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Sections

  • INTRODUCTION
  • Selection of Taxa
  • Acknowledgments
  • Selection of Characters
  • Phylogenetic Analyses
  • RESULTS
  • Relationships among Larger Clades of the Xylocopini
  • KEYSTO THE SUBGENERAOF XYLOCOPA
  • New World Males
  • Subgenus Xylomelissa Hurd and Moure
  • Subgenus Nodula Maa
  • Subgenus Maaiana new subgenus
  • Subgenus Rhysoxylocopa Hurd and Moure
  • Subgenus Nyctomelitta Cockerell
  • Subgenus Ctenoxylocopa Michener
  • Subgenus Xylocopa Latreille (sensu stricto)
  • Subgenus Xylocopoides Michener
  • Subgenus Calloxylocopa Hurd and Moure
  • Subgenus Copoxyla Maa
  • Subgenus Gniithoxylocopa Hurd and Moure
  • Subgenus testis Lepeletier and Serville, New status
  • Subgenus Notoxylocopa Hurd
  • Subgenus Schonnherria Lepeletier
  • Subgenus Dasyxylocopa Hurd and Moure
  • Subgenus Xyhcopsis Hurd and Moure
  • Subgenus Nanoxylocopn Hurd and Moure
  • Subgenus Cirroxylocopa Hurd and Moure
  • Subgenus Xylocopoda Hurci and Moure
  • Subgenus Stenoxylocopa Hurd and Moure
  • Subgenus Neoxylocopa Michener
  • Subgenus Prosopoxylocopa Hurd and Moure
  • Subgenus Zonohirsuta Maa
  • Subgenus Bomboixylocopa Maa
  • Subgenus Xenoxylocopa Hurd and Moure
  • Subgenus Alloxylocopa Maa
  • Subgenus Mesotrichia Westwood
  • Subgenus Koptortosoma Gribodo
  • LITERATURE CITED

1

Scientific

Papers
Natural History Museum The University of Kansas
Number 9:1-47

07 August 1998

A Cladistic Analysis and Classification of the
Subgenera and Genera of the Large Carpenter Bees, Tribe Xylocopini (Hymenoptera: Apidae)^
By

Robert

L.

Minckley^
Museum, and Department of Entomology,

Division of Entomology. Natural History

The University of Kansas, Lawrence, Kansas 66045-2109, USA

CONTENTS
ABSTRACT INTRODUCTION
Acknowledgments
2
3 3 3

MATERIALS AND METHODS
Selection

ofTaxa

3
5

Selection of Characters

Characters List and Coding
Phylogenetic Analyses

7 13

RESULTS
among Larger Clades of the Xylocopini Relationships among Members of the Major Clades andTaxonomic Recommendations
Relationships

14
15

21

KEYS TO THE SUBGENERA OF XKLOCOPA
New World Females New World Males
Old World Females Old World Males DIAGNOSES AND DESCRIPTIONS OF SUBGENERA OF XYLOCOPA
Subgenus Xylomelissa Hurd and Moure Subgenus Nodula

27 27 28 28

29
3

31

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,

© Natural History Museum, The University of Kansas

ISSN No. 1094-0782

Scientific Papers,

Natural History Museum, The University of Kansas
32 32
32 33

QL
"-^

^^
I

(X
-2

A q-

V\
I
I

S'L:>

(

Subgenus Maaiana new subgenus Subgenus Bihinn Maa Subgenus Rhysoxylocopa Hurd and Moure Subgenus Pwxylocopa Hedicke, new status Subgenus Nyctomelitta Cockerell Subgenus Ctenoxylocopa Michener Subgenus Xylocopa Latreille (sensu stricto) Subgenus Xylocopoides Michener Subgenus Calloxylocopn Hurd and Moure Subgenus Copoxyla Maa Subgenus Gnathoxylocopa Hurd and Moure Subgenus testis Lepeletier and Serville, new Subgenus Notoxylocopa Hurd Subgenus Schonnherria Lepeletier Subgenus Dasyxylocopa Hurd and Moure Subgenus Xylocopsis Hurd and Moure Subgenus Nanoxyhcopm Hurd and Moure Subgenus Cirroxylocopw Hurd and Moure Subgenus Xylocopodn Hurd and Moure Subgenus Stenoxylocopa Hurd and Moure Subgenus Neoxylocopa Michener Subgenus Prosopoxylocopa Hurd and Moure Subgenus Zonohirsuta Maa Subgenus Bomboixylocopa Maa Subgenus Xenoxylocopn Hurd and Moure Subgenus AUoxylocopa Maa Subgenus Mesotricliia Westwood Subgenus Koptortosoma Gribodo

33
33

34 34 34
35

35
status

35 36 36

37 37
37

37
38
38

38 39
39

39

40
40

40
41

LITERATURE CITED APPENDICES

42 44

ABSTRACT

Phylogenetic analyses of the genera and subgenera of the tribe Xylocopini (Large Carpenter

Bees) were made using morphological characters coded for 51 species of the ingroup. These species represented all but three of the previously recognized subgenera. Twenty species of bees constituting the outgroup also were

examined

to evaluate primitive

and derived character

states within the Xylocopini.

Four different analyses were
the relationships of the

made because of

uncertainty in hypotheses of character transformations and

among

outgroup taxa. Cladistic parsimony analyses showed that previous classifications were based on combinations of characters and not unique synapomoiphies. Three major clades were supported in most analyses presented here;

two are from

the

Old World, and one

is

from the
this

New
is

World.

A biogeographically mixed
I

group of subgenera are
tribe.

not cleariy placed in this analysis;

among

group are

likely to

be found the basal members of the
conclude

A new

generic and subgeneric classification
testis Lepeletier

proposed for the Xylocopini.

that recognition

of

Pwxylocopa Maa and

and Serville as genera creates a paraphyletic genus o'i Xylocopa: these should be recognized as subgenera of an expanded Xy/oco/ja. A new subgenus. Maaiana. is proposed here forX. hentoni and related species formerly recognized as members of the subgenus Nodiila. The following subgeneric synonymies also are proposed: Xylomelissa Hurd & Moure is expanded to include Epixylocopa Hurd & Moure,

AUoxylocopa Hurd & Moure, Dinoxylocopa Hurd & Moure and Acroxylocopa Hurd & Moure; Ancylocopa Maa is synonymized with Proxylocopa Maa; Schonnherria Lepeletier is expanded to include loxylocopa Hurd & Moure and Xylocospila Hurd & Moure; Stenoxylocopa Hurd & Moure is expanded to include Xylocopina Hurd & Moure; Neoxylocopa Michener is expanded to include Megaxylocopa Hurd & Moure; Bomboixylocopa Maa is expanded to include Minioxylocopa Hurd & Moure; Mesotrichia Westwood is expanded to include Platynopoda Westwood, Hoplitocopa Lieftinck and Hoploxylocopa Hurd & Moure, and; Koptortosoma Gribodo is expanded

Subgenera and Genera of Carpenter Bees. Xylocopini
to include

Ow.wlocopa Hurd

&

Moure, CyphoxYlocopa Hurd
to identify the

&

Moure. Cycmcoderes Hurd

&

Mt)ure and

Afroxxlocopa

Hurd & Moure. A

key

subgenera

i,s

provided.

Ke\i words: Insecta;

Hvmenoptera; Apidae, Xylocopini; Cladistic analyses;

Classification.

INTRODUCTION
The large carpenter bees, tribe Xylocopini, generally are robust, fast-flving insects, some of which are among the largest of all the bees. As their name implies, most members of this group place their nests in dead solid plant material, usually various kinds of wood, but some use the hollow culms of bamboo (subgenus Biluna; Maeta et al.,
stems of Aloe (subgenus Giuitlioxylocopa; Watmough, 1974). Females oi Pwxylocopa construct their nests in the ground. Some species of Xylocopini are important pollinators of particular plants (e.g., Corbet and
1985), or the pithv

catory changes were provided. Nevertheless,

it is

worth

noting that the relationships generated by

Da Cunha's

study should be used with caution, because similarity methods can result in poor estimates of phylogeny (Hillis et al., 1994; Michener, 1970), especially when the data used were intended to diagnose groups and not to infer phylogeny.
In this study,
tic
I

use an exemplar approach in the cladis-

Wilmer, 1980; Gottsberger
restrict its

et al., 1988)

but none seems to

pollen collecting to one or a few plant taxa. Other

I also propose based on my phylogenetic analyses. My approach for taxonomic decisions is to combine terminal taxa that are monophyletic based on the

analysis (see Phylogenetic Analysis).
in the classification

some changes

well-studied features of the biology of the group include

behavior (reviewed in Gerling et al., 1988; Maeta and Sakagami, 1995; Michener, 1990), exocrine gland morphology (Mincklev, 1994; Vinson, 1994) and product chemistrv (Gerling et al, 1988), and male reproductive behavior
its

social

phylogenetic analyses and retain the names of the problematic taxa as they were recognized before this study. Future analyses may result in the recognition of fewer
supraspecific groups than are accepted here.

(Alcock, 1992;

GerUng

et al, 1988;

Minckley, 1994).
This paper
is

Acknowledgments
part of a dissertation presented as partial

The Xylocopini is one of four tribes in the subfamily Xylocopinae. The other three tribes Allodapini, are mostly smaller than the Ceratinini, and Manueliini Xylocopini and build their nests in broken twigs or stems

requirement for the degree of Ph. D. to the Department of Entomology, The University of Kansas. The curators of the institutions listed in the section on Selection of Taxa (below) are gratefully acknowledged for the loan of the material used in this study. Dr. Charles D. Michener provided

(Daly

et al., 1987;

Michener, 1974, 1990).

The purpose of this study is a cladistic analysis of the phylogenetic relationships among the genera and subgenera of the large carpenter bees, tribe Xylocopini. A worldwide revision of this group was last undertaken in 1963
bv Hurd and Moure. They recognized 53 supraspecific taxa in the tribe: Three genera, testis, Pwxylocopa, and Xi/locopa; 48 subgenera in Xylocopa and two subgenera in Proxylocopa. Although their work is the most comprehensive study of the group to date, both with respect to the number of taxa examined and the number of characters considered, they did not provide an explicit hypothesis of phylogenetic relationships for the

generous support and guidance in this project from its inception. Others who were influential in the completion of this project include Drs. B. Alexander, J. S. Ashe, William J.
Bell, E.

O. Wiley,

S.

G. Reyes, R. A. B. Leschen, A. Roig-

Yanega and M. D. Greenfield. Bruce Cutler, who allowed me access to his laboratory and gave considerable guidance in the analysis of some of the internal morAlsina, D.

phological structures used in the study, deserves special commendation. I dedicate this paper to the late Byron

members

of the tribe.

Da Cunha

(1992)

did a quantitative similarity analysis of the subgenera of Xylocopa using the characters of Hurd and Moure (1963). This study was largely concerned with phenetic methodology and was not a taxonomic revision of the Xylocopini,
so no discussion of phylogenetic relationships or classifi-

Alexander; he substantially improved the quality of this study and piqued my appreciation for the importance of systematics in evolutionary biology. This work was facilitated by grants from the American Museum of Natural History (Theodore Roosevelt Fund), Department of Ento-

mology at The University of Kansas (Hungerford Fund), and by the National Science Foundation grant DEB9302186
(C. D.

Michener, principal investigator).

MATERIALS AND METHODS
In order to

Selection of Taxa examine ingroup relationships,

I

used ex-

emplar taxa of all supraspecific taxa of the Xylocopini recognized by Hurd and Moure (1963) for which I could obtain both sexes. This meant the following subgenera of

Xylocopa were not included in the analyses: Euxylocopa, Lieftinckella, loxylocopa, Moiwxylocopa, Diaxylocopa and
Dinoxylocopa. Euxylocopa

was

erected by

Hurd and Moure

(1963) for a species X. (Euxylocopa) fraiidulenta

known only

4
Table
1.

SciENTinc Papers, Natural History Museum, The University of Kansas
List of taxa

examined organized by the subgenera recognized

Table

1

continued

indicates type species; (!) indicates the exemplars used in the phylogenetic analysis. The acronyms, as explained in Materials and Methods, show the sources of the specimens. Subgeneric assignments by previous authors, if different than those proposed here, are shown in parentheses.
herein.
asterisk
(*)

An

Koptortosoma (continued)
X. X. X. X. X. X. X. X. X. X. X. X. X. X. X. X. X. X. X. X. X. X. X. X.
tipnciilis

Smith
Ritsema
!

ariiana anta)ia
caffra

caerulea (Fabricius)
!

(Cyaneoderes)

Nanoxt/locopn Cirroxylocopn Xylocopoda Schonnherria

X. X. X. X. X. X. X.

X.dUata Burmeister *! vestita Hurd and Moure *! elegnns Hurd and Moure *! bambusae Schrottky (Xylocospila) loripes Smith
!

(KSEM)

(Linnaeus) kuhni Friese (Cyphoxylocopa)
!

(USNM)
(KSEM) (KSEM) (KSEM) (KSEM) (KSEM) (KSEM) (KSEM)

nigrita (Fabricius)

*!

(Afroxylocopa)

(Cyphoxylocopa) parva Enderlein (Oxyxylocopa) pubescens Spinola!
ocularis Perez
! !

(KSEM) (KSEM) (KSEM) (KSEM) (RNH) (KSEM) (RNH)
(BM)
(TAV)

macrops Lepeletier
inicans Lepeletier
viridi<^astra
!

!

scioensis

Gribodo
!

Lepeletier
!

!

Dasyxylocopa
Xylocopsis

X.vindis Smith X. clm/sopoda Schrottky X. bumculata Friese *!
X.fimestci

senior Vachal sinensis Smith varipes Smith

(KSEM) (KSEM) (KSEM)
(BM)

!

(Oxyxylocopa)

Uoxylocopa)
'

(USNM)
(KSEM) (KSEM)

testis

aeratus

Smith

!

bombylans (Fabricius)
olivieri (Lepeletier)
*

(KSEM) (KSEM)
(TAV) (TAV) (ZIA)
(ZIA) (ZIA) (ZIA) (ZIA) (ZIA) (ZIA)

Maidl*!

Proxylocopa

Monoxylocopa
Diaxylocopa Notoxylocopa

Xylocopoides

Calloxi/locopa

Neoxyhcopa

X.abbreviata Hurd and Moure * X. truxali Hurd and Moure * X. guatenmlensis Cockerell * X. tabaniformis nndroleuca Michener X. t. nztecn Cresson X. t. tabaniformis Smith X. californica arizonensis Cresson X. cyanea Smith X. pallidiscopa Hurd * X. virginica Linnaeus X. tenuata Smith *! X. aeneipennis (Degeer) * X. b. brasilianonim (Linnaeus) X. dnrivim Cockerell (Megaxylocopa) X.fimbriata Fabricius X.f. frontalis (Olivier) (Megaxylocopa) X.grisescens Lepeletier X. mexicanorum Cockerell
! !

(USNM) (LACM)
(KSEM) (KSEM) (KSEM) (KSEM) (KSEM) (KSEM) (KSEM) (KSEM) (KSEM) (KSEM) (KSEM) (KSEM) (KSEM) (KSEM) (KSEM) (KSEM) (KSEM) (KSEM) (KSEM)

nifa (Friese)
altaica

!

(Popov) (Ancylocopa)

andarabana (Hedicke) (Ancylocopa) nitidiventris (Smith) (Ancylocopa)
nix (Maa) (Ancylocopa) parviceps (Morawitz) (Ancylocopa) pavlovskyi (Popov) (Ancylocopa) przcioalskyi (Morawitz) (Ancylocopa)

!

!

X.varipuncta Patton
Stenoxylocopa
X. artifex

!

Smith

»!

X. m. micheneri

Hurd
!

Ctenoxylocopa

X. rufrcollis Hurd and Moure X.fenestrata (Fabricius) »! X. sukatipes Maa
X.iris ((fhrist)*! X. cantabrita Lepeletier X, uclesicnsis Perez
*!

(Xylocopina)
"

(FMNH)
(KSEM) (KSEM) (KSEM) (KSEM)

synonymized by Maa (1970) under I was not able to examine a male of Lieftinckella. The remaining four monotypic subgenera were examined, but are known from only one sex loxylocopa and Dinoxylocopa from males and Monoxylocopa and Diaxylocopa from females. For some large or seemingly variable subgenera, additional species were included. Character analysis was restricted to these exemplars. The two subgenera of Proxylocopa differ only in one character,
from females and
later

X. (Perixylocopa) en/thrina.

Copoxyla
Rhyso'xylocopa

the size of the ocelli, so

1

included only Proxylocopm

riifa

Gnathoxylocopa
Xylocopa
Biluna

Bomboixylocopa

NyctomeUita

Nodula

Zonohirsuta Prosopoxylocopa Xylomelissa

Alloxylocopa

Xenoxylocopa
Mesotrichia

Koptortosoma

X. kobrowi Brauns X. sicheli Vachal *! X. valga Gerstaecker X.violacea (Christ)*! X. auripennis Lepeletier X. tranquebarorum (Swederus) X.cliinenis Friese! X. rufipes Smith (Mimoxylocopa) X. myops Ritsema X. tranquebarica (Fabricius) *! X. a. amethystina (Fabricius) X. amethystina (Fabricius) *! X. bentohi Cockerell X. dejeanii Lepeletier X. mirabilis Hurd and Moure *! {Dinoxylocopa) X. absurdipes Enderlein X.albifrons Lepeletier! X.carinatn Smith* X. capitata Smith (Acroxylocopa) X. erythrina Gribodo (Perixylocopa) X. lugubris Gerstaecker (Apoxylocopa) X. rufitarsis Lepeletier {Epixyfocopa) X. appendiculata circwnvolans Smith *! X. chiyakensis (Cockerell)*! X. inconstans Smith X. acutipennis Smith! {Hoploxylocopa) X.assimilis Ritsema! (Hoplitocopa) X.flavorufa (DeGeer) (Platynomda) X. latipes (Drury) X. tenuiscapa Westwood (Piatynopoda) " X. torrida (Westwood) *! X. aestuans (Linnaeus)
! !
! ! !

(USNM) (USNM) (USNM)
(KSEM) (KSEM) (KSEM) (KSEM)

(USNM)
(KSEM) (KSEM) (KSEM) (KSEM) (KSEM) (KSEM) (KSEM)

examined and the respective repositories of specimens that I used are listed in Table 1. Because the taxonomic categories recognized by Hurd and Moure (1963) were narrowly defined, these exemplars are likely to provide a good representation of the morphological diversity and a reasonable first estimate
after initial examinations. All the taxa

of the phylogenetic relationships in the group.

(AMNH)
(USNM) (USNM) (USNM)
(KSEM)
(PPl) (PPl)

Specimens used in this study were from the following (acronyms in parentheses): Field Museum of Natural History, Chicago, A. Newton (FMNH); Los Angeles County Museum, Los Angeles, R. Snelling (LACM);
institutions

!

'

National Collection of Insects, Plant Protection Research Institute, South Africa, C. Eardley (PPl); The National

!

Museum

!

!

!

(KSEM) (KSEM) (KSEM) (KSEM)
(BM)

of Natural History, Washington, R. McGinley (USNM); The Natural History Museum, London, G. Else (BM); Rijksmuseum van Natuurlijke Historie, Leiden, C. van Achterberg (RNH); Snow Entomological Museum,

University of Kansas, Lawrence, C. D. Michener (KSEM);
Tel- Aviv University Tel- Aviv, D. Gerling
cal Institute,

(TAU); Zoologi-

!

(RNH) (KSEM) (KSEM) (KSEM) (KSEM) (KSEM)

Academy

of Sciences, Leningrad, Y. Pesenko

(ZIA).

Phylogenetic relationships of the large carpenter bees

with other tribes of Xylocopinae were analyzed by

(Pithitis) smaragdula (Fabricius) C.g. Malyshev. Ctenoceratina tanganyicencis (Strand) Mcgiiccratina sculpturata (Smith) Allodapini Compsomelissa zaxantha (Cockerell) Exonueridia hakariensis Exonuera bicolor Smith Halterapis nigrinen'is (Cameron) Manueliini Manuelia gayatinn (Spinola) M. This hypothetical ancestor has all characters coded as zero. 1 could think of no reasonable criterion that would allow me to decide whether one character or character system (e. concluded that it could be either Xylocopini or Manueliini. 1913). In this study. 1956). Lithurgiini which have a well-devel- Lithurge apicalis Cresson Sakagami and Michener (1987) and Roig-Alsina and Michener (1993). Roig-Alsina and Michener (1993) base of the tibia and a large conspicuous pygidial plate. 1958. (Ceratiiiula) zeteki Cockerell nique of Maddison et al. List of outgroup taxa examined. I For three of the analyses above conditions in Proxylocopa as unique derived features. argued that these structures collectively repOthers have resent an adaptive complex associated with nesting in the ground and have evolved secondarily (Daly et al. I polarized ground-plan states of characters based on a topology of Manueliini as the basal taxon in the subfamily Xylocopinae and a sister-group relationship among Xylocopini and Allodapini + Cera-inini. Furthermore. Ground-plan character states for the Xylocopini were hypothesized using the optimization tech- Cemtina (Ceratina) cucurbitina (Rossi) Kirbv C. such as the Xylocopini. I use the term outgroup to refer to a hypothetical ancestor that represents the inferred ground-plan state of Apidae: Xylocopinae Ceratinini C. Sakagami and Michener (1987) concluded that this group was the Xylocopini. including some characters of the males that are homoplasious in other bees. (1984). in a large group. favored the placement of the Manueliini as the most basal taxon in the subfamily because Xylocopini. (Euccratina) cyanea all characters. it seems reasonable to expect that some characters homoplasious within some clades will serve as synapomorphies of other clades. These conditions are typical for most other ground-nesting bees. Xylocopini TABLE 2. From the family Megachilidae. but the analysis of Roig-Alsina and Michener (1993). 1989). to the They also addressed relationships of groups out- The next more basal taxon Xylocopinae was the family Megachilidae (consisting of the subfamilies Megachilinae.. Parafidelia friesei and Pararhophites orobinus) conlargely wood and Proxylocopa would have secondarily reverted to nesting in ^he ground. Fideliinae and Osmiinae). I used three taxa (Lithurge apicalis. but in the rest of the large carpenter bees the pygidial plate consists of only one or several spines. Both studies supported Ceratinini and Allodapini as sister groups and the terminal members of the subfamily. Unless otherwise and some characters are transstates. 1987). I sensii Hurd. This approach relies on the congruence with other characters to determine the general "goodness of fit" of any single character (Kluge. Allodapini and Ceratinini share a derived shape of sternum eight in the males. which used more characters. Furthermore. (Calloceratina) kela Spinola C. (Lioceratinn) fliwolatcralh Cockerell C. a wood-nesting group. How- because no comparable structures exist elsewhere in sidered basal in the family as additional outgroups. Morphological evidence supports the latter view.. in Proxylocopa the pygidial plate and basitibial plate are swollen lobes and completely lack a line or carina around the family (Xylocopinae lateral edges as is typical of other bees. external sexual characteristics of the male) was phylogenetically more informative than any other. (CreiivHa) rupcstris C. gayi (Spinola) postica (Spinola) Megachilidae Pararhophitini Pannhophites orobinus Morawitz Fideliini Parafidelia friesei Selection of Characters used all characters that fulfilled the conditions of homology (Remane. I Brauns Homology assessment was problematic for some characters in Proxylocopa females of . with a few exceptions (Table 3). of all Names outgroup taxa are listed in Table 2. for the Xylocopini the basal taxon in the subfamily Xylocopinae. I coded all characters associated with the basitibial plate and pygidial plate as missing in Proxylocopa. and the basitibial plate is reduced (some- oped basitibial plate at the apex) and positioned on the The placement and development of these structures in Proxylocopa have led some workers to consider it as the most primitive taxon in the subtimes only represented by its apical two-thirds of the tibia. side the subfamily Xylocopinae. A priori. stated. Preliminary analyses (not shown) made with these features coded as primitive usually prothe ingroup (Xylocopini) formation series with multiple . The sister group to the Ceratinini + Allodapini clade is less certain. M. the inferred ground-plan state chose components of the outgroup for this study based would be that its ancestor nested in treated the on the analysis of Roig-Alsina and Michener (1993). (Certinidia) japonica Cockerell C.Subgenera and Genera of Carpenter Bees. male genitalia vs. is if the Manueliini. ever.

