseXylocopini,Allodapiniand ground and have evolved secondarily (Daly et al., 1987).
Ceratinini share a derived shape of sternum eight in the Morphological evidence supports the latter view; in
males. They also addressed relationships of groups out- Proxylocopathepygidialplateandbasitibialplateareswol-
sidethesubfamilyXylocopinae.Thenextmorebasaltaxon len lobes and completely lack a line or carina around the
to the Xylocopinae was the family Megachilidae (consist- lateraledgesasistypicalofotherbees. Furthermore,ifthe
ing of the subfamilies Megachilinae, Fideliinae and Manueliini, a wood-nesting group, is the basal taxon in
Osmiinae). thesubfamilyXylocopinae,theinferredground-planstate
Ichosecomponentsoftheoutgroupforthisstudybased for the Xylocopini would be that its ancestor nested in
largelyontheanalysisofRoig-AlsinaandMichener(1993). woodandProxylocopawouldhavesecondarilyrevertedto
ForthreeoftheanalysesIpolarized ground-planstatesof nesting in ^he ground. In this study, I treated the above
characters based on a topology ofManueliini as thebasal conditionsinProxylocopaasuniquederivedfeatures.How-
taxonin thesubfamilyXylocopinaeand a sister-group re- ever,becausenocomparablestructuresexistelsewhere in
lationshipamongXylocopiniandAllodapini+Cera-inini. the ingroup (Xylocopini) and some characters are trans-
From the family Megachilidae, I used three taxa (Lithurge formation series with multiple states, I coded all charac-
apicalis, Parafidelia friesei and Pararhophites orobinus) con- ters associated withthebasitibialplateand pygidial plate
sideredbasalinthefamilyasadditionaloutgroups.Names asmissinginProxylocopa.Preliminaryanalyses(notshown)
ofalloutgrouptaxa arelisted inTable2. Unlessotherwise made with these features coded as primitive usually pro-
Scientific Papers. Natural History Museum, The University of Kansas
Table3. DatamatrixusedinthephylogeneticanalysesforAnalysis1,1A,3,and3A.MatricestorAnaly-
ses2,2A,4and4Aareexplained inthetext. Numbersacrossthetopofmatrixcorrespond tocharacter
numbers used in the text. Taxon abbreviations used were Xylocopoides' = X. californica arizonensis,
Xx/locopoides'=X.cyanea,Xylocopa^=X.violacca,Xylocopa-=X.valga,Koptortosoma^=X.sinensis,Koptortosoma-
=X.caffra, Koptortosonm^=X.pubescens,Schonnherria^=X.inridagastra,Schonnherria-=X.micans,Schonnlwrria'
=X.macrops,Nodula^=X.bentoni,No1duln'=X2.amethystina,Notoxylocopa'=X.labaniformisandNotoxylocopci-
=X.guatemalensis.
Character 3 4 5
Taxon 123456789 0123456789 0123456789 0123456789 0123456789 0123456
Outgroup 000?00000 0000000000 00??000000 0000000000 0000000070 0000000
Nanoxylocopa 001000000 0100000001 0100102000 0010002000 0101000111 1020000
Cirroxylocopa 001000000 0100000001 0100101100 0010000000 0101000110 1120000
Xylocopoda^ 001001000 0100000001 0100101100 0000000000 0101000100 1020000
Dasyxylocopa 011000000 0100000001 0100101000 0000002000 0101000112 1120100
Xylocopsis 001000000 0100000011 0100102000 0000002000 0101000111 1120100
Xylocopmdes' 000000000 1210000101 1100004000 0100002000 0210100100 1001000
Calloxylocopa 000000000 0100000001 0100000000 0000002000 0210100100 1000000
Neoxylocopa 101001000 0110000001 0100103100 0020000011 1101000100 1120000
Megaxylocopa 101001000 0110000001 0100103100 0020000011 1101000100 1120000
Stenoxylocopa 001111000 0110000001 0100100100 0020002010 1101000100 1120000
Xylocopina 001111000 0110000001 0100101000 0000000000 1101000100 1120000
Ctenoxylocopa 000100000 1100000111 0000002000 0120002000 1100000100 0000110
Copoxyla 000100100 1100000001 0100000000 0000002000 1000000111 0000100
Rhysoxylocopa 000000000 1101000201 0100000000 0020000000 0100000110 0020000