Notoxylocopa' = X. inridagastra Schonnherria. macrops. Xx/locopoides'= X. Noduln'=X.= X. Table ses 3. labaniformis and Notoxylocopci2. Data matrix used in the phylogenetic analyses for Analysis 1 1 A. Schonnlwrria' = X. valga. Taxon abbreviations used were Xylocopoides' = X. cyanea. Xylocopa-=X. amethystina. sinensis. Koptortosoma= X. = X. bentoni. Nodula^ = X. Numbers across the top of matrix correspond to character numbers used in the text. caffra. Natural History Museum. and 3A. 4 and 4A are explained in the text. Koptortosoma^ = X. guatemalensis. violacca. The University of Kansas . Matrices tor Analy2A. Schonnherria^ = X.1100000111 0000002001 1100000111 0020000001 1100000111 0020000000 1100000110 0020000001 1100000111 0020000001 1100000111 0020000001 1100000110 0020000001 1100000110 0000002001 0101001112 0000002000 0101001112 0000012000 0100000100 0100002000 0210100100 0000000 1020000 1120000 1020000 1120100 1120100 1001000 1000000 1120000 1120000 1120000 1120000 0000110 0000100 0020000 1010100 0001010 1010000 1020000 1110000 0021000 1121000 1021000 1121000 1120010 0011010 1120110 0020010 1121100 1121000 1121000 1120000 1021000 1121000 1021000 0000000 1000011 1120100 1120000 1120000 1120000 0020000 0020000 0020000 0020000 0020000 0020000 0020000 0020000 1010100 1010100 0001010 1001001 . 3.Scientific Papers. Koptortosonm^ = X. micans. Xylocopa^ = X. californica arizonensis. pubescens. Character Taxon 123456789 0123456789 0123456789 0123456789 0123456789 0123456 000?00000 001000000 001000000 001001000 011000000 001000000 000000000 000000000 101001000 101001000 001111000 001111000 000100000 000100100 000000000 001100000 000100000 011000001 000000000 000000000 000000010 011000010 000001000 100001100 000001100 000000100 011000100 000001100 110001100 000001100 001001100 001000000 101000000 101000000 111000000 000000001 000000000 001000000 011000100 001100100 001000000 001000000 011001000 000101010 000000000 000000000 000000000 001000000 000000000 001000000 001100000 001100000 001000010 12 3 4 5 Outgroup Nanoxylocopa Cirroxylocopa Xylocopoda^ Dasyxylocopa Xylocopsis Xylocopmdes' Calloxylocopa Neoxylocopa Megaxylocopa Stenoxylocopa Xylocopina Ctenoxylocopa Copoxyla Rhysoxylocopa Gnathoxylocopa Xylocopa Nyctomelittn Zonohirsuta Prosopoxylocopa Bomboixylocopa Mimoxylocopa Alloxylocopa Xenoxylocopa Mesotrichia Pktynopoda Hoplitocopa Hoploxylocopa Afroxyiocopa Oxyxylocopa Cyaneoderes Cyphoxylocopa Koptortosoma' KoptortoscmaKoptorlosomn^ Pnxylocopa testis Schonnherria} Schonnherria^ Schonnherria^ Xylocospila Epixylocopa Perixylocopa Acroxylocopa Bihinn Apoxyhcopa Xylomelissa Nodula' NodulaNotoxylocopa' Notoxylocopa^ XylocopaXylocopoides^ 0000000000 0100000001 0100000001 0100000001 0100000001 0100000011 1210000101 0100000001 0110000001 0110000001 0110000001 0110000001 1100000111 1100000001 1101000201 1100000111 1210000101 0101000111 0210002001 0210000000 0210000000 0210000001 0210000000 0210000001 0210102000 0210100000 1210100000 0210112000 0210000001 0210010000 1210000000 0210002000 0210000000 0210100000 1210000000 0200000!!! 0101000001 0100001011 0100000011 0100000011 0100000001 1000000201 0000001201 0000001201 0000001000 0000001201 0001000201 0101000201 0110000201 0110000001 0210000101 1210000101 1210000101 00??000000 0100102000 0100101100 010010 1100 010010 1000 0100102000 1100004000 0100000000 0100103100 0100103100 0100100100 0100101000 0000002000 0100000000 0100000000 0100100000 0000001000 0100102000 0111002000 011100 1000 0100001000 0100115000 0111110000 010111! 010 0122115011 0122001001 012210! 001 012211! Oil 0123115100 012311! 010 012310! 010 012311! 010 010211! 010 0123111010 0123115010 1100111000 1100000000 0100102000 0100102000 0101102000 0100101000 0100012000 0100102000 0100112000 0100112100 0100102000 0100102000 0100111000 0111000000 0100110000 0100110000 0000001000 1100004000 0000000000 0000000070 0010002000 0101000111 0010000000 0101000110 0000000000 0101000100 0000002000 0101000112 0000002000 0101000111 0100002000 0210100100 0000002000 0210100100 0020000011 1101000100 0020000011 1101000100 0020002010 1101000100 0000000000 1101000100 0120002000 1100000100 0000002000 1000000111 0020000000 0100000110 0010002000 1100000111 0000002000 0110000100 0000000000 0101000111 0000000000 0100000000 0000000000 0100000101 0000010001 1000000000 0000012001 1100000000 0221111000 0100000000 1020011100 0000000000 1021211110 0110000100 1221211010 0000010100 1221211010 0000010100 1121211000 0000000000 1021211100 1000000102 1221111000 0000000101 1221211100 0000000101 1221111100 0000000101 1121211100 1000000102 1121211101 1000000101 1101211101 1000000102 0000000000 0:00000110 0000002000 0100000111 0220000000 0101000112 0220012000 0101000112 0220002000 0101000112 0020002000 0101000112 0020002001. .

X. Fig. State 2. related to the Xylocopini. Xyhcopa (Megax\/locopm)* fimbriata. which I had permission to dissect. Abbreviations. Hurd 1978. (0) Absent. Mandibular structures are described using the terminology of Michener and Fraser (1978). B redrawn from Hurd and Moure (1964). Numbers in parentheses are the codes used in the data matrix and on the cladograms. (Gnathoxylocopa) sicheli. tawny or ferruginous on at least dorsum is not found in the outgroup. terga and sterna. IB). be fused. a = acetabular groove. 1965) 1 metapostnotum instead of propodeal triangle. Number of teeth on apical margin of female mandible. down the lateral edges. but it is present among other species . the medial tooth is longest.— (0) Two (Fig. State 1 is not found in the outgroup flagellar — exemplars used here. and segments are numbered from anterior to postefollow Brothers (1976) in the use of rior. the Ceratinini. 1993) and analyzed with MacClade version 3. polarity decision is equivocal. supraocellar 8. the characters are given same number in the cladistic analysis and the group descriptions. correspond to the numbers in all the subsequent tables and trees and in the group descriptions. Figs. — (1) Upper rutellar tooth (0) and preapical tooth Separated by an emargination (i. State 1 is not found in the outgroup. lA). is considered the plesiomorphic state. if present. (1) much longer than width of flagellar segment 1 (Hurd and Moure.Subgenera and Genera of Carpenter Bees. 1963. B.2). Examination of the reservoirs of the mesosomal glands of the males required dissection of the posterior part of the thorax using techniques described this was limited to specimens for by Minckley (1994). and therefore not closely State 1 to be apomorphic 3. 18). 'Regarded as a member of the subgenus of Neoxylocopa herein. are shown in Figure cause the IC are coded as having two upper teeth are considered three teeth beto outgroup. 1963. 1. 1963. The cladograms were produced using the the Fig. 1988). of pollex). possibly suggesting that the additional 1 tooth of some Xylocopini has arisen de novo.05 (Maddison and Maddison. IC. in those outgroup taxa with three mandibular teeth (Ceratinini. In some ingroup taxa. present. but in the Xylocopini the ventral tooth (ventral tooth of the rutellum) teeth is longest. 1. Based on outgroup analysis the Malar space of female. 5. 16). X. External views of left mandibles of females: A. C. lA. (Stenoxycolopa) micheneri micheneri. For ready comparison. e. In the pits. except that I use medial (as opposed to lateral) instead of mediolongitudinal for structures on the tibial spine. (0) Absent (Fig. (1) — (0) Punctiform or not punctiform.1 (Swofford. This Subapical tooth or convexity on upper carina of male mandible. Fig. (1) continuously carinate and broadly impunctate (Hurd and Moure. SI. Terga and sterna are designated as T and S. — (1) 1 green. IB).1. S2. a condition — coded as 7. Xylocopini duced trees with Pwx\/Iocopn arising as the basal taxon in the tribe. a carina is present on the upper epistomal suture but it does not extend completely 1 — Subapical tooth on upper carina of female mandible. Twotoothed mandibles 6. fused and forming a continuous edge (Fig. deeply excavated and crater-like. IB). Fig. State 1 is widespread in bees and occurs in the genus Ctenoceratina (Ceratinini) of the outgroup. 51). Pararhophites and Parafidelia have two teeth. it is not likely to be an ancestral group in the Ceratinini — apomorphy is shared by Stenoxylocopa and Xi/locopina and is the same as the "inner tooth" of Hurd (1978). or pale. propodeum. (1) present. lA) were coded 0. (1) present (Fig. Scale = 1 mm. etc. See preceding character. IC). (0) Not raised or broadly impunctate (Hurd and Moure. I infer Epistomal suture of female. Supraocellar pits of female. and Manuelia. Allodapini and Lithiirge (Michener and Fraser. Characters and Coding The characters. sa = subapical tooth. State 1 does not occur in the outgroup. consider three homologous in this analysis and coded the groundapex plan state as equivocal. 1963. Figs. black. Unless noted otherwise. Because Ctenoceratina has a number of derived features (Daly.g. 17. Fig. respectively. 1993). 1978) have three teeth on the apical margin of the mandible. In the outgroup. as numbered below. Terminology generally follows that of Michener (1944. of body. (1) three (Fig. 1. 0. 15. Allodapini and Parafidelia). (0) dark brown. for the Xylocopini. computer program PAUP (Phylogenetic Analysis Using Parsimony) Version 3.e. Color of male vestiture. (Fig. 16). (0) Shorter than width of segment 1 (Hurd and Moure. and Hurd and Moure (1963).. 14. Mandibles as 4. and Michener (1975) and Reyes (1991) in the use of ventroapical plate for the ventral distal projection of the male gonocoxite. (Fig. Furthermore.

State 1 is not found in the outgroup. males of all taxa except Lithurge. (Ctenoxylocopa) fenestrata. Xylocopa sensu stricto). — (0) Yellow on at least i. In the outgroup the basitibial plate. 2B. 3A). As discussed in Character 17. Proxy locopa*riifa. X. 11. 1975. State 1 oc- not occur in the outgroup. — Not modified. Reyes. Xylocopa (Xyhcopoides) cyanea. B. Fig. In all ingroup taxa with well-developed basitibial plates. Number of spines on outer apex of male hind (1) two (Hurd and Moure. en- character 17. — (0) Not modi- fied (Fig. male hind femur. (0) Weakly developed or rising little from surface of leg. character . modifications such as these are known only in some Ceratina (Hirashima. 3B. 37). apex of tibia (Fig. 2B). 2C). but extending to. or tubercles. Those taxa with a single row of tubercles (State 2) on the posterior edge completely lack a carina. All ocelli. (0) Originating at tibial apex and lacking medial carina (Fig. (1) enlarged. E. tibia. State 1 does — Ocelli. in the outgroup that I did not consider (Michener. 3C). 3C). This is a highly variable character in the Apoidea but in the outgroup. 14. (. X. — Anterior and posterior edges of female basitibial plates. This transformation series was coded as additive. 2D. Natural History Museum. 1963.— (0) Number of spines on outer apex of female hind One (Fig. 2B. The University of Kansas ts Fig. In the outgroup. Figs. C. (1) Apex of male basitibial plate. (1) with numerous tubercles on both edges (Fig. The transformation series was coded as additive. 2A. 2A.— (0) None. 18. (2) with numerous tubercles only on posterior edge. State 1 is roughly triangular in ventral view not found in the outgroup. •Regarded as subgenus of Xylocopa herein. testis* bomhylans. was coded as nonadditive. Mid and hind trochanter of male. 38). — . 15.. 2D. the apex of the structure is bifid and usually strongly so. 3B). one. (1 ) — some part of face. and Proxylocopa (sensu outgroup taxa have normal-sized 10.Scientific Papers. State 1 is unknown in the outgroup. bifid. curs in the subgenera Nyctomelitta stricto) of Xylocopa. tibia. when present. 1963. Females of the outgroup 13. invariably has an entire apex. Male facial maculations. (1) without yellow markings. The character was coded with an entire apex as the plesiomorphic condition. This 16. 9. State 1 is found only in taxa with one outer tibial spine. (2) 12. on the anterior edge.Alloxylocopa) circumvolans. (1) ventral surface of (0) (0) Spines abeach with spine. (0) Entire or absent.e. Outgroup taxa lack these spines. 2C. have yellow maculations on the clypeus. (0) Not modiwith basal tubercle (Fig. (1) welldeveloped and originating above. (1) (Fig. 2.g. tirely black. Hurd ) and Moure. sent. The polarity proposed here may be wrong because those ingroup taxa with the most strongly developed basitibial plates (a plesiomorphic condition?) have tubercles along both edges (e. Abbreviation: ts = tibial spines. the polarity of this — Shape of female hind trochanter. 2E). Tibial spine of female. 2A. 1969). Outer view of hind a tibiae of females: A. Scale = 1 mm. 2E). 36. may be the reverse of that presented here. (2) with spine on basal half (Fig. have one or two spines. 1993). D. (1 two (Fig. Ventral surface of fied. Ctenoxylocopa.

Anterior (Afroxylocofm)* nigrita. 27. when present. Male metapostnotum. Beginning of posterior thoracic declivity of fe(0) Profile smoothly rounded. 4A.Subgenera and Genera of Carpenter Bees. 33. See preceding character. Figs. (1) abruptly rounded (Hurd and Moure. Smoothly rounded. position often undefined (Hurd and Moure. 22. Apparently the metapostnotum can be lost in two ways: (1) absence of unique sculpturing and marginal lines. 33. in species 19. but 1 have not considered these 1. 3E). 20. Minckley 1994. 1983. or (2) by an increase in the size of the propodeum relative to the metapostnotum — until the latter is completely absent externally. 1963. 3A. The 24. 2D. In the Ceratinini. 2B. 39. does not overhang metanotum (Hurd and Moure. (1) abruptly rounded (Hurd and Moure. pletely close considered as a to (e. Anterior A. (2) as a transverse carina that Minckley 1994. IC. [Notoxylocopn] guatcmnleiisis). State mesosoma accommodates the mesosomal gland reservoir. Apex of female basitibial plate. 4C. Because states and 1 both occur in the outgroup. be homologous to State 21. Fig. Figs. metapostnotum in bees is highly variable in size and shape and. 27. Fig. 1963. {Ctenoxylocopa) hottentota. X. IC. 2E). Fig. 39. I consider the ground-plan state to be and 3 are unknown in the outgroup apomorphies within Xylocopa. 1994). is distinguished from the propodeum by a change in the sculpturing of the cuticle and /or by a line along its edge with the propodeum. 2E). IE). Figs. Fig. Extending to basal reduced. See discussion of Character 17. Fig. (0) — See discussion in following character. plate. Minckley. Tr = trochanter. 2C). 5E). The metapostnotum in some taxa is well developed and in Xylocopini the metapostnotum invariably other taxa that is vestigial or absent. 2C. (2) sharply angled (Hurd and Moure. 1963. 5A). beginning undefined (Hurd and Moure. 3E). Modified from Eardley. Minckley 1994. Ventral view of the hind trochanter of female Xylocopa view of the femur of the male of X. the . In the a line along the Fig. Minckley. Female basitibial (Fig. foveate and Fig. but do not com. Abbreviations: is delimited by edge with the propodeum and a continuous size-gradation across taxa is evident. 24. 2A. 1963. t = tubercle. Xylocopini 1 is unique to Xi/locopoidcs and Lesfis Other taxa in Xylocopa have lateral edges that converge basally. and no the males of which have an enlarged posterior on apical 3/4 of tibia 2A. fourth of tibia (Fig. *Regarded as a member of the subgenus Koptortoscma herein. Hurd. 1 A. 28. 1963. [Xi/lomelissn] erytliriiin. 2C. 1 A. view of the femur of the male of X. — 1994. These observations suggest that loss of the metapostnotum in the Xylocopini is by a change in the size of the propodeum. male. (1) absent. s = spine. 1994. C. 4C. Figs. (0) Present. Beginning of posterior thoracic declivity of male. 3. ) if closed. 1994. 4E). States 2 formation series was treated as additive. 5C. Minckley. Fig. 23. — (0) Entire (Fig. Fig. 4A. Fig. 4E. Figs. (1) bifid (Fig. This transand are strong equivocal. B. 34. Fig. 1963. 2B). 5A). F = femur. 3C. gradation in size of the propodeum is apparent. 9).g. 3 A. the presence of the metapostnotum is usually made evident by a change in sculpturing. a contiition that could be weak fovea. X. — (0) Open (1 basally or. 3C. (Mesotrichia) latipes. extending onto basal fourth of tibia. 1961. Female basitibial (1) plate. the metapostnotum typically is absent (Minckley. 1963. Minckley 1994. Scale = 1 mm. Furthermore. (3) marked by transverse carina that overhangs entire metanotum (Hurd and Moure. not extending to base (0) — of tibia (Fig 2A.

Fig. Abbreviations: S = scutum. (0) usually abutting in the middle (Minckley. IC). lA. 5E). Shape of the gradulus on Tl. conI sider the presence of a defined metapostnotum to be the T2 and T3. (1) absent. 1994. — Anterior to the 28. (0) Absent. Figs. 43). 4A. 29-34. 42). Female metapostnotum. — Distribution of graduli on female terga. 1963. 24. (f) two widely separated pouches (Minckley. character 32. *Regarded as a member of the subgenus Mesotrkiiin herein. 35. (0) Not modified. 27). forming a spine. 1963. 4C. (1 dorsally elongated. Fig. The presence of a reservoir is a unique character of most Xylocopini. which has a gradulus only on Tl (Hirashima. This character was coded as nonadditive. 31. This apomorphy is unknown in the outgroup. — (0) Linear. Scale = mm. (2) single layer of tubules. Propodeum at its connection with metasoma. 5. (5) numerous. (2) Tl only. 26. Fig. 4D). Fig. 1994. although.10 Scientific Papers. metapostnotum is either well developed or completely absent. (1) a — — Absent. as in Koptortosoma (Minckley. State 1 is not found in the outgroup. 3 A. Fig. . (0) Tl and T2. Figs. (3) highly convoluted tubules. Fig. 5B). Fig. 1963. 41). 23. Figs. (1) — (0) Tl. Minckley. ) — 29. in the Xylocopini. (2) present on Tl only. (0) Present. Figs. 27. IC. 1994. State is not found in the outgroup. or more. 3B). In these difficult taxa I have opted to preserve information and have split the characters into a number of often autapomorphic states rather than simply scoring the character into states of presence or absence. — Modifications of anterior surface of Tl of males. deeply sulcate (Hurd and Moure. 1963. 33. Fig.— (0) Not modified 1 (Fig. IF. 1994. 1944). Minckley. Eardley. 5F). 1963. 1963. The reservoir in at least Modifications of anterior surface of Tl of female. See discus- sion of preceding character. 4A). 44). except for the ceratinine genus Pithitis. Beginning of the thoracic declivity of male. highly convoluted tubules. — (0) present Left tegula of the male of (A) Xi/locopn vaiipuiicta and (B) X. Absent. All members of the outgroup have graduli on at least T1-T3. 5 A. Neither a fovea nor an invaginated mite pouch occurs in other bees. 1 infer from this pattern that in the Ceratinini the metapostnotum is lost by a reduction in its surface ciifferentiation from the propodeum and not by a change in size of the propodeum or the metapostnotum. (Platynopodn)* tenuisaipw. as in Neoxyhcopn (Minckley. 30. Distribution of graduli on male terga. was treated as additive. ground-plan condition 25. The transformation series was coded as additive. 3C. 1969). 5C. The on Tl. 1994). See discussion in preceding character. outgroup. Fig. 1983. There reservoir is tremendous variation in such features as the shape of the tubules that make up the gland and the size of the reservoir (Minckley. IE. (1) elon- gate posteriorly (Fig. the inferred ground-plan state for the ingroup is graduli on all terga posterior to T3. State 1 is not found in the outgroup. including those where it is minute. IB). lA. the medial groove is linear. In the a sex-attractant pheromone. Medial groove of Tl of female. Reservoir of the male mesosomal gland. Fig. (1) on the propodeum (Hurd and Moure. Based on this information and the argument that a gradulus on each tergum is generally considered to be plesiomorphic in bees (Michener. 1994. Male tegula. (1) some species is associated with a gland that produces in the vast majority Fig. the function for this structure has not been examined. 1 Tg = tegula. 27. 4B). 1994. See discussion in following character 34. 4. Fig. (4) single medial pouch with one opening (Minckley. fovea (Hurd and Moure. The University of Kansas of large carpenter bees. 1994. as well as on more anterior terga. 1963. (1) present on Tl and T2. (1) transverse (Hurd and Moure. (2) with an entrance to a acarinarium (Fig. or more. (Da fovea (Hurd and Moure. 42). T2 and T3. (0) propodeum (Hurd and Moure. which makes homology assessment difficult for some taxa. Therefore. Natural History Museum. — (0) Gradulus of Tl con- tinued posteriorly adjacent to lateral margin of metasoma (Hurd and Moure. ID. 5E. 4B. 3D.

This decision is based on the two features not found in other taxa with such gonostyli. 6. testis (Fig. (0) Gonostylus of male genitalia. 8C. (1) strongly developed. flanked by one or spines of many preapical spines (prepygidial 1963). temiiscapm. (0) Not armed with lateral preapical spines (Fig. 1963. 5. 1964. This apomorphy sented by a raised spine that originates before. 9A. sp = subgenus Mesotrichia herein. Scale = 1 mm. on metasomal sternum of female. 7C). arrow indicates anterior. forming a point. (0) Carina not present on all sterna or not continuous. — g = gradulus. The pygidial plate of all wood-nesting is Apex of penis valve. Medial carina 39. Lateral 0. In the ingroup. Distinct dorsal to one or patch of appressed short setae anterior and more metasomal spiracles of male. and Maniielia. (1) protuberance or carina present (Fig. 4). In Ceratinini is and Allodapini. In most taxa these are found on T2. This character was coded as nonadditive. 9E. 9D. At its base. the distal edge of T6. (0) Paired (1) paired setal patches present on at — least one sternum (Fig. — (0) Smooth. A well-defined medial carina on all female sterna clearly is apomorphic for the ingroup. 8A. Apical margin of SI. setal patches absent. not produced. 9C. keel. — (0) Not hirsute (Fig. 38. jection fused to Pregradular areas on sterna of male. although in some outgroup taxa (Ceraiina laeta. B. Fig. a carina is never developed into a strong keel. This character was coded as additive. Xylocopa (Neoxylocopa) varipimcta male: A. Absent. a member present on S6 but is of the outgroup. (1) hirsute (Hurd. spines (Fig. — — gonostylus similar to state 43. . State 1 is not found in the outgroup. 8A. Tl = dorsal surface of tergum as a spiracle. State 1 is found in the spine of T6 of female. Figs. nipestris and Megaceratina sculpturata) a weakly developed medial carina occurs on S6 and in many members of the ingroup. 1. 1975). 9A. C. (0) Absent or not developed into a keel. without a carina (Fig. of S4 Medial carina on S6 of male. 1961. Pregradular area showing patches of appressed setae.T5. 6A). (1) 36. Pygidial — outgroup only in some members 0. Xylocopa (excluding Proxylocopa). but in some taxa they are on fewer terga. (2) with many preapical lateral 41. 9C. Side view of T4 with appressed setal patch. produced (Hurd and Moure. (2) medially emarginate (Hurd and Moure. 88. 8A. g = gradulus. a carina occurs on all sterna. SI = sternum 1. The ground-plan state for the ingroup inferred to be State 44. 7A). 9E. 3. (1) continuous and present on all sterna. the spine may be outgroup. Xylocopini 11 Fig. abbreviations: sp = spiracle. and extends unites Xylocopoides and Cnlloxi/Iocopa in the and does not occur beyond. 1963. 7B). 242. these most often are on S2-S6. — Fig. forming a 37. a developed spine absent although the distal edge of T6 medially elongated. repre- 8C). 9F). 8B. 9F). ^Regarded member of the (0) Entire. All outgroup taxa have a distinct slender 42. (0) A slender proapex of gonocoxite (Figs. 8C). State 1 occurs in all outgroup taxa. edge of penis valve.Subgenera and Genera of Carpenter Bees. 44). 43). State 1 is not found in the outgroup. Abbre\'iations: a = entrance to the acarinarium. 40. 244). Fig. 9B) was coded as having State because its conical gonostylus is different from that coded as 1. 9B. (1) with pair of preapical lateral spines (Fig. When there are such spines they delimit a triangular area that many may be a remnant of is a pygidial plate. although it is weakly developed or not continuous. (1) present (Fig. Anterior face of Tl of a female Xi/locopn (Plnlyiwpodn)* 1. In Ceratina. 6B). a dense brush of medially oriented hairs on the gonostylus and a strong carina on the apex of the gonostylus. Figs. 9B. 9D. — Hurd and Moure. 8B. (1) a rounded or conical setose area at apex of gonocoxite (Hurd and Moure. of the Allodapini is (Michener. but in many taxa they occur on fewer sterna.

(Notoxylocopa) tabmuformis (modified from Hurd and Moure. Figs. 1975. 8A.e. lateral. elonand erect. Abbreviations: Ic = lateral carina of penis valve. 136). X. 9C. Nonetheless.— (0) Reduced or absent 8A. Medial area of posteroventral margin of some Allodapula (Allodapini). 1963. well developed (Hurd and Moure.12 Scientific Papers. 8B. (0) Lacking a dense patch of setae on its apex (Fig. 9E). slender and less than 1/2 as long as gonocoxite (Figs. (2) present. 9C. or entirely fused with gonocoxite and represented only by the rounded or conical setose area at apex of gonocoxite (Character 42-1). 1 is an autapomorphy for Notoxylocopa. 8B. 9A. i. 8C). This condition was coded as 0. 7. Figs. 9D). 9D. 8A. 1963. Dorsal. 8C. and Compsomelissa (Allodapini) and 48. slender gonostylus with numerous setae is considered to be plesiomorphic for bees. ing continuously from — absence of such a structure. Many ingroup taxa have a spine along the ventral margin of the base of the gonocoxite that is more lateral than the condition defined here. Reyes. 49. State characterizes all outgroup taxa. B. 9D). X. 9E. This character is variable in the outgroup.— (0) Absent (Figs. An elongate. over 1/2 as long as gonocoxite (Fig. The outgroup is variable in this respect.. 8B. — (0) Entirely membranous. C. (0) Divergat. (1) 1 posterior projection (Figs. but in each tribe some member has this plesiomorphic condition. (1) present (Fig. 50. 51. 8B. 9F). (Fig. Spatha. 9C).Xylocopoides) californica. X. preapical spine. Ventroapical plate of gonocoxite. the Xylocopini is possibly comparable to that found in the The apex of the gonostylus is entire in all outgroup taxa. (1) present. 134). (. ml = medial lobe of gonocoxite. Although a many groups of bees. 9A. Allodapini (Michener. member of the subgenus Schonnherria Parafidelia and Lithurge) and one member of the Manueliini (Manuelia gayatina). but not carinate and not (2) — well defined (Hurd and Moure. 53. 1991). 9C. 1963): A. X. 9B). (Neoxylocopti) lunipimcta. B. — . 9 A. 9B. in the outgroup. 9F). and ventral views of male genitalia (modi- fied from Hurd and Moure. The ground-plan state has been inferred based on State being a consistent feature of the Ceratinini and Megachilidae (i. iCirroxyloccpn) vestita. *Regarded as a herein. State is with not found 52. Dorsal surface of T6 of females: A. or near. 9A. 9D). 8A. 8. State spatha is present in 47. (0) Absent (Fig. (0) Absent. Parafidelia (Fideliini). C. Xi/lccopm (Xyhcospila)* bnmbusae. 9B. Projection or spine on inner basal margin of under surface of gonocoxite.. (1) roughly parallel from ventral gonocoxal bridge and not diverging until well above ventral gonocoxal bridge (Figs. Abbreviations: flnvorufn. Pararhophites (Fararhophitini). Manuelia (Manueliini). it is not so well developed in the outgroup. 9B.e. Euceratina (Ceratinini). v = ventroapical plate. 8C. (0) Slender.Mcsotrichia) 1963). Gonostylus of male genitalia. carinate (Figs. (. 8C). Fig. 9C). Penis. 135. 9B). (Fig. 8B. 9E. Pararhophites. p = pygidial spine. the inferred ground-plan state of the outgroup is Ventromedial margins of gonocoxites. 8C. (1) present. 9F). 9D). Natural History Museum. The University of Kansas sa Fig. The ventroapical plate of weU developed 8A). (1) sclerotized basally 46. 131. gate (1) Gonostylus of male genitalia. X. well developed (Fig. 8C. — SA. 9B. ventral gonocoxal bridge (Fig. No such projection is found in the outgroup. 8A. State 1 is not found in the outgroup. not — gonocoxite. 8C. Apex of gonostylus with medially projecting lobe. — (0) Not produced (Figs. 9C. (1) with dense apical patch of setae (Figs. 8B. (1) on venter. sa = 45.