Gnathoxylocopa 001100000 1100000111 0100100000 0010002000 1100000111 1010100
Xylocopa 000100000 1210000101 0000001000 0000002000 0110000100 0001010
Nyctomelittn 011000001 0101000111 0100102000 0000000000 0101000111 1010000
Zonohirsuta 000000000 0210002001 0111002000 0000000000 0100000000 1020000
Prosopoxylocopa 000000000 0210000000 0111001000 0000000000 0100000101 1110000
Bomboixylocopa 000000010 0210000000 0100001000 0000010001 1000000000 0021000
Mimoxylocopa 011000010 0210000001 0100115000 0000012001 1100000000 1121000
Alloxylocopa 000001000 0210000000 0111110000 0221111000 0100000000 1021000
Xenoxylocopa 100001100 0210000001 010111!010 1020011100 0000000000 1121000
Mesotrichia 000001100 0210102000 0122115011 1021211110 0110000100 1120010
Pktynopoda 000000100 0210100000 0122001001 1221211010 0000010100 0011010
Hoplitocopa 011000100 1210100000 012210!001 1221211010 0000010100 1120110
Hoploxylocopa 000001100 0210112000 012211!Oil 1121211000 0000000000 0020010
Afroxyiocopa 110001100 0210000001 0123115100 1021211100 1000000102 1121100
Oxyxylocopa 000001100 0210010000 012311!010 1221111000 0000000101 1121000
Cyaneoderes 001001100 1210000000 012310!010 1221211100 0000000101 1121000
Cyphoxylocopa 001000000 0210002000 012311!010 1221111100 0000000101 1120000
Koptortosoma' 101000000 0210000000 010211!010 1121211100 1000000102 1021000
Koptortoscma- 101000000 0210100000 0123111010 1121211101 1000000101 1121000
Koptorlosomn^ 111000000 1210000000 0123115010 1101211101 1000000102 1021000
Pnxylocopa 000000001 0200000!!! 1100111000 0000000000 0:00000110 0000000
testis 000000000 0101000001 1100000000 0000002000 0100000111 1000011
Schonnherria} 001000000 0100001011 0100102000 0220000000 0101000112 1120100
Schonnherria^ 011000100 0100000011 0100102000 0220012000 0101000112 1120000
Schonnherria^ 001100100 0100000011 0101102000 0220002000 0101000112 1120000
Xylocospila 001000000 0100000001 0100101000 0020002000 0101000112 1120000
Epixylocopa 001000000 1000000201 0100012000 0020002001.1100000111 0020000
Perixylocopa 011001000 0000001201 0100102000 0000002001 1100000111 0020000
Acroxylocopa 000101010 0000001201 0100112000 0020000001 1100000111 0020000
Bihinn 000000000 0000001000 0100112100 0020000000 1100000110 0020000
Apoxyhcopa 000000000 0000001201 0100102000 0020000001 1100000111 0020000
Xylomelissa 000000000 0001000201 0100102000 0020000001 1100000111 0020000
Nodula' 001000000 0101000201 0100111000 0020000001 1100000110 0020000
Nodula- 000000000 0110000201 0111000000 0020000001 1100000110 0020000
Notoxylocopa' 001000000 0110000001 0100110000 0000002001 0101001112 1010100
Notoxylocopa^ 001100000 0210000101 0100110000 0000002000 0101001112 1010100
Xylocopa- 001100000 1210000101 0000001000 0000012000 0100000100 0001010
Xylocopoides^ 001000010 1210000101 1100004000 0100002000 0210100100 1001001
Subgenera and Genera of Carpenter Bees. Xylocopini
duced trees with Pwx\/Iocopn arising as thebasal taxon in
the tribe.
Terminologygenerally follows that ofMichener (1944,
1965) and Hurd and Moure (1963), except that I use me-
dial (as opposed to lateral) instead of mediolongitudinal
for structures on the tibial spine, propodeum, terga and
sterna.TergaandsternaaredesignatedasTandS,respec-
tively,and segmentsarenumbered fromanteriortoposte-
rior, e.g., SI, S2, etc. 1 follow Brothers (1976) in the use of
metapostnotum instead of propodeal triangle, and
Michener(1975)andReyes(1991)intheuseofventroapical
plate for the ventral distal projection of the male
gonocoxite.Mandibularstructuresaredescribedusingthe
terminology ofMichener and Fraser (1978).