X. For ease of comparison. X. B. For each analysis the matrix was first analyzed with all characters weighted equally and a second time using successive approximation weighting (Farris. explained by Fitzhugh. testis* aeratus. X. Coding multistate characters in the latter way can result in assignment of an average weight of all states. version 3. Carpenter (1988) provided an example of how these different coding schemes can affect tree topologies. Abbreviations: i swap on all starting were nonminimal. 8C. (Bomboixylocopa) rufipes. iXylocopa) mlga.01 (Swofford. both characters that perform poorly (show high homoplasy) receive more weight and characters that perform well (show low homoplasy) receive a lower weight than they would otherwise. 9D). E. I followed the weighting scheme used by the Hennig86 parsimony program (Farris. I searched by two methods in each analysis. The weighting scheme is applied repeatedly until weights of all characters stabilize. which assigns weights to characters from to 10. if coded in additive binary form. 8B. Before each successive approximation analysis the characters are coded in additive binary form to insure that the weights assigned to states of multistate characters are correct. 9. 8A. ml= medial lobe. Phylogenetic Analyses Four separate sets of analyses (described below) were made. ma = medial apodeme. 1989). (Clenoxylocopa) fenestmta. (Proxi/locopa) rufa. D. In the first search the following settings were used: Starting trees by stepwise addition. addition sequence F Pwxylocopa* = internal medial projection. PAUP provides many options for calculating weights for successive approximations. *Regarded as a subgenus of Xylocopa herein. 1988). slender and parallel (Fig. 1988. which simple. (1 ) — (0) Apically expanded (Figs. State 1 is not found in the outgroup. The second search used the random addition sequence option. The large size of the matrix restricted the search for the shortest trees to heuristic searches in the PAUP program Fig. Though some outgroup taxa have relatively slender penis valves they are not as slender and parallel-sided as in State 1. Because these types of searches cannot guarantee finding the most parsimonious tree for the data. Posterior 13 margin of male T7. my results may differ from those obtained from Hennig86. Ventral views of the right sides of male genitalia: A. Xylocopini 54.Subgenera and Genera of Carpenter Bees. Penis valves. (Prosopoxylocopa) mirabilis. (1) with ma pair of dentiform projections. so I have tentatively inferred this condition to be apomorphic in the Xylocopini. hold 75 trees at each step and trees including trees that . Nevertheless. The alternative coding method for such characters is with all states in one column. — (0) Entire. 1990). Scale = 1 mm. Successive approximation is an a posteriori character weighting procedure that weights each character based on its best performance (= fewest steps) in the previous analysis. 55. C. 9A. X. 9C). because some characters coded as polymorphic in the PAUP data matrix would have to be considered missing in a Hennig86 format (Hennig86 does not accept polymorphic characters) and might be optimized differently or assigned different weights.

successive approximations. successive approximations. statis- Analysis 2A. 1991. For those trees re- sulting from successive approximations weighting.14 Scientific Papers. in Analysis 3. 1988. 2A. The four analyses were undertaken certainties in was done by applying successive approximations 2. Results from these four analyses after successive approximations weighting are followed by the letter A (e. These considerations emphasize that the results of this study are only hypotheses. it seems reasonable that the The influence on tree topologies of strongest hypotheses for relationships are those that were missing (Analyses 1. This analysis was done with Proxylocopa excluded because I had coded some characters as missing for this taxon in Analysis 1 Other than the lack of Proxylocopa. 3 and 4. coding characters as characters coded as nonadditive (unordered) to have the least assumptions. The University of Kansas sis begins searches from trees produced at random by the program. Consensus trees are not accompa- nied with character changes because of problems with Successive approximation weighting has been investigated who used a hypothetical data set with complete character independence and a known phylogeny. Additive coding of character states (Analyses 1. 1992). Topologies largely redundant with those from other analystrict dogram and ses are not presented. 1993. This analyAnalysis lA. 2A etc. the tribes of the Xylocopinae might have polarity assessment of the character states used in this some characters were coded as additive and oth- and ground-plan states for all characters were inferred by assuming Manueliini to be the sister group to Xylocopini (Ceratinini + AUodapini) in the subers as nonadditive). —This analysis was done on the to investigate the role that the uncertain relationships among study. 1985). successive approximations. removing them from the analysis. Natural History Museum. Because islands of equally parsimonious trees were found in Analysis 3. 1 A. characters weighted equally and . For example.g. 1 consider Analysis 3 that had all taxa included.I. This result was because the weight- ing algorithm assigns weights of zero to effectively many characters. to the reduced matrix used in Analysis to investigate unassumptions about the evolutionary changes or homology assessment of characters.e.. Platnick et al. Analysis 1 A. 4. and for weighted tics characters. = retention index and TL. However. used the same matrix as in Analysis 1 with successive approximations character weighting applied. — Analysis 2. For brevity. Assumptions about character ordering were as described in the Character Analysis section (i. This analysis was done using successive approximation weighting on the matrix used in Analysis 4. Five hundred replications w^ere run and set to swap on no more than 30 trees at each step up to and including the fewest steps obtained by the first search. 1). this was the same matrix and character assumptions as used in Analysis 1. . Successive approximation weighting produced more trees than corresponding analyses with all characters weighted equally in all analycharacter optimization ses. or of deleting taxa altogether (Analyses 2 and 2A) remains insufficiently understood (Gauthier et al. 4A) assumes that an a priori knowledge of the evolutionary direction of character change is correct. equal weighting. — Analysis 4. equal weighting. an approach that has been questioned unless ontological data are available (Nelson. — The following abbreviations trees from each analysis: C. -i- Ground-plan states of all characters were determined with the Xylocopini as the sister group to Manueliini -I- (Ceratinini + AUodapini).). Successive approximation weighting was also highly sensitive to the topologies of the initial trees. the four analyses are referred to hereafter as Analysis 1. equal weighting. 4A). sis change of the hypothesized groundplan state ters 25 and 36 from to polymorphic (0. 3A. lA. This analywas done using successive approximation weighting on the matrix used in Analysis 3. sis — — Selection of Taxa. 2. This reanalysis resulted in a for charac- family Xylocopinae. 1995). are used in describing the = consistency index. six analyses were run starting with each of the six islands produced in Analysis 3. = treelength. successive weights. some nodes were left with no character support. 1 discuss each analysis and show one fully resolved cla- consensus trees for unit]ue topologies. except Analysis 2. Hence. 4. —This analy- are given for equally weighted characters to facilitate comparisons with the other analyses.l. Weins and Reeder. Maddison. 3. Analysis 1. equal weighting. This analysis was done using the same matrix as in Analysis 1 but with all characters coded as unordered. 2. three sets of relationships were generated even though the and Coding section). This analysis was based on the full matrix (Table 3). R. and of the relationships of the outgroups discussed above in the section on Analysis 3. Because of reservations concerning the transformation order for several multistate characters (discussed in the Charonly by Farris (1969) acter List when polytomies are present (Maddison and Maddison. and should be treated as such. — Analysis 4A. Analysis 3A. repeatedly supported under different assumptions. RESULTS The eight different analyses the effect that various assumptions have were attempts to explore on tree topology...

in the text Sclioiuiherria. cnffm dently six times in the tribe. (Xylocopa) violacea and X.SlIBGF. and Xylomelissa. Epixylocopm. are further divided into subgroups. in this analysis The Ethiopian and group these New World groups are large and subgroup consisting of Notoxylocopa. Mesotrichia. Fig. the first two exemplars were groupeci together consistently. (Koptortosoma) sinensis and Afroxylocopa. Cyphoxylocopa. be more basal to best consid- phyletic. have given each an infor- mal name. Schonnherria. the emarginate shape of SI. X. Rhysoxylocopa (sensu stricto). Nanoxylocopa. Both characters are lost and gained repeatedly in the tribe and furthermore are functionally correlated. Xenoxylocopa. weighting iteration began with a set of equally parsimonious trees from the same data set.L. Hoploxylocopa. For all other subgenera and genera of the tribe this analysis a single hypothesis of relationships. Together. The monophyly of all members of this clade is supported by Character 36. Nanoxylocopa. For the Ethiopian are. Koptortosoma. The sister group group plus the large group of New World taxa (including Gnathoxylocopa and Nyctomelitta in this topology) is supported by the parallel ventromedial margin of the gonocoxites (Character 50. —Manueliini is basal in the subfamily. If this weighting approach could reliably distinguish characters that were truly homologous from those that were homoplasious. Oxyxylocopa. (1) the Mesotrichia and Xylocospila) but with two Old World members (Gnathoxylocopa and Nyctomelitta). Bilwm. Stenoxylocopa. Mesotrichia. This clade and the next more-derived branch with the species of Koptortosoma varied among analyses. Neoxylocopa. Hoplitocopa. three groups were shown to be monoI (Minckley. use names of all exemplars in the discussion and on . (Xylocopa) valga). thereby suggesting little confi- and and designate X. Noduin (both exemplar species). Neoxylocopa. although it becomes rounded (Character 52-1) in the Rhysoxylocopa group. Those taxa defined that are derived relative to Proxylocopa are Afwxylocopa. (Fig. Hoploxylocopa. T. Xylocopoda.27. Xylocopsis. and Zoiioliirsiita. Thus. Tliis character is highly homoplasious and appears indepen- cludes the Australian phyletic in all analyses. whereas the relationship of the latter species to the other X. 1 refer to those subgenera represented by more than one exemplar by their subgeneric name if the exemplars were shown to be monohi the following text shown to be monophyletic. Three major clades are included herein. (2) the New World group consisting of Cirroxylocopa. and elaboration of the reservoir of the male mesosomal gland from two small pouches to a series of tubules (Character 26-2). Dasyxylocopa. some characters were coded as additive and homoplasy in my analyses. Prosopoxylocopa. = 288. by possession of a carina on the venter of the gonocoxite (Character 52-2). subgroup consisting of and Few characters separate these main groups. Cirroxylocopa. Xylocopoides. species and Koptortosonia For the first two groups. Stenoxylocopa. presence of the is component taxa within these larger clades. AUoxylocopa. Proxylocopa is 15 included. equal weighting. Xylocopsis. Xylocopoda. Bomboixylocopa Cyaneoderes. The most basal group consists of a biogeographically diverse set of taxa that inLestis as the sister group to the North American subgenera Calloxylocopa and Xylocopoides. yet several of these features probably are unique within the Xylocopini Afroxylocopa.l. but each refers to 3). and (3) the Rhysoxylocopa group consisting of Acroxylocopa. a greater level of congruence should be obtained in the weighted trees than was found here. I restrict use of Koptortosomn to X. XyLOCOPINI others as nonadditive. Megaxylocopa. Development of this carina is a strong feature that arises once in this topology. Monophyly was I not consistently supported for the exemplars of Nodiiln. and (2) the Koptortosoma and Platynopoda. = This analysis yielded eight = 0. Relationships In often associated with absence of the mesosomal gland metapostnotum it is among Larger Clades of the Xylocopini most analyses. R. lost in Platynopoda. pubescens. Platynopoda. produced clades are Four major and cladograms. and Noton/locopa multiple exemplar species (Table Xylocopa. for ease of discussion. A thorough study on how multiple islands of trees influence results using successive weighting is beyond the scope of the present paper.) Differences in the topology of 0. the eight trees minimum produced by this analysis were among Cyaneoderes and Cyphoxylocopa and their relationship to Koptortosomn. I refer to a Neoxylocopa subgroup which includes Megaxylocopa. Hoplitocopa. and Bomboixylocopa) and show httle relationship of the Ethiopian . Hoploxylocopa. and Xylocopina. the relationship of Proxylocopa to and the taxa shown ered as unresolved. Cyaneoderes. As such. Apoxylocopa. lOA). and Xylocospila. The following section is a summary of the relationships among the major groups in the tribe. these taxa comprise a sister taxon to the Old World subgenera Copoxyln + (Ctenoxylocopa + X. 1994). are used.I. length trees (C. Analysis 1. Oxyxylocopn. dence should be placed in this grouping. 9B. the Rhysoxylocopa group is basal and the sister group to a clade that includes the Ethiopian group and a clade of taxa that occur primarily in the New World (the Neoxylocopa group. the cladograms. Dasyxylocopa. Perixylocopa.67. Cyphoxylocopa.NERA AND GeNERA OF CARPENTER BeES. In the New World group. Koptortosoma. This is followed by a section discussing the relationships of the genus Proxylocopa are separated by the absence of the male metapostnotum (Character 24-1) and presence of a male mesosomal gland (Character 26-1). Mimoxylocopa. Sclwnuherria. sinensis separately. These are: (1) the Ethiopian group consisting of .

Natural History Museum. The University of Kansas <u .16 Scientific Papers.

In the resolved tree and in the consensus tree. 13B. This analysis yielded 89 minimum length trees (unweighted statistics. Fig. lost in Boiiiboix\/locopn a and derived loss of slender elongate gonostvlus (Character 47-0. 9C. Zonohirsuta + Prosopoxylocopa are part of a polytomy that includes the remainder of the Ethiopian group and other taxa. 13A.27. = 0.I. —Manueliini 15). and is in group. The subapical mandibular tooth is a derived loss in Gnathoxylocopa and is present in groups outside this clade. equal weighting. 4. After five weighting iterations. This analythree groups. = 0.I. Island 5 = 72.25. 5. R. = 0. The topologies of Islands 1 (Fig. C. and more polytomies exist in the consensus trees resulting from these analyses. Proxylocopa included (Figs. (Fig. Zo)iohirsuta + Prosopoxylocopa are a sister group to Notoxylocopa + Proxylocopa and the remainder of the Ethiopian group. wherein the large clades were much the same as those in Analyses 1 and lA. successive basal in the subfamily. 13A) and 6 (Fig. R. This placement of Gnathoxylocopa is the subfamily. 11 A. this analysis recovered the same major groups as in Analysis 1 The monophyletic group positioned at the base of the tribe in Analysis 1 is represented as a grade of five sequential clades along the . groups in Islands 1. 11 A). or "is- produced 48 divided among lands. TL. is associated with the basal lineage of all more derived member of the clade. C. Island 4 = 72. 5. lost repeatedly). Xylocopini and possession of a process or spine 17 on the venter of the gonocoxite (Character 51. These analyses raise the possibility that Zonohirsuta and Prosopoxylocopa are not sister taxa to the rest the Ethiopian clade. Strict minus Zonohirsuta and Prosopoxylocopa.I. The relationships among members within these three major groups are unresolved in the consensus tree. C. weighted statistics T. C. perhaps they are parts of a consensus trees of each of the three islands differ from those of Analysis 1 most notably in the relationship of Biluna to the other members of the Xylomelissa group.49.70). = 0. (Xylocopoides) cyanea). = 282. lost in Calloxylocopa Analysis Manueliini is 2A. Coding all unlikely (ciiscussed below). lOB). were not included with the remainder of the Ethiopian group. All islands of trees converged on the same consensus tree. = 331. = 0. IIB. unordered resulted in 408 trees distributed across 6 islands (T.I. Miiiioxi/locopn).86). —Manueliini is is basal in the subfamily. the two basal taxa. = 362. In the three islands in which monophyly for this group was not supported. = 297. In Island 3 (not shown). Except for the most basal clade. some characters were coded trees as additive and others as nonadditive. = 0. the Etliiopian group plus the Rhysoxylocopa group and the New World assemblage are a single clade. successive approximations. begin- ning of the thoracic declivity abruptly rounded in the fe- male (Character 23-1. Recognition of the major clades was the same as those produced in Analysis 1 Because of the similarity of results to Island 2 Biluna a . = 0. Together. The numbers of trees for each island are: Island 1 = 24. is basal in char- on the mandible of the female (Character 3-1) and a lateral projection on the penis valve (Character 43-1). this clade is the sister group to the remainder of the Ethiopian clade. Proxylocopa Xylocopoides.L. Successive approximations were applied to the matrix used in Analysis 1 A (Fig.l.64. 6. IIA. Proxylocopa sis excluded. These characters are found in other groups. approximations. 3. some characters were and X. 13A.65). All of these taxa and the Ethiopian group have two spines on the hind tibia of the male (Character 11-2) and female (Character 12-1. Analysis is (Notoxylocopa + Giiathox\/locopa) + NyctoDiclitta of a subapical tooth defined 3.L. Proxylocopa is excluded." of 16 trees each (T.65. the results of this analysis are not shown as a cladogram. C.92).23. derived loss of the metapostnotum in the male (Character 24-0) and female (Character 25-0). 15) require . successive approximations were applied.Subgenera and Genera of Carpenter Bees. more inclusive group Island 3 places Rhysoxylocopa (sensu stricto) as a sister that taken together are basal to the rest of the Ethiopian group to the remainder of the Rhysoxylocopa group. Island 6 = 24. = 511. Monophvlv of the Ethiopian group is supported by the possession of two spines on the male hind tibia (Character 11-2) and female (Character 12-1. R. Analysis 2. weighted statistics.I. Notoxylocopa. IIB) placed Zonohirsuta + Prosopoxylocopa as the sister group to the clade base of the cladogram in this analysis. Hypotheses of relationships among members within these larger clades were + Xylocopoides) based on the more numerous than in the equally weighted analysis.L. = 0. regained in the Mcsotricliia and Koptortosonw groups).28. R. = 0. Fig. lost in Afroxylocopn).I.I. = 0. character weights stabilized and produced 25 minimum length trees (unweighted statistics T. — sus trees of all trees in each of these islands generally are not well resolved but found.L.67.I. and sparse setae on the gonostylus (Character 48). Island 2 = 72. Island 3 = 144.l. = 298.L. Taxa supported as a monophyletic group in Analysis 1 form a basal grade of taxa in this analysis. Consenacters Analysis lA. T. but the congruence of characters is support for the mono- phvly of this clade. of the Nev^' The clade composed by the presence World group and is coded as additive and others as nonadditive. all characters were unordered. Zonohirsuta and Prosopoxylocopa. R. and Calloxylocopa are basal to the Ethiopian those in Analysis 1. The Ethiopian group was some unique relationships were shown as monophyletic in three of six islands (Islands 2. In Island 6 (Fig. C. equal weighting. 13B).I. the Ethiopian group in islands. R.I. Xylocopia. Island of Xylocopa + (Calloxylocopa 1 (Fig. 5. = 0. ^ 0. 15).

lit P >< o iK.95 O 5 >< rt] o O o ^c >< O ^ c o ^"5 p ^ o "9-9.9 o p o S O CT3 a§ >< '-> ^ o |g It ^ O CQ «3. ^? o o .2 «o re< c < j-^ a ^ ^ -3 -a SP U. Q.Q >< -e.c (U o en en . en en m &? 009-9-^So. 1^ ^ II^ ^ O l§ § :3 Q_ CL QJ u u IVVJ 2 u -a 1^ c re re ^ c OJ n- ffi < c = " iS re Si <u en C c T3 -13 <0 . o cj y cj o -S o p o Q.CO en en _ _ S Q) Q. c i en "S. p i. The University of Kansas "S .J a c S en 9 <«&.CLO^Cl.i<:0002CL -Q c g eu C en S g m CLcn ^ o. o o O O tl! da oni fc n3 CO v. c o c m .>.-2 o o 9 ej p S ^cn ^ -C ^ 9 <a. i c ^ r In O >. a ^ in re c Xi tn Cl. ^ 5 "en E en < re t^ e:) O C ei) ^1 .enQ) 0>< 0. O $. § a §o 8 en n: etJ E S Cl & 0) ^en ~^ en en C/l t. ^1 ^ Q.2 w re a. 9 t3 a p .i^Cl.18 Scientific Papers.c S.cCl.>< S^S'g '-' ^g-9-i-9^ -* <: P S 3P en en o en —V en 9.en qC « ? CL a D $.J 13 c o c I <U en <u en en en en en en 15.o>.c S = > -2 . en c ei) o2 2§ CO ><Q co>< >< 2. en ^ Cl S S-5 9.><§CLa:l-=i:><. en Q. o o CO p p * ~ ep'S'Q.a la. Natural History Museum. a £i C J^ o ^ S 5 C re IOJ s QJ 00 CD _o i: C U X c ^ •a OJ > "o en 01 .a il O V> ><><e!22><o-jO><><oocuj..'^ o Cl ej o a Q. 9 O O O 3 o p 03 ^ p -2 -5 O 0-5 ej p 9 O o o 55 9 ^'Q u O "D Cl'S. £ en 1.xciBCL'5: ^LV 4M u m^ ^5 -9 §§ 5. -3 fc eu TD >< -C § § 9 3 o CO C c ^ p p ^ -i o -y ej S"^ ^-9 rs o t en o O.'S.2 "S to "3 !.iS en 9.o o -^ Q. & 2 o cl 9- 2 05c "& >._ P R o o O gCD en a 1i :^ 2O 2 Q CT3 S 6 :S £ u >. o e § CL CD S-=i.j.2 CD <j !h re C ra iS 01 . . o o o 9 ?i. Q- Q.ra S.

Proxylocopa is included (Figs. Xylocopini and lost once. that collectively is the sister to the New World Topology 2 divided the members of the more basal clades found in Topology 1 into a number of separate branches. Lestis. Notoxylocopa. = 0.57. Proxylocopa is a basal taxon and the sister group to the RJtysoxylocopa group. 2A. successive basal in the approximations. R.Subgenera and Genera of Carpenter Bees. = 347. {Xylocopa) violacea Zonohirsuta All islands recovered monophyletic Rhysoxylocopa Neoxylocopn groups. the relationships of the other taxa to the Ethiopian group shown in the analysis ^ ^ -C Notoxylocopa Cirroxylocopa Xylocopoda Neoxylocopa Megaxlocopa Stenoxylocopa Xylocopina Rhysoxylocopa Epixylocopa with all characters coded as non-additive may be robust. 14) and X. depending on the analysis. = 359. (Nodula) amethystina X. .22. B) places the Rhi/sox\/locopa group basal in the tribe.I. Xenoxylocopa Mesothchia Platynopoda Hoplitocopa Hopoxylocopa Afroxylocopa Oxyxlocopa Cyphoxylocopa Cyaneoderes Koptortosorva X. multiple gains and losses of this feature must be postulated. C. if the order of character evolution assumed in the that Character 11-2 be gained Nanoxylocopa Xylocopsis Nyctomelilta Schonnherria 19 Dasyxylocopa Xylocospila other analyses proves to be incorrect. = 0.50. respectively. (Xylocopa) valga are progressively more Proxylocopa.L. (n = 700+ trees [stopped before all trees were recovered]. Giiatlioxylocopa) testis. included testis.l. (Koptortsoma) sinensis Analysis Manueliini is 3A. R. 14.353. C. Successive approximations applied to each of the is- Fig. weighted statistics: TL. unweighted statistics: T. 1 2. as the sister group to the large New World group referred to above. In one of these clades Proxylocopa. . Copoxyla + (Ctenoxylocopa + Proxylocopa Copoxyla Gnathoxylocopa Lestis leading to and Nydomelittn are placed along the stem the New World group.89). and consider the relationship Xylocopoides Calloxylocopa Ctenoxylocopa X. = 0. and 6 (n = 110 trees.L.52. R.I.I. 4 A). three of the six analyses generated unique consensus trees. 2. 13 A.L = 0. C. whereas in other topologies.l.xylocopa Prosopoxylocopa. R. The starting islands (from Analysis 3) and the statistics are: Topology 1 (Fig. = 0.I. Copwxyla and Ctenoxylocopa + Giiatlioxylocopa. was poorly resolved with and Cteno.88). unweighted statistics: T. C. Topology 2 was found by starting at Island 2 and 4.88). Island 5 (Fig. Each Old World taxon. C.22. = 0. = 0. the Rhysoxylocopa group. Zonoliirsuta + and the New World group. unweighted statistics: T. This analysis the Rhx/soxylocopa group Pwxi/Iocopa + Notoxylocopa is .I.L. Topology 1 (Fig. . In all analyses with characters coded as additive (Analysis 1. 14) was found by starting at Island 5 {n =700-i.l.L. 4. The second clade is not well resolved.64. = 0. The most basal branch in the tribe consists of Ctenoxylocopa as the sister group to both species of Xylocopa (sensu stricto) and Xylocopoides + Calloxylocopa are associated with the base of the Ethiopian group. 12) divides the tribe into two large clades.l. (Xylocopa) valga outgroup coded as unordered. is the only one in which placed in this position. After the characters stabilized. subfamily. Analysis 3. R. Lestis. island of trees associated the 1 Prosopoxylocopa Alloxylocopa of these subgenera to each other to be unresolved. .l. Copoxyla. = 0. there was support for a monophyletic Ethiopian group. Copoxyla + (Ctenoxylocopa + Gnathoxylocopa). Nydomelitta. weighted statistics: T. Topology 3 (Fig. 12. weighted statistics: T. 3.64. = 344. 1 A. found after successive Island 1 Strict consensus cladogram of six trees approximations weighting. (Xylocopa) violacea + {Calloxylocopa + Xylocopoides) Topology 3 (Fig.) Six analyses were run starting with each of the six islands produced in Analysis 3. however. R. both species of Xylocopa. all characters were — Bomboixylocopa Mimoxylocopa X. as 1 have informally defined it. = 0.22. Biluna Xylocopa (both species) are a grade along the stem of the Ethiopian group. derived taxa. C. Pehxylocopa Acroxylocopa Apoxylocopa Xylomelissa X. along the stem to the Ethiopian group (less Zonohirsuta and Prosopoxylocopa). X.L. = 0. Notoxylocopa + (Gnathoxylocopa + Nyctomelilta) lands produced three very different topologies.L = 0. 12) was found by starting at the trees in Islands 1.343. Howe\'er. (Nodula) bentoni Xylocopoides + Calloxylocopa. = 347. Basal to this New World clade plus Nyctouielitta was a group of Old World taxa that.63. Character 12-1 also occurs in the Neoxiflocopa group and Nanoxi/locopa and as such is weakened.trees [stopped before all trees were recovered]). form a polytomy group.

to <5 to o Q-S o t o-S " a 10 ^5K Q."> u> to 0) c O) . The University of Kansas . O (0 I E o Q.«5 1^ ^§ m . i § o a o o o o la "Ss-c Q.nj II " y ™ O 0) g Q. go 3o>co S-^50-5 SQ.5 P "" o 1 & o .c CO S o O O 0-5 o 5 rag m 5" o 0) 15" <0 to p to to c Q 03 o Q. O qj r Q-O 8 o tj "o >? O o o m j. c sa P .§. o S O O <o CO Q.20 SciENTinc Papers. Natural History Museum. E E § c « ^a Q. •?:>< a ^ 1 § o 8 S ^ to la.1 oi. Q.