Examinationofthereservoirsofthemesosomalglands
of the males required dissection of the posterior part of
thethoraxusingtechniquesdescribedbyMinckley(1994); Fig. 1. External views of left mandibles of females: A. Xyhcopa
this was limited to specimens forwhich I had permission (Megax\/locopm)* fimbriata. B. X. (Gnathoxylocopa) sicheli. C. X.
to dissect. Forready comparison, the characters aregiven s(aSt=ensouxbyacpoilocpaal)tmoiocthhe.nSecrailmei=ch1enmermi..AFbigb.reBviraetdiroanws;nfar=omacHeutrabdulaanrdgMrooouvree,
the same number in the cladistic analysis and the group (1964). 'RegardedasamemberofthesubgenusofNeoxylocopaherein.
descriptions. The cladograms were produced using the
computerprogramPAUP(PhylogeneticAnalysisUsingPar-
polarity decision is equivocal. Furthermore, in those
simony) Version 3.1.1 (Swofford, 1993) and analyzed with
outgroup taxa with three mandibular teeth (Ceratinini,
MacCladeversion 3.05 (Maddisonand Maddison, 1993).
AllodapiniandParafidelia),themedialtoothislongest,but
Characters andCoding in the Xylocopini the ventral tooth (ventral tooth of the
Thecharacters,asnumberedbelow,correspond tothe rutellum)islongest,possiblysuggestingthattheadditional
numbers in all the subsequent tables and trees and in the toothofsomeXylocopinihasarisendenovo.1considerthree
groupdescriptions.Numbersinparenthesesarethecodes teeth homologous in this analysis and coded the ground-
used in the data matrix and on the cladograms. Unless plan state as equivocal.
noted otherwise, is considered—theplesiomorphic state. 5. Upper—rutellar tooth and preapical tooth (i.e. apex
1. Color of male vestiture. (0) dark brown, black, of pollex). (0) Separated by an emargination (Fig. IB);
green,orpale;(1)tawnyorferruginousonatleastdorsum (1) fused and forming a continuous edge (Fig. IC; Hurd
ofbody. State 1 is not found in the outgroup. 1978, Figs. 1,2). State 1 is notfound in the outgroup. This
mal2e. mSaunbdaipbilcea.l—to(o0)thAbosrencton(vFiegx.iItBy);o(n1)upprepseerntc(aFriign.alAo,f aisptohmeorspahmyeiassshtahreed"ibnyneSrtetnoooxytlho"coopfaHaunrddXi/(l1o9c7o8p)i.naTwaon-d
IC). State 1 is widespread inbeesand occurs in thegenus toothed mandibles (Fig. lA) werecod—ed 0.
Ctenoceratina (Ceratinini) of the outgroup. Because 6. Epistomal suture of female. (0) Not raised or
Ctenoceratinahasanumberofderivedfeatures(Daly,1988), broadly impunctate (Hurd and Moure, 1963, Fig. 16); (1)
it is not likely to be an ancestral group in the Ceratinini continuouslycarinateand broadlyimpunctate (Hurd and
and therefore not closely related to the Xylocopini; I infer Moure, 1963, Figs. 14, 15, 17, 18). In some ingroup taxa, a
State 1 to be apomorphic for the Xylocopini. carinaispresentontheupperepistomalsuturebutitdoes
3. S—ubapical tooth on upper carina of female man- notextend completelydownthelateraledges,acondition
dible. (0) Absent; (1) present. See preceding character. 1 coded as 0. State 1 does not occur—in theoutgroup.
4. Number of teeth on apical margin of female man- 7. Supraocellarpits of female. (0) Punctiform ornot
dible.—(0)Two (Fig. lA); (1) three(Fig. IB). Mandiblesas present; (1) deeply excavated and crater-like. In the
shown in Figure IC are coded as having three teeth be- outgroup, supraocellarpits, if—present, are punctiform.