= 0. C. all islands). but the In Epixylocopa Perixylocopa Acroxylocopa relationships among groups are often not strongly estab- Apoxylocopa Xylomelissa Biluna Ushed. — Xylocopini is Schonnherria Cirroxylocopa is the subfamily. R. C. Xylocopini basal in the tribe.I. Strict consensus tree of 700 trees after members of a grade of taxa.I. where there were differences they were among the position of the basal members of this clade. each occurs independently in other Xyhcopa. the mesosomal gland reservoir of males (Character 24) is secondarily lost in Rhysoxylocopa and Xylocopa (Nodula) amethystina. and Biliina. Apoxylocopa. R. = 324. With Biluna as the basal branch. Analysis 3. Lengths of horizontal lines are proportional to number of characters on each internode. With Rhysoxylocopa as the basal branch. Relationships among Members of the Major Clades and In the analyses with both nonadditive . 2. in the subfamily. successive approximations weighting. tive. Xylomelissa. Be- mum-length trees \vere Xylocopoda Neoxylocopa Megaxylocopa Stenoxylocopa Xylocoplna Dasyxylocopa Xylocospila Notoxylocopa Rhysoxylocopa Zonohirsula Prosopoxylocopa X.69). R. a derived loss must be hypothesized for one of two characters shared by most members of this group. Mimoxylocopa Bomboixylocopa X. 2A. Island 5. Pwxylocopn found in this analysis (T. 1 include the following subgenera recognized by Hurd and — characters (Analyses acters 1. either as monophyletic groups with poor character support or as Proxylocopa outgroup Fig. (Xylocopa) violacea X. (Xylocopa) valga genera not placed in the above mentioned informal groupings were placed in most analyses near the base of the tribe. A gradulus on Tl of the female (Character 32-2. Epixylocopa. = 0. The derived members of the Ethiopian group were associated in all analyses. A number of other subof the Ethiopian clade Xylocopoides Calloxylocopa Copoxyla Gnathoxylocopa testis Ctenoxylocopa X. The changes in character coding resulted in few changes among the relationships of the ingroup. 3 [Island 3]).24.I. {Koplortosoma) sinensis = 0. the continuous divergence of the inner ventral margins of the gonocoxites (Character 50. Analysis equal weighting. Rhysoxylocopa. A case can be made in com- favor of this second scenario. (Nodula) amethystina Alloxylocopa cause this analysis provides little new information on relationships.L. Depending on the topology accepted. 4. Nodula.I. Analysis lA.62. (Nodula) bentoni summary. Cyaneoderes Koplortosoma X. ters also ing a row coded as nonadditive).26. and the carinate ventroapical plate of the male gonocoxite (Character 52-2). For example. 14. = 302. = 364.L. This analysis produced 123 Hoploxylocopa Afroxylocopa Oxyxylocopa Cyphoxylocopa most parsimonious trees (unweighted statistics: T. C. weighted statistics: T. other reductions in the basitibial plate occur in Biluna. 2.Subgenera and Genera of Carpenter Bees.92). the reservoir of the mesosomal gland originates once in this clade but the row of tubercles on the posterior edge of the female basitibial plate (Character 17) is lost in Biluna.I.L. —Xylocopini basal Hoplitocopa some characters were coded is as nonaddi- Proxylocopa included. The monophyly of the Neoxylocopn group and the Rhysoxylocopa groups were supported in all analyses. 21 Nanoxylocopa Xylocopsis Nyclomelitta The Rln/foxylocopii is not monophyletic basal in and is placed near the base of the Ethiopian group. I do not present the cladograms or consider the results in the following discussion. Analysis 3. lost in Perixylocopa) was the only synapomorphy for the group in all analyses. some characters were coded as additive and included. Results of this search are not shown for the same reasons discussed for Analysis 4 above. Although these are unusual features for bees. Eight mini- others as nonadditive. the basal branch of this clade was Rhysoxylocopa (sensu stricto). including . Xenoxylocopa Mesotrichia Platynopoda is Analysis 4A. successive weighting. Acroxylocopa. = 0. Taxonomic Recommendations Rhysoxylocopa group In the Rhysoxylocopa group. In the other analyses. Tliis result suggests that the problems in resolution found in the other analyses are not a result of polarity assessment based on the use of different outgroups. Biluna and additive was placed at (all the base of the Rhysoxylocopa clade. these data provide strong support for the monophyly of some groups among the Xylocopini. = 0. various other charac- supported the monophyly of this group includof tubercles on the edge of the basitibial plate (Character 17. Monophyly was supported in all but three islands of Analysis 3. 2A). = 0.66. In Analysis 3 char- Moure (1964): Perixylocopa. lA.I.

15. resolved tree randomly chosen from the 24 trees. (Nodula) amethystina X. shown in Figure 11 .22 Scientific Papers. (Koptortosoma) sinensis Koptortosoma Oxyxylocopa Cyphoxylocopa Cyaneoderes Proxylocopa Notoxylocopa Xylocopoides Calloxylocopa Analysis 3. character distribution shown in Appendix 4. The University of Kansas outgroup Nanoxylocopa Cirroxylocopa Xylocopoda Neoxylocopa Megaxylocopa Stenoxylocopa Xylocopina Dasyxylocopa Schonnherria Xylocospila Xylocopsis Nyctomelitta Lestis Ctenoxylocopa Gnathoxylocopa Copoxyla Rhysoxylocopa Epixylocopa Pehxylocopa Acroxylocopa Apoxylocopa Xylomelissa Biluna X. Island 6. One fully . (Nodula) bentoni X. Natural History Museum. Phylogenetic hypothesis based on another island of trees from matrix Fig. (Xylocopa) valga Zonohirsuta Prosopoxylocopa Bomboixylocopa Mimoxylocopa Alloxylocopa Xenoxylocopa Mesotrichia Hoploxylocopa Platynopoda Hoplitocopa Afroxylocopa X. (Xylocopa) violacea X.

Thus. Therefore. (Nodula) bentoni. iXylouwlissn) ruficollis. Apoxylocopa. support for and questionable. Males of X. in his study of South African species of Xylocopa. {Nodiila) bentoni and Rhysoxylocopa. This derived loss also occurs in Xylocopa {Nodula) amethystina. was close Perixi/locopa. one Asian taxon. 3 [Islands 2. The monophyly of Rhysoxylocopa and Biluna is well supported. with a well-defined spine on the apex of the male hind tibia. amethystina and X. and include one species. They clearly in Xylomelissa. Because yellow maculation varies within other subgenera and intraspecifically in X. this species was grouped with the other taxa that lack this structure.. amethystina and two spines in X. Inclusion of X. RJiysoxylocopa all have extremely small propodeal spiracles in the males. The hind tibia flares at its apex and. number of topologies found for members of the Rhysoxylocopa group. the midtibial spines are functionally replaced by an apical mat of highly modified. The males of Biluna lack spines on the hind tibia. None of my analyses grouped the two species of Nodida As the above discussion indicates. either as the sister group to Zonohirsuta (Analyses ter 1. Maeta et al. bentoni. Thus. the two exemplars I used. and the presence of a the posterior acter 17-2). (Nodula) bentoni be recognized as a new subgenus. the loss of tubercles of the female basitibial plate may be associated with loss of other structures on the leg. the reservoir for the is (Nodiila) ameihystina as Epixylocopn this structure and male mesosomal gland (Character 26) is and Biliiua as lacking it. Acroxylocopa. In all analyses. 1.Subgenera and Genera of Carpenter Bees. it was difficult to code unambiguously a spine as present or absent.ocopini males and reducThese probably are associated with movement along the smocith inner surface of the bamboo culms used for nesting (Malyshev. Support for their monophyly is based on the absence of an angle or spine on the hind tibia of the male (Character 11-0). Xyi. Reduction of the hind tibial spine of the male is approached in Epixylocopn and Biliiun. as do males of Rhysoxylocopa. 2 A). so this character may vary it was mistakenly assessed. 4. 23 panded definition of Xylomelissa. Hurd and Moure (1963) recognized nine subgenera not present in X. have the beginning of the thoracic declivity abruptly rounded in the female (Character 23-1) and few setae on the male gonostylus (Character 48-1). amethystina but present in X. Xylomelissa and its near relatives are more problematic. In females. thickened setae on the outer midtibia. Perixylocopa. intraspecifically. Acroxi/locopa. Eardley (1983). differ notably. In the specimens available to me. (Analyses distinct 2A. the conditions were reversed in these groups. row of tubercles on edge of the basitibial plate of the female (CharCharacter 11-0 is found in four African taxa Apoxylocopa and Xylomelissa and and male mesosomal gland reservoirs consist of a single layer of tubules approximately as deep as they are wide in cross section. bentoni. except Bomboixylocopa Mimoxylocopa. obs. whereas Two characters are found only among members of the RIn/foxylocopa group: the absence or reduction of an apex or spine on the outer apex of the hind tibia of the male (Character 11-0). (Nodida) amethystina within Xylomelissa would entail a greatly ex- of the Ethiopian group. depending on the angle in which it is yiewed. (Nodula) amethystina completely lack a reservoir for the mesosomal gland (Character 26). and females possess a dense mat of hairs on the mid-tibia. all other species lack or have reduced forms of this structure. but I have found no discrete characters that serve as synapomorphies to unite these species. lA. amethystina and smoothly rounded in X. Xylocopa (Nodula) as a monophyletic group. X. bentoni. 3. Recognition of this species as the subgenus Epixylocopa was based on the absence of yellow maculation on the face of the males (Character 10-1) by Hurd and Moure (1963). as a rounded projection. but lacks a carina. and in some analyses (Analyses lA. even though they were aware that the Rliysoxylocopa group was poorly understood. Maa (1938) recognized two groups in his original description of Nodula. I have retained this subgenus to include the group typified by X. 1985). whereas in Zonohirsuta rest of the is and the Ethiopian clade (except Platynopoda) the . the ventroapical plate present. Xylomelissa and Apoxylocopa + (Perixylocopa + Acroxylocopa) are either a monophyletic group or closely allied. the —As discussed above. The Hurd and Moure haying (1963) listed X. 3 [Islands Prosopoxylocopa is from Zonohirsuta based on interpretation of the development of the ventroapical plate of the male gonocoxite. For some specimens of Epixylocopn and Biliina. monophyly of the Ethiopian group varied. All females Ethiopian group. which are represented by the two exemplars I used here. The male genitalia are similar. I have proposed some synonymies. I have not examined the specimens that Hurd and Moure used in their study. outer apex of the hind tibia of the female (Character 12) has one spine in X. In Prosopoxylocopa. {Xylocopoides) californica arizoneusis (Minckley. 6]) or as the sis- group to Zonohirsuta plus the rest of the Ethiopian clade 2. Prosopoxylocopa appears as a basal member of the Ethiopian clade in most analyses. Epixylocopa. or present. (Nodula) amethystina and propose that the group represented by X. the stated that other than this character Epixylocopa to Xylomelissa. this feature does not warrant recognition as a subgeneric character. 1931. pers. Character 17-2 is shared by these same taxa plus X. the spine can appear absent. pointed out that subgeneric rank for some of these of taxa is and the beginning of the posterior thoracic declivity is abrupt in X. 2. plete absence of a basitibial plate in the tion or absence of a midtibial spine in both sexes.). I define Xylomelissa on the basis of structure of the gland reservoir. 5]). Xylocopa (Nodiiln) amethystiim. or Despite the which four were monotypic.

Although superficially dissimilar. Mesotrichia and Koptortosoma groups. which places Xylocopa be a strong character in the Xylocopini. is unreversed if Xenoxylocopa is considered to be the sister group to the thus inferred to be the basal mem- — ber of this clade. In contrast. Most analyses showed a sister group relationship between the Mesotrichia and Koptortosoma groups. 1963). B. I favor the recognition of Prosopwxi/locopa and Zonoliirsiita as separate to sinensis as the sister group to the remainder of the Koptortosoma group and the Mesotrichia group. Char. the gonostylus (Character gate gonostylus and is Hoploxylocopa retains the elon- and the Koptorfosoma group is supported by 4-6 characters in all analyses. but leaves Oxyxylocopa . have a rounded them with certainty into Koptortosoma. and Koptortosoma) were resolved by characters that display considerable homoplasy or are continuously distributed. Because the carina of the ventroapical plate seems characters coded as nonadditive). If further studies find Prosopoxylocopa is a member of Zonohirsuta. 15) and 3A (Fig. 1964 [see to LeVeque. and Platynopoda) is supported by seven inferred the Mesotrichia Bomboixylocopn and Mimoxylocopa. a rounded thorax in the female (a derived loss. 2 (all islands) and 2A. group and the Koptorfosoma group. Females of X. synapomorphies. Despite this evidence. The most constant of these characters.24 Scientific Papers. whereas in all islands of Analysis 3 (Figs. male genitalia of Hoplitocopa and the broadly fused gonostylus and large projection from the posteroventral margin of the gonocoxite) that suggest an examination of the other species oi Mesotrichia and Platynopoda might support a different topology than that shown here. Hurd and Moure (1963) considered Bouiboixylocopa most closely related to Nyctomelitta. Platynopoda have a sister relationship is and Hoplitocopa are shown based on six synapomorphies. Cyphoxylocopa. separate subgeneric status for both taxa will not ably a spurious result because of the high weight as- signed to the carinate beginning of thoracic declivity (Character 23-2. the triangular shape of the hind trochanter in the female (Character 14-1) is strong. two characters are unreversed presence of a fovea on the anterior surface of Tl in the and male (Character 33) and either a fovea or the entrance to an acarinarium on the anterior surface of Tl in the female (Character 34). 23-0). 1963). The perpendicular angle of the beginning of the posterior thoracic declivity in both sexes (Characters 22-2. g. In Analysis 1 (Fig. sinensis be warranted. 8-1). 52-2). As stated have a distinct carina on the ventroapical plate of the male genitalia (Char. 35) and a projection on the inner basal margin of the Mesotrichin all above. the possession of a medial keel pology may be favored numerically. 1 A. The exception was in Islands 1 (Fig. Cyaneoderes. 12) Xenoxylocopa is their sister group. in all of the analyses except Analysis 2A. support a sister group relationship to Mimoxylocopa. This probis subgenera. but found in some species of Koptortosoma (sensu Hurd and Moure.xylocopa. 3. neither to- The monophyly of the Koptortosoma group is supported by the presence of a transverse carina on the scutellum that in most females overhangs the metanotum (Character 23-3). paired setal patches on the pregradular areas of the male (Char. Alloxylocopa. The best supported topology depends on judgment of the strength of three characters. of Xenoxylocopa. Mesotrichia as a monophyletic and Platynopoda + Hoplitocopa are supported group by the reduced length of the male 47). all of which are unicjue to the Xylocopini. and 6 of Analysis 3A on S6 of the male (Character 37). Character 30. but other characters place the venter of the gonocoxite (Char. Mesotrichia. however. The placement of the two basal taxa of this clade differed between analyses. All my analyses. 11 A. supports a clade of . three of these are strong. Cyphoxylocopa. lOA. Natural History Museum. includes Bomboixi/locopn + Mimoxi/locopa. 52-2). including one feature that is uniquely derived the produced apical edge of SI (Character 36-1). see below). The monophyly Mesofrichia group. These include the long malar space in the female (Char. Alloxylocopa is the sister group to the Mesotrichia and Koptortosoma groups.. Maa (1940) originally placed it in the subgenus Zonoliirsiita. Xenoxylocopa. Afroxyloccpa and Koptortosoma (sensu Hvird and Moure. a medial sulcus on Tl (Char. 23-2) provides the strongest support for the monophyly of the clade consisting of thoracic shape. I consider this character to be a derived loss in X. there are similarities of the Mesotrichia (e. The University of Kansas (successive weighting analysis applied to data matrix with all ventroapical plate has a distinct carina (Characters 52-1. B). and the slender penis valves of the male genitalia (Character 55) are found elsewhere in the ingroup only in Xylocopa (sensu stricto) and Xylocopoides. the strongest one the possession of a sclerotized base of the penis (Character 45). 53-1). 1928]). Cyaneoderes. The next most derived lineages within the Ethiopian group. 2. rufipes is the sole species of the subgenus Mimoxylocopa. Alloxylocopa. Hoplitocopa. 39). An elongate tegula in the male (Character 29) is unique and unreversed. (Koptortosoma) sinensis. Relationships among the taxa of this group (Oxyxylocopa. Because characters 33 and 34 could be correlated (similar character states in both sexes). and Xylocopa [Koptortosoma] sinensis. B. X. many features unite and Koptortosoma groups. 12). The Mesotrichia group (Hoploxylocopa. Alloxylocopa is if placed as the sister group to the Mesotrichia Koptortoso7na groups. and patches of appressed setae near the metasomal spiracles (Char 40). 13 A. although this condition is approached by some species oi Mesotrichia (sensu Hurd and Moure. Afro. the shape of the gradulus on Tl.

Xylocopsis. Xylocopina. This character is similar to the condition found in Schojtnherria and Dasi/xylocopa. — New World taxa except Xylocopoides. thereby suggesting independent origins in these taxa.Subgenera and Genera of Carpenter Bees. three of by four inwhich display considercharacter is able homoplasy. differences in the color of the vestiture. 135). such features are the absence of a reservoir of the male mesosomal gland guatemalensis). In my analysis. all characters coded as nonadditive and weighted equally. the beginning of the thoracic declivity on the scutellum of the male {Cirroxylocopa. 1963) I among the analyses that included additive 1. 2. see below) and Neoxylocopa all (sensu lato. B. X. Megaxylocopa). Notoxylocopia. Xylocopini defined by plesiomorphies. Xylocopoda. Koptortosoma (and is allied taxa as discussed herein) the most species-rich broadly attached gonostylus (Character 48). Fig. a slender. . or on the propodeum (Neoxylocopa and includes all of the Calloxylocopa lA. I conclude that these subgen- because the ventroapical plate varies era should be united with the subgenus Koptortosonin. Stenoxylocopa.. to a flattened plate with a sharply-angled carina (Fig. characters (Analyses Many characters serving as found no derived features that unite Oxi/xylocopn. like that of the South African endemic taxon in the Xylocopini. These are: (1) Character 271. Analysis 3. are highlv variable. Mesotrichin (sensu Hurd and Moure. X. strongly suggests their relationship with (Character 49-2. the gonostylus at its attachment to the gonocoxite is thick in Schonnlwyria (sensu lato) and Dnsyxylocopm. continuously across taxa from a rounded condition with carina The four taxa est in the Koptortosoiiin group with the great- number of derived characters (Afroxylocopa. Xylocopa Neoxylocopa group and Schoiuihcrria Naiioxylocopa. C). Xylocopa no evidence of a carina.. Ci/plioxylocopa. Afroxylocopa. One strong feature in Schonnherria (including Xylocospila). tubercles on both sides of the basitibial plate of the female (Character 17-1. 1. obs. In Nyctomelitta. and Nyctomelitta) differ greatly on S6 is and occurs in and Xawxylocopn. but many characters cited above occur in the Old World clades. and the presence or absence of a mite pouch in Tl of the fe- synapomorphies in this group are highly homoplasious. Hurd and Moure. males. and Notoxylocopa. Furthermore. and Cyniicodcres into a single The apomorphies that Hurd and Moure (1963) used to differentiate these taxa. but because it is not found in all species of Koptortosoma (sensu Hurd and Moure. 2[Island 3]). Character 43 is unique to this group. pnibescciif. and Xylocopoda). its biogeography. and relationships stituent taxa among the con1 would require more detailed study than have undertaken. indicated a monophyletic New World group. Character 52. Neoxylocopa. 1983). and they have other features in common with taxa that do not have the sharply-angled ventroapical plate. elon- between the clades and Gnathoxylocopa were variously placed that include Schonnherria and Neoxylocopa. which concerns the ventroapical plate. these features gonostylus (Character are tubercles on both lateral edges of the basitibial plate of the female (Character 17-1) and the thickly hirsute and provides weak sup- port for the its monophyly of the group. the vestiture on the dorsum of the male entirely yellow (Character 1 ). 4). A relatively strong the well- developed medial lobe at the apex of the gonostylus (Character 49-2). Neoxylocopa. the median keel 25 Dasyxylocopa. and (3) Charsmooth zone along the epistomal suture (Megaxylocopa. and the absence of a metapostnotum in the males (Character 24). Dasyxylocopa. 26-0). Characters 3 and 24 are highly homoplasious. (Char. the presence of a tooth on the upper carina of the female mandible (Character 3). (2) Character 48-0 (a loss). However. the strongest is the presence of two spines on the outer apex of the female hind tibia (Character 12). 8A. and slender in Koptortosoma (sensu Hurd and Moure. 1963) fusion of the upper rutellar ruficollis. also see Eardley. Notoxylocopa. 1963) (pers. pubescens logenetically useful ture revision. gate gonostylus with few setae (Megaxylocopa. In the case of Nyctomelitta. 1963). is a strong synapomorphy. and X. analy- They share three synapomorphies. Notoxylocopa and Nyctomelitta may be sister groups to the New World group of taxa. In Notoxylocopa. The shape of the medial lobe of the gonostylus Gnathoxylocopa. 1963. and few characters support these branches of the cladogram. I did not include this as a character in the data matrix. sinensis may prove phyand should be considered in any fu- Some characters suggest groupings within the clade that Neoxylocopa and Schomiherria clades. in Xylocopa and the thickly hirsute and broadly attached 48). and the Old World subgenus Nyctomelitta. In with other analyses (Analyses Nyctomelitta. Xylocopsis. Notoxylocopa. because it shares with Stenoxylocopa (sensu Hurd and Moure. Xylocopina. Fig. sinensis) are defined ferred synapomorphies. and Xylocopoda). Notoxylocopa caffra. Nanoxylocopa and the Neoxylocopa group is a sharply-angled ventroapical plate of the male genitalia. share three characters: a carina or protuberance on the outer lateral edges of the penis valves (Character 43). clade. 1963. Stenoxylocopa is defined here to include The relationships among the taxa associated with the (i. to a rounded condition with a weak (Hurd and Moure. was coded according to the presence or absence of this carina. see below) were closely grouped in ses. lost in males of some members of Koptortosoiiia. such as the shape of the male face and eves. Stenoxylocopa. and Xylocopina). 241) of other Old World taxa. Stenoxylocopa. These taxa. it and Nyctomelitta do not possess these features. the presence of a carina or Stenoxylocopa (sensu lato. acter 6-1.e. and X.

if these are sister taxa this relationship forces two characters. according to the description of Maa. 1963). or the sister group to Schomiherria (sensu lato) (Analyses 1. IC). and Xylocopoides is from the paraphyletic. Xylocopsis. and an angle or projection on the base of the ventral side of the gonocoxite (Char 53). Features of the coded as missing for some of these taxa mesosomal and might progland are vide insight into the relationships of these subgenera to the hneages associated with Schonnherria. to evolve convergently in Calloxylocopa and Xylocopoides. Neither of Two are plesiomorphies —the — these types of characters Stenoxylocopa tirely is useful in establishing relation- Miscellaneous other clades.. str. 41-2) of the penis valves (Char. 11-2 and 12). as the basal taxon to the entire New World group (Analysis 3 [Islands 1. patches of appressed setae on the pregradular area (Character 39). Xylocopa ruficoUis leave these was previously placed in the monotypic subgenus Xylocopina by Hurd and Moure (1963). Until the relationships is polyphyletic or probably more closely related to Ctenoxylocopa because both share many features and have gonostyli that are slender and largely without fined by Hurd and Moure If (1963). Xylocopoides genus Megaxylocopa is distinct. Although I agree that the sub- Other relationships reveal disturbing character conamong these groups such as. valga from the fully fused condition (Char. sister and that X. their allies. based on four characters: shape of the apex of SI and the yellow markings restricted and two are autapomorphies to the lower part of the face the short clypeus and the shape of the tegula. The male genitalia of thus seems unlikely. a series of spines along the lateral edges of the pygidial plate (Char. 41-2) and the hirsute outer apex of the penis valves (Char. Dasyxylocopa. as de- This hypothesized relationship inclusion of X. so. most other Xylocopini are tropical. but it seems unlikely that once lost they would be subsequently regained. and Lestis) have distinct characters. a row of preapical spines The subgenera Neoxylocopa and Megaxylocopa are united by yellow vestiture (Character 11).26 Scientific Papers. and Xylocopoides are also the two most northerly distributed groups in the tribe (Popov and Ponomareva. some of which may be synapomorphies. strongly supported. among Xylocopoides. among these taxa are . 1994. but they also share a number of plesiomorphic features. Naiioxylocopa. Dasyxylocopa is monotypic.: two spines on the outer apex of the hind tibia in both sexes (Char. its recognition clearly ren- ders Neoxylocopa paraphyletic and ignores the strong similarity between Neoxylocopia and Megaxylocopa. Calloxylocopa and Xylocopa s. Three strong apomorphies found elsewhere only in the Ethiopian clade are found in Xylocopoides and Xylocopa s. str. Neoxylocopa. The clade consisting of Dasyxylocopa. The remaining taxa (Xylocopa [sensu stricto]. and was defined by Hurd and Moure (1963) largely by the lack of features found in Schonnherria (sensu Hurd and Moure. found in Schonnherria and warrant inclusion of this monotypic subgenus as a member of that group. as it requires the gain of a well-developed gonostylus in X. I consider it more probable that Calloxylocopa is a derived member of the subgenus Xylocopoides. Fig. or Xylocopina is defined en- The relationship of Calloxylocopa by plesiomorphies. Xylocopa s. This character occurs elsewhere only in Koptortosoma. Hurd and Moure (1963) hmited the subgenus Megaxylocopa to — pygidial spine (Char. Hurd. paraphyletic. Natural History Museum. Minckley. 6]). 44) are features known in no other bees. and X. Gnathoxylocopa. 44). valga is and setae (the latter feature was not coded in this analysis). These four taxa share no strong features with other groups in the Xylocopini. Character 26-3 and a modification of the scutellum forafeatures of the males that accommodates the tubules of the mesosomal gland (Minckley. 1978). is were not well resolved by my analyses. and Xylocospila can be defined by the presence of a well defined ventroapical plate of the male genitalia (Character 52-2) and the morphology of the male mesosomal gland (Character 26-2. flicts men those species having a conspicuous carina over each of the lateral ocelli in the female. In addition to character 53. X. F) are unique synapomorphies of Neoxylocopa and Megaxylocopa. these characters may indicate relationship of this group to the Ethiopian group. I tooth with the preapical tooth (Fig. Ctenoxylocopn. However. 1961. 5). 4 B). Proxylocopa. without including Xi/locopiua. Copoxyla. this assignment was taxa as separate subgenera. 3). 1994. (Xylocopa) violacea. The monotypic subgenus Nanoxylocopa was placed as a basal taxon of this group (Analysis 3A [Islands 1. Most relationships among these taxa — ships (Wiley. and may be plesiomorphic members of the Xylocospila share all features is complicated by the subgenus Xylocopa. 42) found in X. Like Nanoxylocopa and Xylocopsis. not found in other bees. (Xylocopa) violacea represents the group to Xylocopoides. This might be evidence that Xylocopa s. and Xylocospila share a well-developed medial lobe on the apex of the gonostylus (Character 49-2). The University of Kansas further clarified (see further discussion below). 5. 5E. 2 [Islands 1. (Xylocopa) violacea completely lack gonostyli and have a foveate basitibial plate (known elsewhere only in Lestis and X. str. Xylocopsis. As previously discussed. and the distinctive shape of the tubules and large size of the male mesosomal gland (Character 26-3). 3. Gonostyli are commonly Icwt in bees. (Xylocopa) convexior Hedicke. 2. and setae on the outer apex The relationship proposed Schonnherria. Fig. valga in the Schonnherria lineage. 1954). Xylocopoides + Calloxylocopa. Schonnherria. 1954). str.