cause the two upper teeth are considered to be fused. In 8. Malar space of female. (0) Shorter than width of
the outgroup, the Ceratinini, Allodapini and Lithiirge flagellar segment 1 (Hurd and Moure, 1963, Fig. 51); (1)
(Michenerand Fraser, 1978) havethree teeth on theapical muchlongerthan widthofflagellarsegment 1 (Hurd and
margin of the mandible, and Manuelia, Pararhophites and Moure, 1963, Fig. 16). State 1 isnotfound in theoutgroup
Parafidelia havetwoteeth. Based on outgroupanalysis the exemplarsusedhere,butitispresentamongotherspecies
Scientific Papers, Natural History Museum, The University of Kansas
ts
Fig.2. Outerviewofhind tibiaeoffemales:A.Xylocopa (Xyhcopoides)cyanea. B. X. (Ctenoxylocopa)fenestrata.C.
testis*bomhylans. D.Proxylocopa*riifa. E.X. (.Alloxylocopa)circumvolans.Abbreviation: ts=tibialspines.Scale= 1 mm.
•RegardedasasubgenusofXylocopaherein.
—
in the outgroup that I did not consider (Michener, 1975; 15. Mid and hind trochanter of male. (0) Spines ab-
Reyes, 1993). sent; (1) ventral surface of each with spine. State 1 does
—
9. Ocelli. (0) Not modified; (1) enlarged. State 1 oc- notoccur in theoutgroup. —
curs in the subgenera Nyctomelitta andProxylocopa (sensu 16.Ventralsurfaceofmalehindfemur. (0)Notmodi-
stricto) of Xylocopa. All outgroup taxa have normal-sized fied;(1)withbasaltubercle(Fig.3B);(2)withspineonbasal
ocelli. half(Fig.3C).Intheoutgroup,modificationssuchasthese
10. Male facial maculations.—(0) Yellow on at least are known only in some Ceratina (Hirashima, 1969). This
some part of face; (1) without yellow markings, i.e., en- characterwas coded as nonadditive.
tirely black. This is a highly variable character in the 17. A—nteriorand posterioredges of female basitibial
Apoidea but in the outgroup, males of all taxa except plates. (0)Weaklydevelopedorrisinglittlefromsurface
Lithurge, haveyellow maculations on the clypeus. ofleg; (1)withnumeroustuberclesonbothedges(Fig.2A,
11. Number of spines on outer apex of male hind 2B); (2) with numerous tubercles only on posterior edge.
tibia.—(0) None; (1) one; (2) two (Hurd and Moure, 1963; Thepolarity proposed heremaybewrong,becausethose
Fig.37).Outgrouptaxa lackthesespines.Thetransforma- ingroup taxa with the most strongly developed basitibial
tionseries was coded as additive. plates (a plesiomorphic condition?) have tubercles along
both edges (e.g. Ctenoxylocopa, Xylocopa sensu stricto).
12. Number of spines on outer apex of female hind Those taxa with a single row of tubercles (State 2) on the
tibia.—(0)One(Fig.2B,2C,2D,2E);(1)two(Fig.2A;Hurd posterior edge completely lack a carina, or tubercles, on
and Moure, 1963, Figs. 36, 38). Females of the outgroup theanterioredge.State1 isunknownintheoutgroup.This
haveone or two spines. — transformation series was coded as additive.
—
13. Tibial spine of female. (0) Originating at tibial 18.Apexofmalebasitibialplate. (0)Entireorabsent;
apex and lacking medial carina (Fig. 2A, 2D,2E); (1) well- (1) bifid. As discussed in Character 17, the polarity ofthis
developed and originating above, but extending to, apex character may be thereverse ofthat presented here. In all
oftibia (Fig. 2B,2C). State 1 is found onlyin taxa with one ingroup taxa with well-developed basitibial plates, the
outer tibial spine. State 1 is not found in t—he outgroup. apex of the structure is bifid and usually strongly so. In
14. Shape of female hind trochanter. (0) Not modi- theoutgroupthebasitibialplate,whenpresent,invariably
fied (Fig. 3B, 3C); (1) roughly triangular in ventral view hasanentireapex.Thecharacterwascoded withanentire
(Fig. 3A). State 1 is not found in the outgroup. apex as the plesiomorphic condition.
Subgenera and Genera of Carpenter Bees, Xylocopini
1 isuniquetoXi/locopoidcsand Lesfis.OthertaxainXylocopa
have lateral edges that converge basally, but do not com-
pletely close (e.g. X. [Xi/lomelissn] erytliriiin, X.
[Notoxylocopn] guatcmnleiisis), a contiition that could be
consideredasaweakfovea,but1havenotconsideredthese
tobe homologous toState 1.
—
21. Female basitibial plate. (0) Extending to basal
fourthoftibia (Fig. 2B); (1) reduced, notextending tobase
oftibia (Fig 2A, 2C, 2E). See discussion ofCharacter 17.
—
22.Beginningofposteriorthoracicdeclivityofmale.
(0) Smoothly rounded, beginning undefined (Hurd and
Moure, 1963, Figs. 24,27;Minckley 1994, Figs. 1A, IC,3A,
3C, 4A, 4C, 4E); (1) abruptly rounded (Hurd and Moure,
1963, Figs. 39; Minckley 1994, Fig. 3E); (2) sharply angled
(Hurd and Moure, 1963, Fig. 33; Minckley 1994, Fig. 5A).
See discussionin following character.
23. Beginning of posterior thoracic declivity of fe-
—
male. (0) Profile smoothly rounded, position often un-
defined (Hurd and Moure, 1963, Figs. 27, 28; Minckley,
1994,Figs. 1A, IC,3A,3C,4A,4C,4E,5C,5E);(1)abruptly
rounded (Hurd and Moure, 1963, Fig. 39; Minckley 1994,
Fig. 3E); (2) as a transverse carina that does not overhang
metanotum (Hurd and Moure, 1963, Fig. 33; Minckley,
1994, Fig. IE); (3) marked by transverse carina that over-
hangs entire metanotum (Hurd and Moure, 1963, Fig. 34;
Minckley, 1994,Fig.5A). Becausestates and 1 bothoccur
in the outgroup, I consider the ground-plan state to be
equivocal. States 2 and 3 are unknown in the outgroup
and are strong apomorphies within Xylocopa. This trans-
formation serieswas treated as additive.
—
24.Malemetapostnotum. (0)Present;(1)absent.The
metapostnotuminbeesishighlyvariableinsizeandshape
and, when present, is distinguished from the propodeum
by a change in the sculpturing of the cuticle and/orby a
line along its edge with the propodeum. Apparently the
metapostnotum can be lost in two ways: (1) absence of
uniquesculpturingandmarginallines;or(2)byanincrease
inthesizeofthepropodeumrelativetothemetapostnotum
until the latter is completely absent externally. In the
Xylocopini the metapostnotum invariablyis delimited by
a linealong the edge with the propodeum and a continu-
Fig. 3. A. Ventral view of the hind trochanter of female Xylocopa ous size-gradation across taxa is evident. The
({MCetseontorxiyclhoicao)pal)athioptetse.ntBo.taA.nCt.erAinotrervioirewvioefwtohfethfeefmeumruorfofthtehemmaalleeooffXX.. metapostnotum in some taxa is well developed and in
(Afroxylocofm)* nigrita. Abbreviations: F = femur, t = tubercle, Tr = tro- other taxa is vestigial or absent. Furthermore, in species
chanter,s=spine.ModifiedfromEardley,1983.Scale=1 mm.*Regarded themales ofwhich haveanenlarged posteriormesosoma
asamemberofthesubgenusKoptortoscmaherein. that accommodates the mesosomal gland reservoir, the
— metapostnotumtypicallyisabsent(Minckley,1994).These
19. Apex of female basitibial plate. (0) Entire (Fig. observations suggest that loss of the metapostnotum in
2D,2E); (1)bifid (Fig.2A,2B,2C).—Seeprecedingcharacter. theXylocopiniisbyachangeinthesizeofthepropodeum.
20. Female basitibial plate. (0) Open basally or, if In the Ceratinini, the presence of the metapostnotum is
closed,extendingontobasalfourthoftibia;(1)foveateand usually made evident by a change in sculpturing, and no
onapical3/4oftibia(Fig.2A,2C;Hurd, 1961,Fig.9).State gradation in size of the propodeum is apparent; the
10 Scientific Papers, Natural History Museum, The University of Kansas
oflarge carpenterbees, the function for this structure has
notbeenexamined.Thereistremendousvariationinsuch
featuresastheshapeofthetubulesthatmakeupthegland
reservoir and the size of the reservoir (Minckley, 1994),
whichmakeshomologyassessmentdifficultforsometaxa.
In these difficult taxa I have opted to preserve informa-
tion and have split the characters into a number of often
autapomorphicstatesratherthansimplyscoringthechar-
acterintostatesofpresenceorabsence.Thischaracterwas
coded as nonadditive.