sometimes cleft apically [southeastern Canada and northwestern United States Xylocopoides to southern Mexico] Labrum with 4 or fewer tubercles 7 Labrum tral with 5 tubercles (abundant long setae) [cenXylocopsis Argentina to southern Brazil] Frontal carina elongate open at the base. Nonetheless. KEYS TO THE SUBGENERA OF XYLOCOPA New World 1. Clypeus at least half as long as broad. 48) and the gonocoxites diverge continuously from the ventral gonocoxal bridge (Char.. The male geniPwxylocopa are most similar to those of the Rhi/foxi/locopa clade and/or Copoxx/ln. str. 3 (1). 1 A) [southern South America to southwestern United States. in part. Except for Table 1. median tubercle of labrum narrow (body elongate. The absence of a defined metapostnotum in the female (Char. An example might be the basitibial plate as it is known to readily respond to changes in body size in other bees (Reyes. Medial longitudinal carina on all metasomal sterna. because of my inability to homologize features of the basitibial plate and pygidial plate. median clypeal 9 clypeus usually bounded by continuous impunctate 4 ridge length longer than the distance between clypeus and anterior ocellus Medial longitudinal carina is not on is sterna or entirely absent. transverse. Structures that are readily influenced features that also occur Xi/lomelissa among some in Copoxyla. closed basally (Fig.. or ridge entirely absent . Lestis and Pwxylocopa are hereafter treated as subgenera of Xylocopa. median clypeal tinuous impunctate ridge. From here on in the paper. which should reveal new I characters or duction). 25) and one tibial spine in the male (Char. The relationships require further analysis before any subCnlloxylocopn. 11) are gize suggest that talia of These analyses show that many of the characters hisused to define the subgenera and genera of the Xylocopini arise repeatedly in the tribe and many groups are defined by unique combinations of characters rather than by characters that do not occur in any other group. IC) [southern Arizona in the United States to southern Argentina] of T6 7C) 2 Upper tooth marked by numerous spines and not strongly divergent anteriorly.. stantiated statements of relationships can be preceding sections are accepted. A possible problem with these analyses is the use of outgroup comparison for structures that might be drastically influenced by a tremendous increase in size. tuberculiform and not extending between antennal sockets [central Argentina Nanoxylocopa to southern Brazil] 8 (7). Pref- example. have retained these groups as subgenera of Xylocopa. subcylindrical) [Costa Basitibial plate 7 (6). median tubercle of labrum broad. tooth (Fig. up to this point the taxa recognized CtenoxyJocopa. introduced on Hawaii and Neoxylocopa 6 to Guam] 5 (3). is thought by some to be the most ancestral taxon in the tribe (see hitroIts relationship to the other Xylocopini is unresolved in this analysis. 52). 7B) Pygidial plate with lateral margins not . not foveate... Proxylocopn. as are the nomenclatural results. Basitibial plate foveate. In general. rather globose. sometimes represented by a spine that often has a shorter preapical spine on each side of its base and thus 3 not or scarcely represented on disc of T6 (Fig. Copoxyla. Xylocopini The ground-nesting group. clypeus all metasomal bounded by discon- Clypeus less than half as long as broad. conclude that among these taxa {Xylocopn s. For made. 8 Rica to southern Mexico] Calloxylocopa Frontal carina short. I by the large size may make determination of the groundplan states for these characters difficult or impossible. outer Notoxylocopa apex of hind tibia with two spines. Upper tooth of mandible much wider than lower Stenoxylocopa marked by numerous spines and strongly divergent anteriorly on disc of Pygidial plate with lateral margins (Fig. Feniales 4 (3). 2A). Co- 2 (1). of members of the group and not evidence of close relationship. outer apex of hind tibia with long curved spine. Gnathoxylocopa. mandible as wide as or narrower than lower tooth (Fig. and may be further Copoxyhi and Pwx\/locopn lack the ventroapical plate of the gonocoxite. the relationships indicated in the Proxylocopn and testis) the most plesiomorphic members of the tribe are likely to be found. 1993). those structures could homolo- 27 more of the species erably future analyses should include in these groups. I it is a plesiomorphic taxon. The gonostylus of these taxa is thickly hirsute apically and broadly attached to the gonocoxite (Char. Propodeal triangle present Propodeal triangle absent [western United States lombia] 6 (5). Xylocopoides + by Hurd and Moure (1963) have been used. largely redefine all some of those used in this study.Subgenera and Genera of Carpenter Bees. the members of the Xylocopini are many times larger than most memtorically bers of the outgroup. 5 length shorter than the distance between clypeus and 10 anterior ocellus .

highly polished tubercle [south central Brazil to Paraguay] . C) Face with yellow maculations extending to vertex Stenoxylocopa Apex 3 (2). Natural History Museum. across ventral surface of tibia [central Argentina southern Brazil] Xylocopoda Hind tibia without lamella. Scientific Papers. body inbrilliantly metallic [southern United Schonnherria States to Argentina] Eyes at least equally convergent above and below. 3D. propodeum Disc of scutellum not entirely subhorizontal. produced [extreme southern Dasyxylocopm base of hind tibial with high. parocular area not modified as above 12 12 Disc of scutellum entirely subhorizontal. Apex of SI entire. inner hind tibial spur absent. sometimes very weakly metallic 2 Graduli on Tl. ferruginous or fulvous pubescence. posterior declivity of thorax beginning on the metanotum or 10 Thickly plumose hairs on the disc of the first metasomal tergum. southern Australia to northern China] tral surface . Lieftinckella) 4 (3). Integument mostly black at least on dorsum with blackish. whitish. without a longitudinal yellow Apex of basitibial plate asymmetrically or symmetri3 of basitibial plate entire (Fig. tegula punctate throughout or thickly pubescent Sparsely plumose hairs on the disc of first metasomal 10 tergum. The University of Kansas Hind 7 (6).. thin lamella beginning near spur and extending obliquely for- Brazil] ward to tegument often Eyes more convergent above than below. Anterior half of T3 broadly depressed on each side Beginning of thoracic declivity represented by a gently or abruptly rounded angle 2 (1). posterior declivity of thorax undefined (antennal scape yellow. B. T7 without dentiform projections on apex 5 Beginning of thoracic declivity represented by a transverse carina that divides the scutellum into a dorsal and vertical surfaces T7 with dentiform projections on apex [western United States to Colombia] 2 Notoxylocopa 5 (4).. 3A. 6 Body length generally less than 25 mm [Africa. introduced on Hawaii and Guam]. of basally carinate median line anterior margin of each 3 depression bearing transverse row of very long. Mandible with three teeth on apical margin. flattened. eyes more convergent below than above [southMouoxylocopa central Brazil] circummarginally) [central Argentina to southern Brazil] Nanoxylocopa (9). 8 (6). posteriorly directed plumose hairs which overlie the depression [Costa Rica to southern Mexico] Calloxylocopa Body length generally greater than 25 mm [Africa. 11 cally bifid (Fig.. integument usually non-metallic. eyes equally convergent above and below [central Argentina to southern Brazil] Xylocopoda (11). integument bare on discs of T1-T5 and largely eburneus [Argentina] Schonnherria (in part) Old World Females (Excluding 1. integument usually brilliant metallic (often with apex of basitibial plate symmetrically or asymmetrically bifid) [southern United States to Argentina] Schonnherria (in part) 2(1). Mesotrichia Indian subcontinent to Indonesia and north to China] T3 not modified as above 6 (5).. Integument mostly ferruginous or yellowish with 1.28 9 (8). tegula black. body integument at most feebly metallic in bright light 11 . apex of tibia beneath with large. east to Koptoitosonia Hind tibia with spines or high. 4 of head [southern Arizona in the United States to south- Vestiture chiefly black. tegula ferruginous. scape entirely ferruginous or with longitudinal yellow frontal stripe [southern South America to southwestNeoxylocopm ern United States. integument on discs of TITS black and covered with dark setae [central Argentina to ern Argentina] Face with yellow maculations extending to just above antennal sockets [extreme southern Brazil] Dasyxylocopa southern Brazil] Xylocopsis Vestiture chiefly brownish. thin lamella on ven7 the Pacific. not produced [south-central BraCirroxi/locopa tibia not modified zil to Paraguay] of SI distinctly Hind tibia Apex 10 (8). E) frontal stripe 11 (10). New World Males T2 or more terga. Graduli absent on T2-T5. or griseous pubescence. scape largely or entirely black. parocular area with a large elongate depression adjacent to lateral sides of antennal sockets [south-central Brazil] One spine on outer apex of hind tibia Two spines on the outer apex of the hind eastern 9 tibia [south- Canada and northwestern United States to Diaxylocopa 9 southern Mexico] (8). Cirroxylocopa 11 (10). Xylocopoides Mandible with two teeth on apical margin.

one or both lateral edges of the basitibial plate crenulate (Fig. Parocular area without tubercle. Turkestan region to the Indian subcontinent] 12 (5). ment equal Parocular area with tubercle below antennal socket. medial groove of propodeum indistinct [northern Africa. 7B) (lateral line of Tl transverse. IB) [Africa Gnathoxylocopa (basitibial plate Basitibial plate absent. lateral line of Tl transverse [Africa Ocelli normal. 2A) 6 (5). Old World Males 1 (Exclliding Liefinckella) (Fig. Beginning of thoracic declivity represented as an abrupt rovmded angle in scutellar profile 14 i. to InBiluna donesia and west to Indian subcontinent] 4 (3)..Subgenera and Genera of Carpenter Bees. 2A. medial groove of propodeum flagellar 10 (9).] plate bifid. if present. 10 segment shorter than combined lengths of succeeding three segments.e. Is. represented by a spine that often has a shorter preapical spine on each side of its base and thus 5 not or scarcely represented on disc of T6 (Fig. lateral edges of the basitibial plate entire (Fig. tibia with two spines or con12 15 (14). thoracic shape being evenly rounded in pro- Pygidial plate with lateral margins strongly divergent anteriorly on disc of T6. posterior part of the lateral line of Tl curved posteriorly. integument reddish- Anterior face of Tl lacking a vertical sulcus altogether or. diameter greater than twice the width of the scape (basitibial plate bifid. Beginning of thoracic declivity broadly rounded. lateral line of Tl curved posteriorly. E) (supraclypeal base (Fig. maxillary palpus to distinct [Africa] Xylomelissa first flagellar seg- six-segmented [northwestern India Turkestan] southern Maaiana Discal hairs of Tl thickly plumose.. sulcus not as well developed as above. B. Anterior face of Tl with medial. at base 7C) 15 Minor to Turkestan Pygidial spine without a preapical spine on each side at region] Proxylocopa tibia base (Fig. C) [Tibet through the tropical Orient to Pygidial spine without a preapical spine on each side at Apex of basitibial plate simple (Fig. apex asymmetrically or sym3 with tubercles along both lateral edges (Fig. C. (Fig. metrically bifid (Fig. forming a conspicuous posteriorly-directed lobe that overhangs entire metanotum Nodida [Sri Lanka and peninsular India] Scutellum not forming a conspicuous posteriorly-directed lobe Length of metanotum subequal from side to side [Mediterranean Basin to central Asia] 18 (17). 2B)) [Africa south of the Sahara. 2B) Dorsal surface of Tl angulately (or subangulately in Alloxylocopa) onto anterior surface. D. E) southern China] Outer apex of the hind vexities (Fig. maxillary palpus four-segmented [northern and eastern Australia] 11 (8). 1 A) 11 Length of metanotum greatest medially 19 Scutellum modified.. to black with ferruginous pubescence) [Indian Nyctomelitta more or less 16 (15). Bomboixylocopa 13 (12). 2C. Xylocopa first Discal hairs of Tl simple or sparsely plumose. Pygidial spine with a preapical spine on each side (Fig. 7B. area strongly swollen. lateral 5 line of Tl transverse rounded Dorsal surface of Tl sloping or rounding onto declivous anterior surface. or its apex entire (Fig. file not 17 defined. Xylocopini 3 (1). Mandible with two teeth on apical margin 9 17 (13). parallel to lateral margin of metasoma . 2B. southern Europe into Asia Minor] Rhysoxylocopa Mandible with hook-like process on the lower edge south of the Sahara] . 29 Midtibia with a spine on outer apex 4 bounded on Midtibia lacking spine on outer apex (anterior ocellus either side by an impunctate. bigibbose) [Madagascar] Prosopoxylocopa 8 (7). vertical sulcus that is broad and deep. Sulawesi (Fig. 13 . Lestis to or longer than the succeeding three segments. more than seven times wider anteriorly than posteriorly [Asia 14 (13). Ctenoxylocopa Minimum length of malar space less than length of second flagellar segment. 7B) [Europe to central Asia] Copoxyh (Fig. IB) . brown 7 (6). crescent- Minimum length of malar space more than length of second flagellar segment.. Pvgidial plate with lateral margins not strongly divergent anteriorly. 2E) 2 Mandible without hook-like process Basitibial plate present. sometimes feebly so Zouohirsuta Pygidial spine with a preapical spine on each side (Fig. 2D. Outer apex of the hind with one spine or con6 fold of anterior face of Tl broad to and deep) [Middle East Alloxylocopa vexity (Fig. C) 2 (1). Mandible with three teetli on the apical margin 9 (8). 16 subcontinent to Java] Apex of basitibial B. diameter much less than twice the 7 south of the Sahara] Xenoxylocopa wddth of the scape Ocelli enlarged. C) . E) [northeastern China and Korea] shaped swelling) [Taiwan and southern China. vertical 5 (4). at base 7C) 8 2A.

2B. outer margin convex.. abruptly and 7 angulately divided from anterior surface Apex 16 (15). 20 Tegula punctate throughout [Tibet through the tropiZonohirstita cal Orient to Sulawesi] 19 (17). frontal carina siilciform. Eyes closest medially. inner orbits convex. beginning of thoracic declivity represented as abruptly rounded angle in scutellar proPwsopoxylocopa file [Madagascar] Inner orbits of eyes concave. The University of Kansas more 3 (2). 4B). Clypeus with yellow maculations 15 (14). lar Minimum malar length shorter than second flagelsegment. Beginning of thoracic declivity indicated by an 18 Outer apex of mid tibia with a spine 9 abruptly rounded angle in scutellar profile Outer apex of mid tibia without a spine (anterior ocellus bounded on either side by impunctate. glabrous and shining (northwestern India to southern Turkestan) Outer apex of the hind tibia tibia with one spine (Fig. T7 with pair of dentiform spines [Europe to central Asia] Copoxyla Dorsal surface of Tl sloping forward. thorax evenly rounded in profile 19 18 (17). towards antennal sockets. not modified. diameter less than twice the width 4 Nyctomelitia 12 Ocellus enlarged. Posterolateral third of tegula impunctate. Xylocopa (sensu stricto) Ragellum cylindrical. and shining [Africa. diameter greater than twice the width of the scape [Indian subcontinent to Java] . to InBiliina donesia and west to Indian subcontinent] 9 (8). or less parallel to lateral margin of metasoma . at least laterally) [China] Hind femur without tubercle. E) 10 Outer apex of the hind 10 (9). latbounded laterally by conspicuous transXylomelissn (in part) verse ridge (Africa) Minimum malar flagellar length as long as length of second segment. Turkestan region to the Indian subcontinent] Labrum apically entire 13 lum Labrum apically emarginate 14 13 (12). not elevated at distal apex. 8 longer posteriorly than anteriorly 12 of the scape Ocellus normal. posterolateral third impunctate. subangulately rounding onto anterior surface (tegula punctate mum Propodeal spiracles small and asymmetrical. Beginning of posterior thoracic declivity on scutel(posterolateral lobe of pronotum prolonged posteriorly so as to extend well onto mesepisternum and nearly attaining preepisternal groove) [Africa south of the Sahara. Propodeal triangle defined [Mediterranean Basin central and palpus five or six forewing much Europe to central Asia] Bomboixylocopa. rounded behind (Fig. Bomboixylocopa (in part) 14 (12). second submarginal cell of Proxylocopa . testis 4 (3). Indian subcontinent to Indonesia and north to China] 6 (5). face with two parallel stripes running along the inner margins of the eyes from the vertex to the bases of the mandibles (entire metanotum exposed dorsally and base of propodeum horizontal. Beginning of thoracic declivity not defined. Hagellum crenulate on inferior surface. crescentshaped swelling) [Taiwan and southern China. D. face entirely black or with yellow maculations only on areas below the lateral ocelli 11 eral ocellus not Hind femur with tubercle on ventral surface. glabrous and shining 6 rounded in profile [northeastern China and Korea] .. curved inward towards central eye axis. minimum interorbital distance shorter than length of antennal Xylomelissn (in part) scape [Africa] (14).. bowed Cteuoxylocopa Beginning of posterior thoracic declivity on metanotum Gnathoxylocopa [Africa south of the Sahara] 5 (2). Propodeal spiracles normal. glabrous.. Maaiaiui C. with two spines (Fig. frontal carina cariniform. maxillary palpus four-segmented.. scutellum Tegula normal. Clypeus black 15 17 16 Tegula elongate. southern Rhysoxylocopa Europe into Asia Minor] of the Sahara] Anterior surface of Tl without a fovea [Africa south Xenoxylocopa Anterior surface of Tl with a fovea [Africa. outer margin at least slightly concave (Fig. 2A) . miniinterorbital distance equal to or greater than throughout) [Middle East to southern China] AUoxylocopa 7 (6). lateral ocellus often bounded laterally by conspicuous transverse ridge (Sri Nodiiln Lanka and peninsular India) 20 (9). second submarginal cell of forewing nearly in form of parallelogram (integument brightly metallic) [Australia] (11). 4A). in part (X. Australia to northern China] 17 8 (2). posterolateral third may or may not be impunctate. rufipes) 11 (10). east to the Koptortosoma Pacific. length of antennal scape [northern Africa. Mesotrichia Apex of of T7 entire Dorsal surface of Tl nearly horizontal. Natural History Museum.30 SciENTinc Papers. maxillary Propodeal triangle absent [Asia Minor gion] to Turkestan re- segmented..

bv original designation. (8) first flagellar Apex of T7 of male entire. except in X. (54) Female malar space shorter than width of segment. 1963: 236. (2) Upper carina of male mandible without a tooth. (26) Mesosomal gland reservoir a male on scutellum. (28) single layer of parallel running tubules. Hurd and Moure. New synonymy. (17) Posterior edge of female basitibial plate with a Hurd and Moure. (40) Patches of appressed setae present near spiracles on T2-T6 of males. 1963: 230. erythrina. (50) Females are similar to those of Rhysoxylocopa but can be distinguished by the presence of a distinct medial groove on the propodeum. Bihiiin is prob- ably not a member all of the clade. except for X. (43) Outer edge of penis valve not modified. (29) Male tegula not elongated posteriorly. (4) — Two teeth on apical which Upper rutellar tooth and preapical tooth of female mandible separated by emargination. (49) Gonostylus apex with weakly developed. and X. (15) Male mid. Emylocopn Hurd and Moure. (13) Tibial spine of female originating at tibial apex and punctiform. (7) Supraocular pits of female bridge or nearby. (18) Apex of male basitibial plate en- species: Xi/hcopti airlnatn tire. capitata Male metapostnotum absent. (23) Thoracic profile of female smoothly rounded. without lateral projection. (35) Medial groove of Tl of female linear. Maa (1968. bv original designation. 1874. New synonymy. by original designation. capitata. bv original designation. (11) Outer apex of male hind tibia without spines. Smith. not triangular. 1854. with a carina. 1793. (47) Gonostylus broadly attached to gonocoxite. Xylocopa absuniipes. except in X. (although in some taxa) and the presence of a basitibial tinued posteriorly adjacent to lateral margin of the metasoma. 106) effectively synonymized Perixylocopa with Xylomelissa in his species citation. Type species: Xylocopa capitata Smith. without a ridge. (37) Medial carina restricted to posterior half or absent on S6 of male. (16) Male hind femur with basal tubercle. 1963: 223. reduced. (20) Female basitibial plate open basally. Description. capitata Medial ventral margin of gonocoxites diverging continuously from dorsal gonocoxal epistomal suture smooth. (27) Beginning of thoracic declivity of at its Propodeum connection with metasoma not modified. Tvpe species: Xylocopa en/thrina Gribodo. Type row of tubercles. (24) rufitarsis. 1963: 232. (53) Medial projection or spine at base of ventral gonocoxite absent. (51) Gonocoxite lacking a projection or spine from posterior ventral margin. (48) Gonostylus broadly attached to gonocoxite and apically with dense patch of setae. New synonymy. (36) Apical margin of SI entire. Hurd and Moure. albifrons. (34) Anterior surface of Tl of female without a foveal pit or entrance to acarinarium. Type species: Xylocopa absurdipcf Enderlein. except in X. (25) byMaa. (1) Color of vestiture on male dorsum predominately dark brown or black. in which it is medially emarginate. Pcriw/locopii 1968: 106. Type species: Apis amcthystina Fabricius. rufitarsis and X. (30) Gradulus on Tl con- Tliis subgenus is closely related to Nodiiln. (33) Anterior surface of Tl of male without a fovea. as discussed below. 1894: 232. (55) Penis valves apically expanded. all and Rhysoxylocopa. even though in the discussion he stated that he left the subgenus Perixylocopa intact. \^b3: 219. except in Xylocopa eiythrina. (6) Female margin of female mandible except has three teeth. by the combination of the complete on the outer apex of the hind tibia. (39) Pregradular areas on S3-S5 of males with paired patches of long setae. (5) for X. Type species: Xylocopa fraiuiulentn Gribodo. (10) Male with yellow facial maculations. (42) Gonostylus of male genitalia distinct although fused to gonocoxite. not extending to near base of tibia. 1841. rufitarsis. (32) Gradulus only on Tl of female except in X. Xylocopini 31 DIAGNOSES AND DESCRIPTIONS OF SUBGENERA OF XYLOCOPA The following descriptions are presented in the order used for characters in the Character List and Coding section.and hind trochanter not modified. All species are African. the only species placed in Dinoxylocopa and males. (12) Outer apex of female hind tibia with one spine. bv Maa. enjthrina which has graduli on T1-T5. erythrina and X. capitata and X. rufitarsis. rufitarsis. Type species: Xyhcopn lugiibris Gerstaecker. not produced except in X. New synonymy. Dhioxuhivpa Hurd and Moure. albifrons. bv original designation. Males can be distinguished from other members of the tribe lack of spines this is t"lared plate. numbers are the same as those used there.Subgenera and Genera of Carpenter Bees. Type species: Xylocopa rufitarsis bv original designation. although. 1963: 226. (38) Medial carina on sternum of females weakly developed. (41) Pygidial spine of female with pair of preapical lateral spines. Unique apomorphies are in bold face type. Subgenus Xylomelissa Hurd and Moure XylonwUiSii without a medial carina. (14) Hind trochanter of male subcylindrical. Synonymy with Xylomeliffa in part. 1894 by original designation. erythrina. (52) Ventroapical plate of gonocoxite rounded. (45) Penis entirely membranous. Acwxylocopa Hurd and Moure. 1963: 232. medially projecting lobe. 1857. . beginning of thoracic declivity undefined. shares of the known only from this apomorphies of group and so should be included in Xylomelissa. (31) Graduli present on Tl to T5 of male. 1968:106. (3) Upper carina of female mandible without a tooth. Female metapostnotum present except in X. (21) Female basitibial plate Lepeletier. 1938: 290. (44) Penis valve apex not hirsute. (46) Spatha absent. except in X. Subgenus Nodula Maa Nodida Maa. except in X. (9) Ocelli not enlarged. Synonymy with Xyloinelissa Epixxflocofa (19) Apex of female basitibial plate bifid. rufitarsis and X. In some analyses presented here the Oriental subgenus Biluna was placed in this group. (22) Thoracic profile of male smoothly rounded. except in X. 1903. beginning of thoracic declivity undefined. Aphm/hccpii Hurd and Moure. by original designation. Maaiaua.

X. (11 Outer apex of male hind tibia with one spine. Hurd and Moure. Description. Females of Maaiana can be distinguished from those of the subgenus Rhysoxylocopa by the presence of a carina dorsal to each lateral ocellus and a sharply rounded bentoni. X. by 1963: 178. Nodula. Natural History Museum.ales can be distinguished by two features of the mid-tibia: a dense mat of short.) and Rhysoxylocopa. (39) Pregradular areas of male sterna without lateral setal patches. as scutellum.32 Scientific Papers. was given in Hurd and Moure (1963). Rhysoxylocopa. that Nodula. 1841. (23) Beginning of posterior thoracic declivity of female abruptly rounded. Type species: Xylocopa imsalis Westwood. Hurd and This subgenus far as Moure (1963) considered Bilu)ia to be closely related to the subgenus Xylocopa. as known. Etymology. lat. ametln/stiiia. anterior mar- determined by deeply impressed line. (19) Metapostnotum of female absent. (26) Male without a mesosomal gland ) — . Associated with the very smooth inner surface of this nesting substrate are morphological niodifications of the legs and mandibles. —As in Xylomelissa." exemplars of both were included my analysis. Nodula is closely related to Maaiana. remota. Fem. Females of Rhysoxylocopa can be distinguished from those taxa by the absence of a medial groove on the propodeal triangle. (22) Beginning of posterior thoracic declivity of male abruptly rounded. ramakrishnai Biluna Maa. The evidence suggests subgenus as a Outer apex of male hind tibia with one spine. except as follows: and Nodula. male absent. Type species: Xylocopa cantabrita Lepeletier. madurensis. lower extremity broadened. 1938: 270. although the male genitalia are phenetically similar. except as follows: (11) two "natural groups. Maaiana consists of the following species. forming an elongate triangular flap whose lower anterior margin projects strongly forward. A key to the species. X. X. (12) Outer apex of female hind tibia with two spines. tranquebawrum. nasalis and X. Males can be distinguished from those of all other taxa by the combination of lacking a basitibial plate and of spines on the outer apex of the tibia. asymmetrical. angulosa. and X. Xylomelissa (s. (16) Male hind femur without basal tubercle or spine. X. X. punctigena This group is closely related to Xylomelissa. As in Xylomelissa. original designation. C. (49) Gonostylus apex entire. (13) Female hind tibia with tibial spine strongly developed. male on propodeum in X. except as follows: plate of (25) Basitibial plate of female with edges slightly raised from tibia. stout setae and absence of spines on the outer apex. as presently recognized. as the punctigena group of Nodula. bicristata. This leaves Nodula consisting of the following species: X. A is a group of large. (49) Gonostylus apex entire. 1842." gin. modified from Maa (1938 and 1954). Males oi Maaiana can be distinguished from those of Nodula by the abrupt beginning of the posterior thoracic declivity on the scutellum. was presented by Maa (1954) and in modified form by Hurd and Moure (1963). (27) Beginning of thoracic declivity of Subgenus Maaiana new subgenus Type species. this subgenus is clearly allied scutellum. Males are easily distinguished from all other taxa by the unusual propodeal spiracles which Hurd and Moure (1963) described as "small. (49) Gonostylus entire. without medially projecting lobe. meyeri. elongate bees that. p^lmnerocephala. is either a paraphyletic or polyphyletic assemblage and have recognized the group typified by X. The males of Nodula can be distinguished from those of Xylomelissa by the presence of a spine on the outer apex of the hind tibia of the male and from those of Rhysoxylocopa by the unmodified propodeal spiracles. The females of Nodula can be distinguished from those of the subgenus Rhysoxylocopa by carinae dorsal to the lateral ocelh and a sharply rounded and X. (18) Basitibial Apex of female basitibial plate entire. 276. (13) Hind tibial spine of female strongly developed. I found no characters that define members this of monophyletic group. Maaiana and Biluna. Description. except as follows: (10) Male face entirely black. (24) Male metapostnotum evident. (26) Mesosomal gland reservoir present. (11) In most analyses. (16) Male hind femur without basal tubercle or spine. nigwtarsata. metanotum in X. prashadi. The University of Kansas is Maa (1938) noted that this subgenus represented by in Description. Xylocopa bentoni Cockerell 1919: 172. (26) Mesosomal gland reservoir absent. — As in Xylomelissa. nest solely in bamboo. with a clade consisting of Xylomelissa. X. punctilabris. from Xylomelissa by the presence of a spine on the outer apex of the hind tibia. Maa for his important work on Old World Xylocopini. X. In my analyses. surface of —As in Xylomelissa. key to them and those in the subgenus Maaiana. originating well before tibial apex and with medial carina. X. Biluna is shown as the most (17) Outer apex of male hind tibia with one spine. Biliina Maa Maaiana. without a medial lobe. basal taxon of this clade. —The name Maaiana is in recognition of T. X. Subgenus Rhysoxylocopa Hurd and Moure Rliysoxylocopa The relationship of this group to Nodula and allied subgenera is discussed above under Nodula. auripennis. As defined herein. Description. and from Rhysoxi/locopa by the unmodified propodeal spiracles. without medially projecting lobe. bentoni as a separate subgenus 1 Subgenus by original designation.

(52) Ventroapical plate of gonocoxite absent. scalelike process. (52) Veiitroapical plate of gonocoxite weakly developed. Oenoxylocopa is apparently most closely related to some members of the subgenus Xylocopa (sensu stricto). except as follows: (4) Apical edge of female mandible tridentate. (40) metasomal spiracles. diameter greater than that of antennal socket. (2) Ocelli enlarged in some species. 193. (26) Mesosomal reservoir of male present. (43) Penis valve with lateral projection. Both sexes of the species of Nyctomelitta can be distinguished from all other members of the genus Xylocopa by their large size and greatly enlarged ocelli. Maaiana. the recognition of Proxylocopa as a genus makes Xylocopa paraphyletic. Pwxykwfm Aita/kicopa {Proxyhcopa) Maa. Type species: Apis fcnesfrata Fabricius. i954: 190. ovate. Apex of gize with structures of other Xylocopini are not generi- from the other taxa in the tribe. Description. 1798. Xylccofn iProxyhccpa) Hedicke. Those characters of Proxylocopa that 1 can homoloProxi/locopa lies primarily group is not well established by Although seemingly allied to a clade consisting this study. olroieri (=Xiihxopa ctivieri Lepeletier. olivieri of Proxylocopa (sensu stricto) and all species of Ancylocopa. posteriorly directed. 1878. (39) Pregradular areas of male sterna without setal patches. Subgenus ProxyJocopn Hedicke. Proxylocopa. 1938: 192. of the majority of New World species. New status Brll. Subgenus Ctenoxylocopa Michener Ctempoda Maa. and create an unequivocally monophyletic genus Xylocopa by assigning subgeneric rank to cally distinct . (11) Outer apex of male hind tibia with one spine. by original designation. 1938. (39) Pregradular area of male 33 without lateral entire. new name for Ctenopoda Maa. (50) Medial ventral margins of gonocoxites roughly parallel distad from ventral gonocoxal bridge. without medially projecting lobe. the ocellar diameters and the malar space grade continuously together. This subgenus face in females as this feature notable in having a distinctively long compared members of the tribe. (16) Male hind femur without basal tubercle or spine. synonymize Ancylocopa with the subgenus Proxylocopa. (40) Male without patches of short appressed setae near metasomal spiracles. the distinctiveness of with the structures associated nesting. (16) Male hind femur without basal spine or tubercle. Furthermore." of short appressed setae near metasomal spiracles. 198. Males of this subgenus are distinctive. and (3) hind tibia with one spine on its outer apex. (25) Female metapostnotum absent. Examination of other species in the subgenus should help to clarify these matters. Ocellar diameter and length of the malar space were the primary Baam Sandhouse 1943: 530. by original designation. Gonostylus apex without Male without patches medially projecting lobe. (11) mandible with two teeth on its apical edge. era. widely of antennal socket. 1929: 303. —As in Xylomelissa. (26) Mesosomal gland reservoir consisting of two small. some structural features and its biogeography suggest that this group is closely related to Kliysoxylocopa and its allies. Males can be recognized by the set of characters including short. riifa and P. and two spines on the outer apex of the hind tibia. Xylocopini reservoir. netic relationship of this Description. Nodula. (49) Gonostylar apex entire. (3) Upper carina of female mandible with subapical tooth. The phyloge- no more that three times as long as wide. (9) Ocelli enlarged. The male genitalia suggest that this group is most closely related to Copoxyla and/or the clade oi Rhysoxylocopa Biluna.^4: 116: 190. except as follows: (9) bercles along each lateral edge. (10) Male face . Proxylocopa (sensu stricto) and Aucylocopa. by original designation. 1841). Type species: Xyhcopa nilidiventrh Smith. 1938. is New synonymy. 1938: 270. (39) Pregradular areas of male sterna without setal patches. to other by original designation. and Xylomelissa. Subgenus Nyctomelitta Cockerell Ni/chvnclitta Cockerell. diameter greater than that Outer apex of male hind tibia with two spines. Based on my examination of P. 1933). 19. new name for Cteiwpodn Maa.Subgenera and Genera of Carpenter Bees. Females are easily distinguished by the well-developed basitibial plate located at the base of the hind tibia and by the welldeveloped pvgidial plate. (40) Male without patches ot short setae near (49) separated pouches. As discussed above. 285 (preoccupied by Ctenopoda McAtee and Malloch. characters used I eral lobes of the pronotum are prolonged posteriorly so groove on T3 bear an elevated. Type species: Bomlms tranqucharicn Fab- Maa. and there is morphological and phylowith soil genetic evidence that these structures have evolved secondarily. Type species: "X. 1804. (32) Female with graduli on T1-T5. the posterolat- by Maa (1954) to distinguish these groups. setal patches. obvious that there is no clear distinction between the two presently recognized subgenit is Ctenoxylocopa Michener. — As in Xylomelissa. —As in Xylomelissa. ricius. Therefore. 1942: 282. and the spiracles of: (1) the well-developed basitibial plate with a row of tu- Description. Females can be distinguished from all other groups by the combination that they are nearly in line with the preepisternal on the mesepisternum. (32) Graduli on T1-T5 of female. (18) male basitibial plate bifid. I favor emphasizing the homogeneity of the tribe. The recognition of Proxylocopa as a genus requires that it be the sister group to the rest of the genus Xylocopa. parapsidal lines. probably is associated with loss of the strong mandibular musculature for gnawing in wood. except as follows: (2) Upper carina on male mandible with subapical tooth. If Proxylocopa is more closely related to any of the derived subgenera of Xylocopa.

Xylocopa Latreille. Males can be distinguished from all other Old World groups by the combination of face without yellow maculations. Females of both species can be distinguished from all other members of the tribe by the combination of the two spines on the outer apex of the hind tibia and the tridentate apical edge of the mandible. (24) Male metapostnotum present. (40) Metasomal spiracle without associated setal patch. (48) completely fused to gonocoxite in X. dentate. cyanea. californica arizonensis. Generic Names in Opinion 743. (24) Male metapostnotum present. (48) Gonostylus slender and digitiform. 1738. (17) Lateral edges of female basitibial plate with- Description. Xilocopa Latreille. (32) Gradulus present on T1-T5 of female.. (55) Penis valves slender. (16) Male hind femur without basal tubercle or spine. (55) Penis valves slender Subgenus Xylocopoides Michener Xylocopoides Michener. (39) Pregradular areas on male sterna without paired setal patches. Description. (52) Ventroapical plate of gonocoxite absent. (53) Mesal margin of venter of gonocoxite with tooth or projection. Females share with the subgenus . imlga. 1802:431 Rejected Comm. Comm. (18) Upper carina of female mandible without a tooth. (41 Pygidial plate margined by two rows of many preapical lateral spines converging at male hind tibia base of pygidial spine. (42) Gonostylus completely fused and indistinct from. (17) Anterior and posterior edges of female basitibial plate each with row of tubercles. The subgenus Xylocopa is in Upper carina of female mandible without tooth except in (8) Malar space of female shorter than width of segment 1 except in X. flagellar or spine. Nomencl. 1954: 105. (35) Medial groove linear on Tl of female except in X. valga probably is most closely related to the Old World subgenus Ctenoxylocopa. with very few apical setae. (11) Outer apex of with two spines. (52) Ventroapical plate of male gonocoxite absent. (40) Setal ) patches absent near cept in X. 1963: 142. long. Smith 1874. (49) Male hind tibia with one outer apical Male liind femur without basal tubercle or spine. and the lack Xylocopoides Xylocopa (sensu stricto) probably spect to Xylocopoides. 1840:86. (48) Gonostylus slender and digitiform. (42) Gonostylus of male genitabasitibial plate bifid. with very few se- This is a monotypic subgenus and probably derived from Xylocopoides. gonocoxite. i>alga. The University of Kansas tae at apex. (50) Medial ventral margins of gonocoxites roughly parallel beyond ventral gonocoxal bridge. except X. slender. 1965. with very to. (31) Tl and T2 of male with graduli. (31) Graduli on Tl and T2 of male. (16) Male hind femur without basal tubercle X. (18) Apex of male basitibial plate bifid. recognizable. except as follows: tri- Apex of male basitibial plate bifid. outer apex of the Latreille (sensu stricto) by Internatl. pygidial plate on the surface of T6 and a foveate basitibial Males can be distinguished from those of tibia. and the two spines on the outer apex of the hind tibia. —As in Xylomelissa.34 Scientific Papers. (52) Ventroapical plate of male gonocoxite absent. (3) —As in Xylomelissa. (39) Pregradular areas of male sterna without paired setal patches. by the two spines on the Subgenus Xylocopa . violacea shows clear affinities with the New World subgenus Xylocopoides. violacea. Type species: Apis vir^itiicn Linnaeus. both share several highly derived synapomorphies and of a distinct gonostylus. (32) Graduli present on T1-T5 of female. (20) Female basitibial plate closed basally (21 ) Female basitibial plate reduced. ex(4) without paired setal patches. of numerous preapical spines delineating the all without medially projecting lobe. Gonostylus slender and digitiform. 1771. (49) Gonostylar apex entirely black. (11) spine. (11) Outer apex of male hind tibia with two spines. (36) Apical margin of SI medially emarginate. Natural History Museum. fused but and with few setae in X. (24) Male metapostnotum present. (18) Apex of Male metasomal spiracles of male. by original designation. (17) Lateral edges of female basitibial plate each with a row of tubercles. Emended name placed on Official List of Zoology by Internatl. The two species used in this study suggest that this subgenus is paraphyletic or polyphyletic. whereas X. (36) Apical margin of SI medially emarginate. the dark blue metallic coloration. (53) Ventral surface of gonocoxite with basal angle or spine. few apical setae. (10) Male with yellow facial maculations except most specimens of X. (12) Outer apex of female hind tibia with two spines. the rounded profile of the posterior part of the thorax. (44) Penis valve apex hirsute. (12) Outer apex of female hind tibia with two spines. (36) Apical margin of SI medially emarginate. violacea. (39) Pregradular areas on sterna of males out tubercles. (21) Female basitibial plate extending to near base of tibia. (26) Mesosomal gland present. other Xylocopini. except as follows: (3) need of further study before its phylogenetic affinities can be clarified. not extending to base of tibia. by sub- Although 1 have left and Xylocopa (sensu stricto) as separate groups.. (26) Male mesosomal gland present. Opinion 743. Zool. Type species: Apis violaceae Linnaeus. (10) Apical edge of mandible of female face entirely black. (55) Penis valves slender Subgenus CaUoxylocopa Calloxylocopa Hurd and Moure Type species: Xylocopa Icnualii Hurd and Moure. (49) Gonostylar apex without medially male Female basitibial plate extending to near base of tibia. 1965. 1802:379. cyanea. is paraphyletic with re- sequent designation of Westwood. Nomencl. valga. Zool. Females of the subgenus Xylocopoides uniquely possess two rows plate. by original designation. X. (54) Male T7 with pair of apical dentiform processes. (16) Gonostylar apex without medially pro- jecting lobe. (21) lia projecting lobe.

(39) Pregradular areas on male sterna without paired setal patches. Females are unique in having a large. well-developed with numerous convoluted tubules. 1828: 795. among Description. and by the presence of a defined metapostnotum. iris Female mandible with tooth on upper carina. (52) Ventroapical plate of male gonocoxite absent. (26) Reservoir of mesosomal gland of male absent. the second submarginal cell of the wing which is very nearly a parallelogram and not attenuate on the posterior edge as in all other Xylocopa.. Description. (54) Male T7 with pair of apical dentiform processes. and indistinct from. (32) Graduli present on T1-T5 of female. (49) Gonostylus of male with a weakly developed medially projecting lobe. (16) Male hind femur without basal tubercle or spine. Subgenus Gniithoxylocopa Hurd and Moure Gnatiwxylocctpn Hurd and Moure 1963: 182. and covered by long hairs that originate on its anterior margin. and strong metallic integument of both sexes. The gonostylus of the male genitalia is almost completely fused to the gonocoxite and has a spinelike medial lobe at the . gonocoxite. and Type — 182. Nevertheless. (39) Pregradular areas on sterna of males without paired setal patches. (21) Female basitibial plate extending nearly to base of tibia. (48) Gonostylus slender and digitiform. — Subgenus testis Lepeletier and Serville. (26) Reservoir of mesosomal gland present. by origi- subgenus Copoxyla are among the smallest of the Xylocopini and are closely related to Ctenoxylocopa. (16) Male hind femur without basal tubercle or spine. (31) Graduli on Tl and T2 of male. Vachal. (54) Male T7 with pair of apical dentiform processes. 1898. except as follows: (4) (7) Apical edge of female mandible tridentate. (40) Male metasomal spiracles without nearby setal patches. (50) Medial ventral margins of gonocoxite roughly parallel beyond ventral gonocoxal bridge. (18) Apex of male basitibial plate bifid. (55) Penis valves slender. New status testis Lepeletier species: Apis bombylans Fabricius. (49) Gonostylar apex Apex of male basitibial plate bifid. Type species: Apis nal designation. (52) Ventroapical plate of male gonocoxite absent. relationships sidered as tentative. which is depresseci on the anterior half. by original designation. with very few apical setae. These features and two others the entirely black face of the male. (31) Graduli present on T1-T3 of male. Bull.1963). (32) Graduli on T1-T5 of female. (44) Penis valve apex hirsute. swollen tubercle on the parocular area below the antennal socket. (ID Outer apex of male hind tibia with one apical spine. 799. (17) Anterior and posterior edge of basitibial plate of female each with row of tubercles. Males of Copoxyla can be distinguished from Ctenoxylocopa bv their small size. hut can he distinguished from that group is open basally and barely Males are easily distinguished by the third sternum. Subgenus Copoxyla Maa Cofwxyta Maa. and by the bifid apex of the basitibial plate. Nomencl. This group seems to be relatively plesiomorphic and most closely related to the long. 1779 (Opinion 657. (39) Pregradular areas on male sterna without paired setal patches. forming two deep depressions. (52) Ventroapical plate of gonocoxite absent. rises abo\'e the leg surface. (11) Male by the four-segmented maxillary palpi. the posterior thoracic declivity begins on the posterior half of the metanotum). (24) Male metapostnotum present. 1954: 191. (4) Apical edge of female mandible tridentate. (41) Description. (10) Male face entirely black. (36) Apical margin of SI medially emarginate.e. (41) Pygidial plate margined by two rows of numerous preapical lateral spines converging at base of pygidial spine. and the presence of three teeth on the apical margin of the mandible. 800. (36) Apical margin of SI medially emarginate. (42) Gonostylus of male genitalia fused completely to. until cies of Ctenoxylocopa. 1791. (17) Anterior and posterior edges of female basitibial plate each with row of tubercles. (31) Graduli on T1-T3 of male. (24) Male metapostnotum present. (18) Apex of male basitibial plate bifid. The species of the margins of gonocoxites roughly parallel beyond ventral gonocoxal bridge. (17) Both lateral edges of basitibial plate barely rising (18) — Preapical spines of female pygidial spine absent. the absence of preapical spines beside the pygidial spine. and the single spine on the outer apex of the hind tibia will differentiate males of this group from all others. except as follows: (3) without medially projecting lobe. Females can be distinguished by the combination of a smoothly rounded thorax. face entirely black. As in Xi/louicli:^sn. except as follows: (10) Face of male entirely black. pits deep. (16) Male hind femur without basal tubercle or spine. Xylocopa. without tubercles. (50) Medial ventral Christ. Female basitibial plate extending nearly to base of tibia. (11) Outer apex of male hind tibia with one apical spine. (21) Females of Giiathoxylocopa are easily distinguished by hook that arises from the ventral margin of the mandible. (55) Penis valves slender. —As in Xylomelissa. 20:181Serville. more speand other allied groups are these taxa should be con- examined.. Type species: Xylocopn sidifli above tibial surface. (10) —As This group of two Australian species is distinguished in Xylomelissa. curved Ctenoxylocopa and Copoxyla. Supraocellar Outer apex of male hind tibia with one apical spine. Zool. Xylocopini Xi/locopoidcs the 35 numerous preapical spines margining which the pygidial plate. (36) Apical margin of SI medially emarginate. Males can be distinguished by the lengthened thoracic dorsum (i. 211.Subgenera and Genera of Carpenter Bees. by the basitibial plate.

macrops and X. Spatha present. Type species: Xylocopa bambusae by original designation. guatemalensis. Type species: Xylocopa micans Lepeletier. (26) reservoir absent. Both of This is a speciose New World are brightly metallic and fast-flying. within this group are uncertain. Misspelling of Schonherria Lepeletier. except (3) and Calloxylocopa. originating before tibial apex and with a longitudinal medial carina. Type species: Xylocopa chrysopoda Hurd and Moure. 1902. based on three male genital characters (strongly developed medial lobe of the gonostylus. My analysis does not place this group firmly with any other subgenus. (39) Pregradular areas on male sterna without paired setal patches. (50) Medial ventral margins of gonocoxites roughly parallel beyond ventral gonocoxal bridge.36 Scientific Papers. Schoniherria 1841. (24) Male metapostnotum present. (52) Ventroapical plate of gonocoxite absent. Subgenus Schonnherria Lepeletier Schonherria Lepeletier. (17) Anterior and posterior edges of female basitibial plate weakly developed or rising little from surface of tibia in X. a strongly bifid apex of the gonostylus and the spine on the ventral apex of the gonocoxites. by original designation. 1902. (55) Penis valves apically slender. (40) Setal patches near male metasomal spiracles not present. loxylocopnj 1963:109. tribe. 1841: 207. New synonymy. would make Mesosomal gland on T1-T4 of male. It is clearly a if recognized as a genus. (16) Male hind femur without basal tubercle except in X. with tubercles along Description. two spines in X. Therefore. (7) Supraocellar pits of female punctiform or deeply excavated in X. and Xylocopsis these taxa possess the define Schonnherria (sensu 1 likely that Notoxylocopa is the sister group to a group that as subgenera. Schonherria Lepeletier. (20) Female basitibial plate foveclosed basally.e. Description. (11) Outer apex of male hind tibia with one spine. hence 1 consider it two strongest synapomorphies that Hurd and Moure. the weakly developed ventroapical plate of the gonocoxite suggests this relationship is not close. (3) Tooth on upper carina of female mandible. (26) Mesosomal gland reservoir present. except as follows: (11) Patches of setae near male metasomal spiracles absent. (12) Outer apex of female hind tibia with two spines. thickly hirsute apex of the tribe. 1943: 598. 1 841 : 207. However. Hurd and Moure (1963) considered Notoxylocopa to be most closely related to the Old World subgenus Rhysoxylocopa. (36) Apical margin of SI medially emarginate. 1963). 1956: 2. i. a unique feature in the This group. the Neoxylocopm group and the Schonnherria group are more firmly established. Dasyxylocopa. Ashmead. subgenus. (17) Anterior and poste- . (50) Medial ventral margins of gonocoxites roughly parallel beyond ventral gonocoxal bridge. I consider testis to patches occur in some subspecies of X.. New synonymy. although setal lateral edges in X. and especially the outward projection on the penis valves). Most species but some species are decidedly drab and hairy. except as follows: (2) Subapical tooth on upper carina of male mandible absent except in X. but not bifid. 1841 by designation of Sandhouse. They can be distinguished from all other New World groups by the presence of graduli only on Tl in females and by the bifid apex of the gonostylus and a large spine on the ventroposterior margin of the gonocoxite in males. (32) Graduli present on T1-T4 of female. until the relationships between Notoxylocopa. (43) Conspicuous carina on outer edge of penis valve. includes the rest of the Xylocopoides New World Xylocopa. —As in Xylomelissa . and Xylocopsis) erected by Hurd and Moure (1963). micans. Dasyxylocopa. (36) Apical margin of SI medially emarginate. although its affinities relict group. (52) Ventroapical plate of male gonocoxite weakly developed. except as follows: Tooth on upper carina of female mandible. The University of Kansas both apex. Natural History Museum. As defined here. (49) Medially projecting lobe of gonostylar apex well developed. Description. (32) Gradulus present on T1-T5 of female. 1854. guatemalensis. Schrottky. tabaniformis. Schrottky. or spatha. Hurd and Moure. Subgenus Notoxylocopa Hurd Notoxylocopa Hurd. viridigastra which has three. 1899: 71. This New World subgenus is defined by the posses- sion in males of an elongation of the dorsal bridge of the penis valve. micans. 1963: 116. (31) Graduli present the genus Xi/locopa paraphyletic. differ from Schonnherria primarily in the number of graduli on the terga of the female and may eventually be shown to be plesiomorphic members of the Schonnherria clade. Xylocospila male genitalia with dense setal patch. Type species: Xylocopa tnbaniformis Smith. —As in Xylomelissa. gonostylus. this group is expanded to include Xylocospila and loxylocopa. Three other monotypic subgenera (Nanoxylocopa. (17) Basitibial plate not margined with tubercles. (48) Gonostylar apex of ate. viridigastrn. recognized previously as a genus. (49) Gonostylar apex with medially projecting lobe. seems to be most closely related to the other groups allied to Xi/locopa (sensu stricto). (54) Apex of T7 of male with pair of dentiform projections. (39) Pregradular areas on male sterna without paired setal patches. but. (46) Outer apex of male hind tibia with one spine. (4) Two teeth on apical margin of female man- — dible except in X. (40) be a subgenus of Xylocopa. However. have retained Nanoxylocopa. by original designation. tabaniformis. tabaniformis. it could be most closely related to the New World group that includes Schonnherria. As in Xylomelissa. a feature shared with no other mem- ber of the Females can be distinguished from all other New World Xylocopa by the absence of a defined metapostnotum. although. (11) Outer apex of male hind tibia with one spine in X. (13) Tibial spine of female strong.

by original designation. entire in X. The phylogenetic position of this group is discussed under Schonnherria. (50) Medial ventral margins of gonocoxites roughly parallel beyond ventral Subgenus Dasyxylocopa Hurd and Moure Dasyxylocopn gonocoxal bridge. (26) Mesosomal gland reservoir not examined. Subgenus Xyhcopsis Hurd and Moure Xylocopsis Subgenus Cirroxylocopa Hurd and Moure Cirroxylocopn Hurd and Moure 1963: 124. rising little from surface of tibia. Burmeister. (54) Gonocoxite with a conspicuous Apex of male T7 Friese. (16) Male hind femur without basal tubercle. forming a bifid apex. (49) Mediallv projecting lobe on gonostylar apex elongate. (43) Lateral edge of penis valve with strong carina. Hurd and Moure 1963: 113. Type species: Xylocopa cilinta from males of that group by the greater number of graduli in Dasyxylocopa (T1-T5) than on Schonnlwrria (Tl only). (18) Apex of male basitibial plate bifid. (36) Apical margin of SI medially emarginate. (40) Patches of appressed setae near spiracles of male metasoma absent. (40) Patches of appressed setae near spiracles of male metasoma absent. (16) Male hind femur without basal tubercle or spine. rising little from surface of tibia. (54) Apex of male T7 with pair of dentiform projections. (7) Supraocellar pits of female punctiform. Xylocopa vestita Females of Xylocopsis can be distinguished from all other New World taxa by the possession of five tubercles on the Hurd and Moure (1963) suggested that Cirroxylocopa was a basal taxon. of the lineage that . (50) Medial ventral margins of gonocoxites roughly parallel beyond ventral gonocoxal bridge. (54) Apex of male T7 without pair of dentiform projections. by original designation. (36) Apical margin of SI medially emarginate. 1912. The phylogenetic position of this group is discussed under Schonnherria. (43) Strong carina present on lateral edge of penis valve. Tooth on upper carina of female mandible. (51) Gonocoxite with conspicuous spine on posterior ventral margin. (17) Anterior and posterior edges of female basitibial plates weakly developed. (26) Mesosomal gland reservoir present as a single layer of parallel-nmning tubules. by original designation. 1963: 102. 1963) shape of the frontal carina in females. (43) Lateral edge of penis valve with strong carina. (26) Mesosomal gland reservoir present. 1903. (51) Gonocoxite with conspicuous spine on posterior ventral margin. (4) Two teeth on apical margin of female mandible. (31) Gradulus on Tl of male. (26) Mesosomal gland reservoir present. — As in Xylomelissa. except as follows: (2) (3) Upper carina of male mandible with tooth. (40) Patches of appressed setae near spiracles of male metasoma absent. except as follows: (3) emarginate. viridigastra. or rising little from surface of tibia. (35) Medial groove of female Tl linear. (39) Setal patches of pregradular areas of male sterna absent. Description. (36) Apical margin of SI medially rior 37 labrum. or abruptly rounded in X. Males can be distinguished by the combination of a distinctive yellow stripe on the scape. (40) Patches of appressed setae near spiracles of male metasoma absent. Xyi. (51) Gonocoxite with a conspicuous spine on posterior ventral margin. (17) Anterior and posterior edges of female basitibial plates weakly developed. except in X. Type species: Xyhcopsis ftmesta Maidl. Hurd and Moure. an abruptly rounded beginning of the thoracic declivity and two spines on the apex of T7. (36) Apical margin of SI medially emarginate. by original designation. (11) Outer apex of male hind tibia with one spine. bniiibnsac. except as follows: (3) Upper carina of female mandible with tooth. (18) Male basitibial plate bifid at apex. (32) Gradulus on Tl of female. (50) Medial ventral margins of gonocoxites roughly parallel beyond ventral gonocoxal bridge. (23) Beginning of female posterior thoracic declivity smoothly rounded. nuKwpf. 1963. (50) Medial ventral margins of gonocoxites roughly parallel beyond ventral gonocoxal bridge. the sexes of this with pair of dentiform processes. near Nanoxylocopa. As noted by Hurd and Moure species are distinctive because of the thick pubescence. —As in Xi/Iomelissa.ocopini edges of female basitibial plate weakly developed. (39) Setal patches of pregradular areas of male sterna absent. This subgenus is similar to Schonnherria but can be distinguished Subgenus Nanoxylocopn Hurd and Moure Nanoxylocopa Hurd and Moure. (43) Lateral edge of penis valve with strong carina. (51) Type species: Xylocopa bimaculata spine at posterior ventral margin. This monotypic subgenus can be distinguished from the other New World groups by the evenly punctate tegula Description. rising little from surface of tibia. (31 ) Tl of male with gradulus. viridigastrn. 1976. — As in Xylomelissa. rina of female der Schonnherria. (49) Medially projecting lobe on gonostylus apex weakly developed. Type species: Hurd and Moure. Description. Tl and T2 in X. (1963). 1963: 99. (11) Outer apex of male hind tibia with one spine. (11) Outer apex of male hind tibia with one spine. (32) Tl and T2 of female with graduli. micans. (32) Graduli present on T1-T4 of female.Subgenera and Genera of Carpenter Bees. (17) Anterior and posterior edges of female basitibial plates weakly developed. (16) Male hind femur without basal tubercle. deeply impressed in X. (39) Setal patches of pregradular areas of male sterna absent. (39) Setal patches absent on pregradular areas of male sterna. Upper ca- males and by the "transversely tuberculiform" (Hurd and Moure. The phylogenetic position of this group is discussed unin mandible with tooth.