—
27. Beginning of the thoracic declivity of male. (0)
Anterior to the propodeum (Hurd and Moure, 1963, Figs.
29-34; Minckley, 1994, Fig. lA, IC); (1) on thepropodeum
(Hurd and Moure, 1963, Fig. 27; Minckley, 1994, Fig. 5C,
5E). State 1 is not found in the outgroup.
—
28. Propodeumatitsconnectionwithmetasoma. (0)
Notmodified;(1)dorsallyelongated,formingaspine.This
apomorphy is unknown in the outgroup.
29. Male tegula.—(0) Not modified (Fig. 4A); (1) elon-
gate posteriorly (Fig. 4B). State 1 is not found in the
outgroup.
—
30.ShapeofthegradulusonTl. (0)GradulusofTlcon-
tinued posteriorly adjacent to lateral margin of metasoma
(Hurd and Moure, 1963, Fig. 43); (1) transverse (Hurd and
Moure, 1963,Fig. 44). State 1 is not found in theoutgroup.
—
31.Distributionofgradulionmaleterga. (0)present
Fig.4. Lefttegulaofthemaleof(A) Xi/locopnvaiipuiictaand (B) X. on Tl, T2 and T3, or more; (1) present on Tl and T2; (2)
(Platynopodn)*tenuisaipw.Abbreviations:S=scutum,Tg=tegula.Scale= present on Tl only. All members of the outgroup have
1 mm.*RegardedasamemberofthesubgenusMesotrkiiinherein. graduli on at least T1-T3, except for the ceratinine genus
Pithitis,whichhasagradulusonlyonTl (Hirashima,1969).
metapostnotum is either well developed or completely Based on this information and the argument that a
ambesteanpt.os1tinnofetrufmriosmlotshtisbypaattreerdnuctthiaotninintihtes sCuerraftaicneicniiifftehre- gradulus on each tergum is generally considered to be
plesiomorphic in bees (Michener, 1944), the inferred
entiationfromthepropodeumand notbya changeinsize
ofthepropodeumorthemetapostnotum.Therefore, con- ground-plan state for the ingroup is graduli on all terga
I posterior to T3, as well as on more anterior terga. The
sider the presence of a defined metapostnotum to be the
characterwas treated as additive.
ground-plan condition in the Xylocopini. —
— 32. Distribution of graduli on female terga. (0) Tl,
25. Female metapostnotum. (0) Present, including
thosewhere it is minute; (1) absent. Seediscussion in pre- T2 and T3, or more; (1) Tl and T2; (2) Tl only. Seediscus-
sion ofpreceding character.
ceding character. —
— 33.ModificationsofanteriorsurfaceofTl ofmales.
26. Reservoir of the male mesosomal gland. (0) Ab-
sent; (f) two widely separated pouches (Minckley, 1994, (0) Absent; (1) a fovea (Hurd and Moure, 1963, Fig. 42).
See discussion in following character
Figs. IC, ID, IE, IF, 3A, 3B); (2) single layer of tubules, —
usually abutting in the middle (Minckley, 1994, Figs. 3C, 34.ModificationsofanteriorsurfaceofTloffemale.
3D, 4A, 4B, 4C, 4D); (3) highly convoluted tubules, as in (0)Absent; (Da fovea (Hurd and Moure, 1963, Fig. 42); (2)
Neoxyhcopn (Minckley, 1994, Fig. 5E,5F); (4) singlemedial with an entrance to a acarinarium (Fig. 5; Eardley, 1983,
pouch with oneopening (Minckley, 1994, Fig. lA, IB); (5) Figs. 23, 24, 27). Neither a fovea nor an invaginated mite
numerous, highly convoluted tubules, as in Koptortosoma pouchoccursinotherbees. Thetransformationserieswas
(Minckley, 1994,Fig. 5A,5B).Thepresenceofareservoiris coded as additive. —
a uniquecharacterofmostXylocopini. The reservoir in at 35. Medial groove of Tl of female. (0) Linear; (1)
leastsomespeciesisassociated withagland thatproduces deeply sulcate (Hurd and Moure, 1963, Fig. 41). In the
asex-attractantpheromone,although,in thevastmajority outgroup, the medial groove is linear.