(40) Setal patches present near spiracles of male metasoma. not with a dense brush at its apex. (16) Male hind femur without basal tubercle. except as follows: (3) Tooth on upper carina of female mandible. the short clypeus and the shape of the — — tegula. —As in Xylomelissa. nificollis. Females can be distinof the tribe by the combi- Xyhcopina Hurd and Moure. 1963: 105. by original designation. by original designation. except in X. 1963: 160. —As in Xylomelissa. rise to the Schonnherria it is Natural History Museum. except as follows: (3) Subgenus Xylocopoda Hurci and Moure Xylocopoda Upper elegaiis Hurd and Moure. (50) Medial ventral margins of gonocoxites roughly parallel beyond ventral gonocoxal bridge. (11) Outer apex of male hind tibia with one spine. by original designation. (43) Lateral edge of penis valve with strong carina. nificollis. males of all species have two parallel yellow stripes running from the vertex of the head to the base of the mandibles. Mcf^axylocopm Hurd and Moure. rising little from surface of tibia. consisting of several layers of discshaped tubules. (51) Gonocoxite with conspicuous spine on posterior ventral margin. 1960: 809. (48) Gonostylus with few setae. guished from a continuous other members subgenera. Type species: Xylocopa nificollis Hurd and Moure. (27) Beginning of thoracic declivity of male on propodeum. (49) Medial lobe on apex of gonostylus absent. (27) Beginning of thoracic declivity of male on propodeum. nificollis. Type species: Apis frontalis Ohvier. (17) Anterior and posterior edges of female basitibial plate weakly developed. distinguishes females of this group from all others in the tribe. —As in Xylomelissa except as follows: (3) carina of female mandible with tooth.)iV/'rns/7/(im)n(m Linnaeus. New synonymy. Stenoxylocopa Description. (36) Apical margin of SI medially emarginate. Description. apex of SI and yellow markings restricted to the lower part of the face and two autapomorphies. Subgenus Neoxylocopa Michener Neoxylocopa Michener. (49) Apex of gonostylus without medial lobe. (49) Medial lobe of gonostylar apex absent. (17) Anterior and posterior margins of female basitibial plate weakly developeci. Males can be distinguished uniquely by the presence of a high. (11) Outer apex of male hind tibia with one spine. 1874a. (51 ) Gonocoxite with a conspicuous spine on posterior ventral margin. 157-158. (51) Gonocoxite with a conspicuous spine on posterior ventral margin. Except for Xylocopina nificollis. leading to the Neoxi/lacopa lineage. by original designation. impunctate ridge along the epistomal suture. (17) Anterior and posterior edges of female basitibial plate weakly developed. which are abruptly raised near the lower margins of the eye. Type species: Hurd and Moure. entire and not produced in X. (50) Medial ventral margins of gonocoxites roughly parallel beyond ventral gonocoxal bridge. (50) Medial ventral margins of gonocoxites roughly parallel beyond ventral gonocoxal bridge. (32) T1-T5 of female with graduli. My This group is exclusively New World. by original designation. but not produced. 1963. (12) Outer apex of female hind tibia with two spines. (6) Female epistomal suture continuously carinate and broadly impunctate. Xylocopa This is a monotypic subgenus.38 gave Scientific Papers. Type species: /i. (35) Medial groove of Tl of female linear. Xylocopoda is the sister group to and Neoxylocopa. (26) Mesosomal gland reservoir not examined. The University of Kansas and Neoxi/locopa clades. Description. except in Xylocopina nificollis. (38) Medial carina on feniale sternum continuous across all segments. Upper as wide or wider than the lower tooth. (4) Female mandible with three teeth on apical margin. . (48) Gonostylus with few setae. without dense brush of setae at its apex. (6) Female epistomal suture continuously carinate and broadly impunctate. apex of pollex) fused and forming a continuous edge. (11) Outer apex of male hind tibia with one spine. rising little from surface of tibia. Females also are easily identified by the parocular areas. 1789. (40) Patches of appressed setae near spiracles of male metasoma absent. (43) Lateral edge of penis valve with strong carina. (36) Apical margin of SI medially emarginate. (16) Male hind femur without basal tubercle.e. (40) Patches of appressed setae near spiracles of male metasoma absent. (26) Mesosomal carina of female mandible with tooth. 1963: 151. Type species: Xylocopa artifex New World all Smith. New synonymy. In my analysis. rising little from surface of tibia. (27) Beginning of thoracic declivity on male propodeum. and most spe- analysis suggests that the basal member of the clade cies are Neotropical. gland reservoir present. (43) Lateral edge of penis valve with strong carina. (39) Pregradular areas on male sterna with setal patches. Subgenus Stenoxylocopa Hurd and Moure stenoxylocopa This subgenus uted of the is the most speciose and widely distrib- Hurd and Moure. (32) Tl and T2 of female with graduli. 1954:155. (16) Male hind femur without basal tubercle or spine. Xylocopina nificollis was considered to be subgenerically distinct from Stenoxylocopa by Hurd and Moure (1963) based on two plesiomorphies the entire. (39) Setal patches of pregradular areas on male sterna absent. thin lamella that begins near the base of the hind tibial spur and extends obliquely forward and across the ventral surface of the tibia. (26) Mesosomal gland reservoir not examined. An enlarged upper mandibular tooth. (5) Upper rutellar tooth and preapical tooth (i. nation of two teeth on the apical margin of the mandible. present but not continuous in X. 1767.. Males can be distinguished by the combination of two spines on the outer apex of the hind tibia and the tegula being impunctate on its posterolateral third.

(40) Setal patches present near spiracles of male metasoma. rising little from surface of tibia. Description. (52) Ventroapical plate of gonocoxite present. (22) Beginning of posterior thoracic declivity of male abruptly rounded. (26) reservoir not examined. which represented by a single. other . Recognition of Mcgaxi/locopn makes the subgenus thoracic declivity ( 1*^63) Mesosomal gland on T1-T5 of female. (16) Male hind femur without basal tubercle. (49) Gonostylar apex without medially projecting lobe. (52) Ventroapical plate of gonocoxite weakly developed. Zonohirsuta is that both of the lateral Description. on the propodeum. dcjccmii Lepeletier. without row of tubercles. Hurd and Moure based the subgenus Mc.^iix\/locopn on their extremely large size and conspicuous carinae over the lateral ocelli. (22) Beginning of posterior thoracic declivity of male abruptly rounded. (48) Gonostylus slender. 1841). male on propodeum. (26) Reservoir of male mesosomal gland extremely large and consisting of many layers of highly convoluted tu- An additional apomorphy of metanotum. (23) Beginning of posterior thoracic declivity of female abruptly margin. both on the pregradular areas of the metasomal sterna and near the metasomal spiracles. (47) Gonostylar designation. (19) Female basitibial plate apex entire. rounded. (40) Setal patches near male metasomal spiracles absent. by original is 215. Females can be distinguished by the abruptly rounded beginning of the thoracic declivity is and by the basitibial plate. (51) Gonocoxite with a bifid spine originating at Description. The subgenus Zoiioliirsiitn is similar to Prosopoxylocopa. (43) Lateral edge of penis valve with strong carina. —As in Xylomelissa. (23) Beginning Females of this subgenus can be distinguished from all members of the Xylocopini by the combination of long malar space (longer than the width of first flagellar segment) and two spines on the outer apex of the tibia. —As in Xi/loiuclissn. 1939: 155. and by two spines on the outer apex of the tibia. bridge. 1841. 1963: 203. Type species: Xylocopa colltuis Lepeletier. sometimes visible through c1ecli\'ity cuticle of (27) Beginning of thoracic of edges of the female basitibial plate are strongly carinate and project perpendicularly from the tibial surface. Subgenus Prosopoxylocopa Hurd and Moure ProS'Opoxi/locopn mirabilif without paired setal patches. (11) —As in the Xylomelissa. (16) Male hind femur without tubercle or spine. (38) Medial carina on sternum of female continuous across all segments. (17) Posterior edge of female basitibial plate with carina arising perpendicularly from leg. except as follows: (1) Male vestiture on dorsum of body entirely tawny or fer- the ventroposterior margin (Fig. (6) Female epistomal suture a continuous ridge and broadly impmictate. (24) Male metapostnotum present. Males are distinguished by their yellow vestiture (most species) and the beginning of the posterior 39 and a strong of posterior thoracic declivity of female abruptly rounded. Type species: Xylocopa bomhoides Smith. (32) Graduli present of gonocoxites roughly parallel from ventral gonocoxal Ncoxi/locopn paraphyletic. 9C) and the continuous yellow maculations from the labrum to the supraclypeal area of the face of the it male BomboixylocopiJ 1879. New synonymy. (17) Posterior edge of female basitibial plate with carina arising perpendicularly from leg.ocopini medial carina along the entire sternum of the metasoma. (17) Anterior and posterior edges of female basi tibial plates Subgenus Zonohirsuta Maa Zonohirsiitn Maa. (48) Gonostylus with few setae. Mimoxylocopa Hurd and Moure. This subgenus likely the sister group to the remain- der of the Ethiopian clade. without row of tubercles. Riginous (most species). (12) Outer apex of female hind tibia with two spines. except as follows: Outer apex of male hind tibia with two spines.Subgenera and Genera of Carpenter Bees. Males can be distinguished by the presence of appressed setae. by original designation. (39) Pregradular areas of male sterna without paired setal patches. The strongly produced projection of the posteriormost ventral part of the gonocoxite (Fig. 9C). 1789 (= X. with a carina. 300. Type species: Xylocopa nifipes Smith. (16) Male hind femur armed with basal spine. weaklv developeci. (51) Gonocoxite with a conspicuous spine on posterior ventral Outer apex of male hind tibia with one spine. (39) Pregradular areas of male sterna without paired bules. (50) Medial ventral margins of gonocoxites roughly parallel beyond ventral gonocoxal bridge. (50) Medial ventral margins of gonocoxites roughly parallel beyond ventral gonocoxal bridge. 1852. Xyi. (50) Medial ventral margins (24) Male metapostnotum present. (36) Apical margin of SI medially emarginate. very strongly projecting carina. (12) Outer apex of female hind tibia with two spines. Subgenus Bomboixylocopa Maa Maa. 1963: Hurd and Moure. (26) Mesosomal gland reservoir present. They differ by the basally open basitibial plate in females of Zonohirsuta and by the single spine on the outer apex of the hind tibia of males. not Olivier. (39) Pregradular areas on male sterna near male apex without dense setal patch. without dense brush of setae at its apex. (49) Medial lobe on gonostylar apex absent. Type species: Xylocopa metasomal spiracles absent. (40) Setal patches Hurd and Moure. by original designation. 1938: 270. except as follows: (11) setal patches. (3) Tooth on upper carina of female mandible. (12) Outer apex of female hind tibia with two spines. (11) Outer apex of male hind tibia with one spine. distinguish from all other large carpenter bees. (32) Graduli present on T1-T5 of female. by original des- ignation.

(33) Anterior surface of Tl of male with fovea. New synonymy. extending to propodeal pit. Xenoxylocopa and/ or Alloxylocopa are/is the sister group to Mesotrichia and Koptortosoma. (50) Medial ventral margins of gonocoxites roughly parallel beyond ventral gonocoxal bridge. 1908. on posterior ventral margin. (11) Two spines on outer apex of male hind tibia. (16) basal spine or tubercle. chinensis. Gonostylus lacking dense setal patch on apex. Plntynopoda Westwood. (36) Apical margin of SI medially emarginate in X. (7) Supraocellar pits of female normal. (32) Graduli present on T1-T5 of female. (26) Mesosomal gland reservoir present. forming a keel. (30) Gradulus of Tl nearly coincident with lateral ventral margin of metasoma. The subgenus Alloxylocopa is similar to Xenoxylocopa but by the lack of preapical spines beside the pygidial spine and in males by the dark coloration and presence of a fovea on the anterior face of Tl. (53) Gonocoxite with projection along inner mesal margin of base. Emendation of Plntynopoda Westwood Hoplitocopa Lieftinck. New synonymy. (48) Gonostylar apex lacking dense setal patch. Auiiiuctia Lepeletier. Xylocopa acutipennis Smith. Type species: Xylocopa appendiculnta Smith. nifipes. by original designation. (34) Ante- rior surface of Tl of females with fovea. except as follows: (1) tawny or ferruginous over entire dorsum Female epistomal suture with a continuous — smooth ridge. synonym of Platyiwpoda by designation of Sandhouse. this group is often placed as the sister group to Mesotrichia and Koptortosoma. (1 7) Anterior and posterior edges of female basitibial plates rising little from surface of tibia. (23) Beginning of posterior thoracic declivity of This subgenus is the sister group to the subgenus all Koptortosoma. 1899: 71. 202.40 Description. (25) Female metapostnotum present. Females can be distinguished from all other members of the tribe by the abruptly rounded beginning of the posterior thoracic declivity and by the presence of a pair of preapical spines at the base of the pygidial spine. (49) Gonostylar apex without medially projecting lobe. (26) Mesosomal gland reservoir not examined. (3) Upper carina of female mandible without tooth. by original designation. 1854. chinensis. As in Xenoxylocopa. 1840: 271. 1939: 155. 27. by designation of Ashmead. (See section on Relationships among Members of the Major Glades and Taxonomic Recommendations. (8) Malar space of female longer than width of first flagellar segment. (40) Setal patches near male metasomal spiracles absent. Type species: Mesotricliia torriiia Westwood. (41 ) Pygidial spine absent. 1880. by original designation. Type species. 9D). (39) Pregradular areas of male sterna without setal patches. (47) Gonostylus of male slender and elongate. differs in females Subgenus Xenoxylocopa Hurd and Moure Xenoxylocopa Hurd and Moure. (6) As in Xylomelissa. (53) Gonocoxite with projection along inner mesal margin of base. (22) Beginning of posterior thoracic declivity of — chiyakensis Cockerell. (28) Propodeum at its connection with by the perpendicular angle of the be- ginning of the thoracic declivity. (48) Maa. (51) Gonocoxite with spine on posterior ventral margin. Type species. 1896. base in X. 1943. over half as long as gonocoxite (Fig. (35) Medial groove of female Tl deeply sulcate. except in Xylocopa riifipes. 529. (7) Supraocellar pits of female deeply excavated and craterlike. Subgenus Mesotrichia Westwood Westwood. (49) Gonostylar apex without medially projecting lobe. (31) Gradulus only on Tl of male. rounded apex of female hind tibia. rufipes. Females can be distinguished from large carpenter bees other female abruptly rounded. (12) Two spines on outer Male hind femur without and posterior edges of female basitibial plates rising little from surface of tibia. Mesotrichia Male vestiture of body. (28) Propodeum at its connection with metasoma not elongated. 1963: 260. chinensis. forming a spine. (35) Medial groove of Tl of females deeply sulcate. over half as long as gonocoxite (Fig. (19) Apex of female basitibial plate entire. (30) Gradulus of Tl obliquely curved on anterior ventral margin of metasoma. (19) Female basitibial plate apex entire. these spines present in X. ntfipes. 1773. —As in Xi/lomelissa. Apis latipes Drury. absent in X. chinensis. 1 955. 1773. by original designation. 1838: 112. or asymmetrically bifid in X. divergent in X. New synonymy. Hoploxylocopa Hurd and Moure. 1841: 203. except in X. male abruptly rounded. Type species: Apis latipes Drury.) Description. Males can be distinguished by the combination of sharp beginning of the declivity of Tl (as occurs in Koptortosoma and Mesotrichia). (51) Gonocoxite without a spine and the nearly entirely punctate tegula. lack of a fovea on the anterior surface of Tl. 1838. in Jardine. original description. (24) Male metapostnotum present in X. the deeply excavated (craterlike) supraocellar pits and triangular hind tro- . In the phylogenetic analysis. (37) Medial carina on S6 of male strongly developed. (11 ) Two spines on outer apex of male hind tibia. 9. The University of Kansas metasoma dorsally elongated. (25) Female metapostnotum absent. nifipes. (17) Anterior in X. Platinopoda Dalla Torre. Scientific Papers. (47) Gonostylus slender and elongate. Type species: Xylocopm assimilis Ritsema. (41) Basal pair of pygidial spines present. except as follows: (1) Male vestiture black or dark brown on sides and pale on dorsum. (50) Medial ventral margins of gonocoxites roughly parallel beyond ventral gonocoxal bridge. (16) Male hind femur without basal spine or tubercle. nifipes. (41) Pygidial spine lacking lateral spine at Subgenus Alloxylocopa Maa Alloxylocopa 1852. Type species. Mesotridua Description. Natural History Museum. (12) Two spines on outer apex of female hind tibia. except as follows: (2) Upper carina of male mandible without tooth. 1963: 243. (51) Gonocoxite with a spine on posterior ventral margin. 9D). (36) Apical margin of SI entire and produced.

This is a structurally diverse group chanter. Xylocopini Males can be distinguished by the posteriorly elongate tegulae (Fig. Cyphoxylocopii Hurd and Moure. and sclero- tized basally beneath in X. Maa. assimilis. ocularis. by original designation. rounded lobe in X. acKtipcniiis. Cyaneoderes Ashmead. acutipennis. assimilis. (53) Upper carina of a tooth. acutipennis. (24) Male metapostnotum absent. New synonymy. in (Among specimens of X. Type species: C\yaneoderes] fairchildi Ashmlead. but not so in X. and X. except in X. (See section on Relationships the Major Clades among Members of and Taxonomic Recommendations. (29) Male tegula elongated posteriorly. acutipennis. acutipennis. assimilis. Koptorthosoma Dalla Torre. by original designation. Unjustified emendation of Koptortosoma Koptortcisonm Gribodo. (7) Supraocellar pits of female deeply excavated and craterlike. (37) Medial carina on S6 of male present but not strongly developed. 1899: 70. Males share the tegula. (50) Medial ventral margins of gonocoxites roughly parallel beyond ventral gonocoxal bridge. latipes and X. (23) Beginning of posterior thoracic declivity of female a transverse carina not extending posteriorly beyond metanotum. forming a spine in X. —As in Alloxylocopa. torrida and X. Maiella Michener. 1943: 561. Upper carina of female mandible without a tooth. 1894. except in X. 1963: 275. latipes and X. which has a spine. (42) Gonostylus of male genitalia slen- Male vestiture blue in Xylocopa caerulea to fer- pubescens. except as follows: tawny or segments. The apomorphies identified herein and by Hurd and Moure all (1963) that de- fine the monophyly of Oxyxylocopa. (54) Apex of T7 of male entire. (3) but still recognizable in X. Cyaneoderes.) (25) Female metapostnotum present in X. torrida X. 1901. acutipennis. latipes and X. completely undifferentiated from gonocoxite in X. Upper carina of female mandible with tooth except in X. and broadly fused pubescens. assimilis. except in X. widely separated pouches that are not developed into tubules in X. X. 1841. torrida. —As in Alloxylocopa. (10) Male with yellow facial macula tions except in X. 41 as the description indicates. latipes. is the sister group to Mesotrichia (sensu The pronounced transverse carina on the scutellum that completely overhangs the metanotum of females of most species distinguishes these taxa from females of all This subgenus lato). (41 Pygidial spine lacking preapical lateral spines at base. sinensis.and hind trochanter of Gonocoxite without projection on inner mesal margin except in X. Afroxylocopa Hurd and Moure. assimilis. assimilis. 1938. T1-T3 in X. latipes. torrida. cius. nigrita and X. Unjustified emendation of Koptortofoma Gribodo. New synonymy.Subgenera and Genera of Carpenter Bees. nigrita. male without spine or tubercle except in X. tenuiscapa and X. 4B). except as follows: male mandible without (3) (3) Ventroapical plate of gonocoxite present or absent in X. torrida and X. acutipennis. 1854. without this projection in X. 1896: 202. ined. assimilis. Type species Xylocopa varipes Smith. 1775. acutipennis. Subgenus Koptortosoma Gribodo Type species. (16) Male hind femur with basal spine in X. latipes. (22) Beginning of posterior thoracic declivity of male sharply angled. (30) Gradulus of SI virtually coincident with anterior ventral margin of metasoma. assimilis. by original designation. Copforthcsoiim Perez. 1938.) (1) carina on sterna of female continuous and present on all Description. Apis nigrita FabriOrbitclla . the (34) Anterior surface of Tl of female with entrance to a mite pouch. torrida. der in X. except in X. (55) Penis valves slender and parallel. torrida. present but not continuous in X. Oxyxylocopti Hurd and Moure. 1915). X. assituilis but dorsally elongated. 305 (not OrbitcUa Douville. acutipennis. except vestigial. torrida. caerulea and X. with few setae in X. ocularis. 1899] [= Bombus caeruleus Fabricius. assimilis but produced only slightly as a emphasized. tenuiscapa 1 which it is have exam- metapostnotum is exposed in some individuals and absent in others. acutipennis and only Tl in X. torrida and X. torrida and X. and highly convoluted mass of tubules in X. yet a number of characters provide evidence for its monophyly. (48) Apex of gonostylus setose. (2) Male mandible without tooth on upper carina. New synonymy. nigrita. (38) Medial Gribodo and Koptortlwsoma Dalla Torre. is an equivalent taxon to Koptortosoiua: by combining the four subgenera recognized by Hurd and Moure (1963). 1942: 50:282 Replacement name for Orbitelln Maa. torrida and X. and Afroxylocopa are highly homoplasious and do not justify recognizing these taxa as separate sub- genera. (7) . nigrita. two. (26) Reservoir of male mesosomal gland present and variable. assiiuilis. latipes. 1963: 283. torrida. KofHortosoma galwnica Gribodo. other large carpenter bees. (52) Description. except in X. (31) Distribution of graduU on male terga variable: T1-T5 in X. (51) Gonocoxite with strong spine from posterior ventral margin in X. latipes and X. Euryapis Sandhouse. (45) Penis entirely in X. Cyphoxylocopa. as recognized herein. 1901:3. Type species. (47) epistomal suture without continuous ridge and devoid of setae. by original designation. (6) Female membranous and X. with dentiform projections in X. 1894: 271. caffra ruginous over entire dorsum of body in X. Xylocopa ocularis Perez. by designation of Sandhouse. 1963: 264. (6) Epistomal suture of female continuously carinate and smooth in X. divergent in X. New synonymy. 1938: 270. latipes and X. the evolutionary relationships of the species are Gonostylus of male slender and less than half as long as gonocoxite (also see Character 42 above). latipes and X. but absent in X. latipes. caendea and X. Type species. (28) Propodeum at its connection with metasoma not modified in X. Mcsotricliin. 1943: 551 Replacement name for Orhitella Maa. (15) Mid. many features with those of Mesotrichia and can be separated most easily by which is not modified in Koptortosoma and is elongated posteriorly in Mesotrichia. latipes and X. (14) Female hind trochanter roughly triangular in ventral view. X. except in Xylocopa assimilis. in X.

pubescens. Amniote phylogeny and the importance of fossils. caendea. 1793. except in X. varipes. M. ocularis. Gottsberger. Fabricius. H. Annals and Magazine of Natural Fiistory (10)4:296-304. M. (32) Graduli present only on Tl of female or Tl and T2 in X. caffra. Enderlein. J. Leipzig: Engleman. Hedicke. E. except in X. 1440]+[l]-30. University of California Publications in Entomology 108:1-^9. 100. H. 1969. D. 1976. The carpenter bees of California.. (30) Gradulus on SI virtually coincident with anterior ventral and X. Drury. S. Akademie der Wissenschaften. F. T. Frankfurt am Hafniae: Proft. pubescens and X. October 1857. Botanische Jahrbiicher fiir Systematik. D. nigrita.. Berliner Entomologische Zeitschrift 48:45-64. Hirashima. Descriptions and records of bees. S. R. Jr. Entomologist 41:34-36. Friese. G... C. deeply excavated in X. [Bees and wasps collected in Mozambique). Fitzhugh. C. Carpenter. (54) T7 of male with apex entire or with pair of dentiform spines on X. 29:460^64. A bee-polJ. X. Observations on the nesting habits of some New World carpenter bees with remarks on their importance in the problem of species formation. 535 pp. J. 1894. Pflanzengeschichte und Ptlanzengeographie 109:469-500. 1899. 1938. Systematic Zoology 18:374-385. Annals of the Entomological Society of 832 pp.and hind trochanter of male without spines. K. Catahgus Hymenopterorum Vol. Cambridge 95:655-666. Neue Arten der Bieneiigattung Xylocopa Latr. nigrita. Fabricius. Deutsche Entomologische Zeitschrift 1938:186-196.. Cladistics 4:105-209. D.. Hurd. with basal spine in X. (50) Medial ventral margins of gonocoxites roughly parallel beyond ventral gonocoxal bridge or divergent in X. 1919. 1903. (53) Gonocoxite with projection along inner mesal margin of base. 1955. (37) Medial carina of S6 of male strongly developed. 1775. except in X. Fabricius. (19) Apex in X. D. D. linated tropical community: The beach dune vegetation of Ilha de Sao Luis. Corbet. Pollination of the yellow passionfruit: nectar. (26) Reservoir of male mesosomal gland a highly convoluted. Copenhagen. P. nigrita. L. except in X. (27) Beginning of thoracic declivity of male on metanotum. X. puhescens and X. P. Cockerell. nigrita. sinensis. and I. pollen and carpenter bees. Bulletin of the California Insect Survey 4: 37-72. 1992. 10. Anthophoridae). Gribodo. G. 1988. de. Main: Hermann.. nigrita. ocularis. varipes and X. A. (28) Propodeum at its connection with metasoma dorsally elongated forming a spine except in X. Farris. Daly. Gautnier. 1929. ocidaris LITERATURE CITED Ashmead. with setae at of male sharply angled. nigrita.42 Scientific Papers. (34) Anterior surface of Tl of female with entrance to mite pouch or with fovea in X. (12) Outer apex of female hind tibia with one spine. Synopsis of the genus Pithitis Klug of the world (Hymenoptera: Anthophoridae). which it is on the propodeum. except in X. 1956. Cockerell. 1959. G. D. Notes on the subgenera of the New World carpenter bees of the genus Xylocopa. Bees in the collection of the United States National Museum. (Distributed by the author. (15) Mid. Y. Bees of the new genus Ctenocerntina in Africa south of the Sahara (Hymenoptera: Apoidea). nigrita. Republic of South Africa Department of Agriculture. pubescens. A. many-layered mass of tubules. T. 1988. Systema Entomotogie. Proceedings of the United States National Mu- Cockerell. iiigrita and X. G. S. varipes. Cunha. Braunschweig.) Port Farris. Jr. Some nomcnclatorial problems in the genus Xylocopa . Natural History Museum. D.49- Brothers. nigrita and X. varipes. 1802 (Hymenoptera. 1908. caendea. (39) Pregradular areas of male sterna with paired setal patches. (25) Female metapostnotum absent or present in X. 1857. besonders aus der Ausbeute des Freiherrn von Erlanger in Galla und Somalica. Gottsberger. S.C. Berlin. except in Male with yellow facial X. 2. Systematic Entomology 1:177-182. Some bees collected by Dr. ocularis. New York. Classification of the bees. except in X. American Museum Novitates 1776:1-7. J. Klnssificntio)i und Nomenclature der Insekten vom Bienen-Wespeti-und Ameisengeschlecht. Cladistics in the fast lane. varipes. in apex. (22) of female basitibial plate entire. caendea. J. A.. (47) Gonostylus of male genitalia less than half as long as gonocoxite. Syst Systema Piezatorum. 1896. The University of Kansas margin of metasoma. C. 1988. Journal of the New York Entomological Society 97:234-241. (14) Female hind trochanter roughly cylindrical. 1989. forming a keel.. (10) maculations. C.. Comparing phenetic structures: An application to subgenera of Xylocopa Latreille. Rowe. Brazil. Wilmer. 1969. Note imenotterologiche.. America 51:365-375. Cladistics 4:291-296. Jr. absent in X. Modification of the metapostnotum and origin of the propodeal triangle in Hvmenoptera Aculeata. P. viii+643 pp. Fabricius. Nnturgeschichte. P. caffra. p^d^escens and X. viii+519 pp. sinensis. 1988. 1983. varipes. Christ. Neue und weniger bekannte afrikanische Xylocopen. W. chinensis. D. V. 1773. Pacific Insects 1 1 :649-669. except Beginning of posterior thoracic declivity in X. Uber einige Apiden vom Hindu-kusch. ocularis. Dalla Torre. J. A. BuUettinodella Societa Entomologica Italiana (Florence) 26:262-314.). J. Entomologia Systematica Emendata et Aucta. (23) Begin- preapical lateral spines at base. 1791. Journal of Agricultural Science. Kluge and T. 1980. C. and lobe producing a bifid point in X. Choosing among multiple equally parsimonious cladograms. Maranhao. (29) Graduli present on male Tl and T2 or only on Tl in X.5. A. (48) Gonostylus lacking dense setal patch on apex. Hurd. (41) Pygidial spine lacking pair of Supraocellar pits of female punctiform. A taxonomic revision of the genus Xylocopa Latreille (Hymenoptera: Anthophoridae) in southern Africa. Gerstaecker. sinensis. xiv+[15]- A successive approximations approach to character weighting. C. except ning of posterior thoracic declivity of female a transverse carina that overhangs entire metanotum but rounded in X. D. Zeitschrift fiir systematische Hymenopterologie und Dipterologie 3:202-208. Vol. 1804. Monatsberichte. CXIX. version 1.. aus der neotropischen und orientalischen Region (Hym. Nota 11. 1988. except in X. J. G. A. London: White. 3. Hurd. (51) Gonocoxite with spine on posterior ventral margin. D. Wellman in West Africa. Entomologiae Systematicae Supplementum. A. H. Hennig86. Reyista Brasiliera de Entomologia 36: 465-473. xxvii+90 Jefferson. Transactions of the American Entomological Society 26. F Camargo. nigrita. (40) Setal patches present near metasomal spiracles of males. or the superfamily Apoidea. Flensburg and Leipzig: Korte. J. Eardley. Jr. 1903. (49) Gonostylar apex with medially projecting lobe in X. X. (35) Medial groove of Tl of females deeply sulcate. Nuovi generi e nuove specie di Imenotteri antofili ed isservazioni sopra alcune specie gia conosciute. Illustmtioiiti of Natural History. G. (16) Male hind femur without spine or tubercle in most species. caerulea and X. 572 pp.J. A. 1798. seum 55:167-221. and P. C. except in X. not triangular in ventral view. X. 1958. H. Hurd. Entomology Memoir 58:1-67. D. T.

-K. Journal of the Entomological Society of South Africa Zoologischen 37:261-281. 1968. Cladistics 7:337-343. D. Kjobenhavn 116:189-231. Duncan. Calcutta: Government of India. Abeille. J. Massachusetts.). xi-i-398 pp. Notes on the carpenter-bees fauna of the Soviet Union (Hymenoptera. and other additional matters by ]. Maddison. Y. Castes in xylocopine bees. F.I 975. Japanese Journal of Entomology 63: 347-375. Smith.D. 1984. D. pp. Monograph of the genus Xylocopa Latr Transactions of the Entomological Society of London 1854:247-307. Perez. Scientific Results of the Second Yarkland Mission: Based on the Collections and Notes of the late Ferdinand Stoliczka: Hymenoptera. 1901. The carpenter-bees (Xi/locopn Latr. Zoogeographical character of some Palearctic species of the bee genus Xylocopa Latr (Hymenoptera: Apoidea) and their interrelations with melittophilous plants. 1974. Vol. D. G.Subgenera and Genera of Carpenter Bees. C. Missing data versus missing characters in phyloJ. Estratto Dagli Annali del Museo Civico Reyes. 1943. Reyes. D. Verhandlungen der Naturforschenden Gesellschaft in Basel 66:5-32. 1954. 1879. Bulletin of the American of Natural History 104:1-176. Maa. 12th ed. Diderot et al. W. Museums. 1993.. Smith. and A. 1880. Anthophoridae. F. Reeder 1995. P. Pp. B.'(i Latreille (Hymenoptera: Apoidea). 1840.[= Maa] 1938. 10. A. A comparative anatomical study of the mandibular structure in bees. Subgenus Bomhoixylocopa novuiii. Insectes. and M. M. Outgroup analysis and parsimony. with systematic characters of each species. and D. Jr. Pp. Russkoe Entomol. A taxonomic study of African allodapine bees (Hymenoptera. Apoidea). 1879. Version 3. Lingnan Science Journal (Canton) 18:155-160. Lepeletier de Saint-Fargeau. C. Vol. Part 2. 1991. and D. T. Tlie Naturalists' Library. A. Westwood. C. T. F. T. Maeta. Observations on a nest aggregation of the Taiwanese bamboo carpenter bee Xylocopa {Biluna) tranquebarorum tranqueharoruni (Hymenoptera. Maa. Die Xylocopen (Holzbienen) des Wiener Hofmuseums Ein Beitrag zu einer Monographie dieser Gattung. Paris and Liege Vol. Paris. W. A. 1767. Michener. Xylocopa and halictine bees. London: Henry G. J. Catalo'^ue of the Hymenopterous Insects of the Britisli Museum. Uni\'ersity of California Publications in Entomology 29: 1 365. 1. Apoidea). 680 pp. pp: 1-334. University of Kansas Science Bulletin 54:179-207. Ritsema. Transactions of the Entomological Society of di Storia Naturale Genova 77:103-114. Serpentes). Entomologicheskoe Obozrnie 40:393^04 [In Russian]. 1913.. Combining data sets with different numbers of taxa for phylogenetic analysis. Trudy Instituta Zoologii. Westwood. Michener. Griswold. Descriptions of some new and apparently undescribed species of hymenopterous insects from North China. 1985. 1828. A New World subgenus of bamboo nesting carpenter bees belonging to the genus Xi//('ai. Westwood. A revision of type specimens of Xylocopine bees in the Genova Museum Maa. [In E.-c.. 1825. and T W. [Articles in] M. Pan-Pacific P. Xylocopa orientalia critica (Hymen. Revision of the bee genus Braunsapis in the Oriental Region. Museum 1 838. Malyshev. Donovan] Natural History of the Insects of India. Description of a new genus of exotic bees. N.-c. Hurd. Moure. in Encyclopedic Mcthodique. Bulletin of the American Museum 53:80^-821. S. J. Xylocopini Hurt). 1842. F... G. 1955. Kluge. A. Oophagy and egg replacement in artificially induced colonies of a basically solitary bee. I. A. Engels (ed.. Systematic Biology 42:576-581.). E. Sunda Islands and Taninibar (Hymenoptera. containing upwards of two hundred and twenty figures and descriptions.-c. 1-22. J. Coddington.-c. O. Journal of the Kansas Entomological of Natural History 155:67-240. in. 26. revision of the subgenus Ctenoxylocopa Anthophoridae). E Moure. 2nd ed. 1978. J. In J.enetic analysis. Weins. H. Lieftinck. P. T. de savans et d'artistes. Ceratina (Ceratinidia) okinawana (Hymenoptera. Edinburg: W.. B. 40:1-58 [in Russian with English summary]. J. The Indian species of the genus Xylocopa Latr. J. D. London (Hymenoptera: 2:33-48. Smith. the large carpenter bees. Social Insects: An Evolutionary Approach to Castes and Reproduction. A. Systema Naturae. Histoire Natiirelle. pp. M. Vol. E. H. Vol. A classification of Olivier G. V. J. Anthophoridae. 0. brought down to the present state of the science. A.). London: British Museum. A Maddison. Ma. Descriptions ofNeiv Species of Hymenoptera in the Collection Taschenberg. 1%3. Maddison. 1990. of the British Museum. D. collected by Robert Fortune. Smith. 1874. S. Bohn. and j. Vachal. Maa. 1954. P. R. 1898. and S. Linnaeus. Tlie Natural History of Bees. 1912. viii-i-17-301 pp. Naturhistorischen Hofmuseums 26:249-330. 1942. P. C. lusivtes. R. Videnskabelige Meddelelser fra Dansk Naturhistorisk Forening i Akademii Nauk SSSR 1:29-52 [In Russian]. University of Kansas Science Bul- Michener Society 51:46-76. Jr. J. 1991. A. and J. The type species of the genera and subgenera of bees. C. collection. A concern for evidence and a phy logenetic hypothesis of relationships among Epicratcf (Boidae. V. Zeitscnrift fur de Gesammte Naturwissschaft 52:563-599. 1878. 43 Entomologist 35:135-148. Notes Leyden Museum 2:220-224. 1852. University of Kansas Science Bulletin 55:97-122. Sinauer Associates.). Systematic Zoology 38:7-25. Serville. P. and ]. Smith. Esq. dison.) of the Lesser P. (Horae Societatis EntomologicaeRossicae). C. Sunderland. Lepeletier cie Saint-Fargeau. R. Euci/ch^pciiic Mctlwdiqiic. xxi-i-240 pp.ens de lettrcs. Annals of the Entomological Society of America Latreille.. The xylocopine bees (Insecta) of Afghanistan. S. and A. Materiaux pour une revision synoptique des especes Europeennes et Africaines du Xylocopa Latr Annales de la Societe Entomologique de France 67:92-99. O. Donoghue.G. F. 1995. Sakagami. 1789. 6.. Watmough. Records of the Indian Museum 40:265-329. Annalen K. Contribution a etude des xylocopes. Sandhouse. Actes de la Societe 1 Linneenne de Bordeaux (6)6:1-128. 1961. Eraser 1978.Verlag. Histoirc NntureUe des InsectesHyinciwvtcnv. Publiee par unc societc de i. 2. W. 30 pis. F. Bamboi>-nesting carpenter bees (genus XyUiccpa Latreille) of the subgenus Stcnow/lihopn Hurd and Moure (Hymenoptera: Anthophoridae). F. F. Transactions of the Entomological Society of London 2:112-113. Jardine (ed. Pacific Insects 12:723-752. Platnick. 1-264 pp. 123-146 in W. (Hymenoptera). 1841. cclxxxviii-Hccclxxii+331 pp. Westwood. Holmiae: Laurentii Salvii 12: 901-1068. Description of three new exotic species of the Hymenopterous genus Xylocopa. Sakagami. MacCladc: Analysis ofPhylogenyand Character Evolution. Maidl. synonyms. letin 51:463^82. Michener. Taxonomic observations on bees with descriptions of new genera and species. M.I 989.-c. W. 1%0. Michener. Systematic Biology 44:548- 559. G. and A. . C. Popov. Y. Latreille (ed. Xylocopinae). Jr. Apidae): A comparative and experimental study. On missing entries in cladistic analysis. A. Revision of the genus Braunsapis in the Australian Region. O. Systematic Zoology 33:83-103. a new edition. 1993. 199^65. Ponomareva. indexes. Biology and behaviour of carpenter bees in Southern Africa. Schiokawa. M. Bees of Panama. London: British Museum. Ceralinini). Hisloire Naturcllc. Berlin: Springer. S. M. L. Hurd. Life and instincts of some Ceratina-bees (Hymenoptera. Proceedings of the United States National Museum 92:519-619. Entomology. D. Lizars. Journal of the Kansas Entomological Society 58:3641. Journal of the New York Entomology Society 50:273-282. 1939. S. 4. Popov. L. 1992. Anthophoridae). 1854. Paris: Roret. 1970. T. Maddison. Die arten der Gattung Xylocopa Ltr des Halle' schen Maeta. with a comparison of social behavior among Ceratina. 1-102. 1947.

2. 52. Penis valves M . 20. 22. 39. numbered in the text. Penis valves basal margin of venter of gonocoxite 33. NODES 50^49. M NODES 50. 12. M (1—>0). 12. 26. Medi(0—>1). M=male. 26. Mesosomal gland reservoir (2=>5). 53. Graduli on Tl-3 F (0=>2). The University of Kansas APPENDIX Character changes and node support for the tree shown Accelerated transformation 1 in Figure 10 A (Analysis 1. Epistomal suture F (1^0). Metapostnotum not defined F (0—>1). 1 Number of spines on outer apex of hind of thoracic declivity F 47. occur among sub- sequent nodes. Appressed setae near metasomal spiracles 54. 40. Supraocellar pit not deeply excavated F (1—>0). Pregradular areas of sterna with setal areas M {0— >1). NODES 73—>72. Facial macu- 76->62: 22. Penis 72-^71 23. 21 Basitibial plate reduced F (0—>1). Vesfiture (0=>1). Triangular hind trochanter F (0=>1). M M M (1 =>2). Supraocellar pit F (0->l). (0=>2). Epistomal suture F (0=>1 ). short M (0^1). Appressed setae near metasomal spiracles (0=>1). Shape of gradulus on Tl F (0—>1). Graduli on Tl-3 F metasomal sterna F {0=>1). 26. Subapical tooth on upper carina of mandible F (0=>1). 54. 32. Medial groove of Tl F (0—>1). ally projecting lobe on apex of gonostylus NODES 71 —^70: 3. apex F (1=>0). Upper rutellar tooth and preapical tooth F (0—^1). numbers in parentheses indicate character states (0=>1). 7. 49. (2=>1). (0=>1). Spine on ventral surface of hind femur M (0=>2). gonocoxite NODES 75->63: 8. 49. 38. Metapostnotum F (0—>1). character weights equal) M (ACCTRAN) of characters has been assumed. Gonostylus (0=>1). 11. Medial carina of metasomal sterna F (0—>1). Distribution of tergal graduli (2=>1). M metasomal spiracles 17. 48. 34. NODES 54^53: 18. Many spines along pygidial plate margin F (1=>2). ). 22. 51. Setae on apex of (1=>0). : M NODES 92—>91 17. Inner (1—>0). 7. Length of gonostylus (0=>1). M racles M (0=>1). NODES 80^79: NODES 94-^93: NODES 93->89: Upper 11 . Spine on posteroventral margin of gonocoxite (1=>0). 24. Metapostnotvim (1=>0). GraduH on Tl-3 F M (0—>1). 43. 44. 40. 55. Ventromedial (0=>1). edge of penis valve modified NODES 61^55: 36. Appressed setae near metasomal spi- M M M M M M (0=>1). 49. 28. 40. Subapical tooth on mandible (0—>1). Ventroapical plate of gonocoxite NODES 52—>51: 26. Appressed setae near M Tubercles on posterior margin of basitibial plate F M Metapostnotum 39. Tubercle on ventral surface of hind femur M (0=>1 NODES 83—>82. Malar space F (0-^1). 31. Appressed setae near metasomal spiracles (1 ^0). (1—>0). 47. NODES 87^86: NODES 86—>77: (0=>1). Tl (0=>1). 54. ning of thoracic declivity NODES 60->59: 6. Basitibial plate foveate F (0^1). Pregradular areas of sterna M (0—>1). 55. Arrows in parentheses with single lines (—>) indicate losses and gains in the character do not occur along subsequent nodes. 23. M NODES 84-^83: NODES 83-^78: 25. 3. Arrow Ln parentheses with double as they are lines (=*) indicate that reversals or gains NODES 74^73: 19. Apical margin of SI B (0->2). 12. Lateral (0=>1). NODES 76-^75. 42. Basitibial plate Apical margin of SI B (0=>1). Beginning of tho- racic declivity M (0=>1). Gonostylus (0=>1). M (2=>1). Spine on posteroventral margin of gonocoxite NODES 77-^61: 3. Spines on posterior margin T7 (0=>1). Node numbers are those shown in Figure 10. M (0=>1). Setae on apex of gonostylus M tibia F (0^1 ). Ventromedial margin of gonocoxites diverge (0=>1). One spine on outer apex of hind tibia . M (0-^1). NODES 66-^64: 16. M NODES 77^76: Two spines on (0—>1 ). . 31 NODES 94-^87: Metapostnotum not evident M >1). 31. 1. 32. Mesosomal gland reservoir (0—>1). Medially projecting lobe (1=*2). M NODES 93—>92: lations absent 4. Mesosomal gland reservoir (2-^3). NODES 72^66: 14. 36. Medially projecting lobe on apex of gonostylus M (0=>1 NODES 81 ^>80: 16. NODES 95—>94: 41. Two spines on of gonocoxite outer apex of hind tibia F (0=>1). 41 Marginal spines of pygidial plate F (1—>0). Metapostnotum not defined M (0—>1). Epistomal suture not raised F (1=>0). 45. 0. 24. Beginning (1-^0). B=both sexes. Setae on apex of gonostylus (1=>0). Medially projecfing (1=>2). NODES 53^52: 17. Medial carina of NODES 57^56. 35. 51. No spines on outer apex of hind tibia M (1=>0). Tubercles along margins of basitibial plate F (0=>1). 40. Ap- NODES M (0— 21. Modified margins of basitibial plate F (0->l). Number of spines on outer apex of hind tibia F (0=>1). lobe on apex of gonostylus NODES 55->54. Begin(0=>1).44 Scientific Papers. 40. Apex of mandible tridentate F (0=>1). 48. 53. gonostylus NODES 59-^58: 4. 47. Setae on apex of gonostylus M (1=>0). Mesosomal gland reservoir M (1-^2). Beginning of thoracic declivity M (0^1). Distribution of tergal graduli (0=>I). 24. Distribution of tergal graduli M (0=>2). 5. ing lobe on apex of gonostylus NODES 55^50: 32. Apex of mandible bidentate F and preapical tooth F (1—^0). (0=>2). Length of gonostylus (1—)0). 11. Beginning of thoracic declivity F (2=>3). Pregradular areas of sterna M (0=>1). Anterior face of Tl F (1—>2). Mesosomal gland reservoir M (0— 26. 25. Beginning of thoracic declivity F (1=>2). 49. NODES 61—>60: 26. and the direction of character state change. Two spines on outer apex of hind tibia M (1=>2). 53. Ventroapical plate of gonocoxite M (0=>2). 38. Mesosomal gland reservoir M distinct F (1=>0). 34. Apex of mandible tridentate F (0->l). Medially project(0=>1). M NODES 91—>90: 11. margins of gonocoxites diverge NODES 89-^88: 41. 37. 19. Subapical tooth on upper carina of mandible F (0=>1). Anterior face of M M M M M M Basitibial plate apex bifid F (0^1). Basitibial plate apex bifid (0=>1). Subapical tooth on mandible F (0=>1). 36. NODES 70->69: 6. 32. 29. rutellar tooth 4. Inner basal margin of venter M M (0=>1). slender. Anterior face of Tl F (0=>1). Mesosomal gland reservoir lost (2^0). 27. >1). Inner basal margin of venter of (0—>1). Tegula shape (0=>1). Natural History Museum. 30. outer apex of hind tibia Epistomal suture raised F (0=>\). 52. 23. 28. Propodeal tooth lost B (1^0). 50. pressed setae near metasomal spiracles NODES 75^74: 6. 26. Vesfiture M (0=5'1). M M Extent of basitibial plate F(1=>0). NODES 82^81 : ). NODES 66—>65: Metapostnotum 6. 40. 48. distinct (0=>2). Spine lost on posteroventral margin of gonocoxite : M M M M M M M M M(l^O). 49. NODES 58-^57: 32. 5. 39. on apex of gonostylus NODES 68—>67: 51. Medial carina of S6 (O-^l). Epistomal suture F (0=>1). 23. Paired marginal spines of pygidial plate F (0—>I). 48. Apical margin of SI B (0^2). Beginning of thoracic declivity (1=>2). (5^1). Beginning of thoracic declivity F (1—>0). Propodeal tooth B (0—>1). F=female. Spines on posterior margin T7 M (0— >1). 24. M NODES 86—^85: NODES 85—>84. NODES 69-»68: 1. Graduli on Tl-3 F (0=>2). Number of tergal graduli F (0=>1). Posterior margin T7 M (1—^0). M(0->1). Bifid apex of penis valves 6. 25.

Triangular hind trochanter F (0=>1). 53. Inner basal margin of (1—>0). 53. Lateral edge of penis 24. 39. Anterior face of Tl F (2=>1). 50. Ventroapical F (0—>1). 10. Facial NODES 63-^60: 13. Appressed M (0=>1). Mesosomal gland reservoir Epistomal suture not raised F (1=>0). 12. Medially projecting lobe on apex of gonostylus NODES 83^82: 6. NODES 91^77: NODES 77—>75: basitibial plate 24. 25. M M (1=>0). 40. Spine on ventral surface of hind femur M (0^2). 36. M Island 1. Beginning of thoracic declivity M (0=>2). Basitibial plate M (0=>1). 11 A (Analysis 3. 50. M (5^1). M (1=>0). M (0=>1). M Metapostnotum defined F 10. Two spines on outer apex of hind tibia F (1=>0). Two spines on outer apex of hind tibia F (0=>1). 48. NODES 53—>52: gonostvlus 6. Inner basal margin of Ventroapical plate of gonocoxite M M M NODES 72—>63: margin SI (0—>2). Ventroapical plate of gonocoxite (1=>0). 35. Graduh on T1-3'f (0=>2). gonocoxites diverge NODES 60—>59: 3. Marginal spines of pygidial plate F (1=>2). 2. Ventroapical plate of gonocoxite valve modified M M M (0->l). . Medial carina of S6 M (0=>1). Graduli on Tl-3 F metasomal sterna F (0=>1). NODES 69—>68: NODES 68^67: (0=5. Beginning of thoracic declivity F (2->3). diverge (0=>1). Medial groove of Tl F (0=>1).Subgenera and Genera of Carpenter Bees. Distribution of tergal graduli M (0^1). (0=>1). Two spines on outer tibia F (0=>1). venter of gonocoxite (1—>0). Appressed setae near metasomal spiracles M (0=>1). Apical margin of SI B (2^0). Subapical 23. Ventroapical plate of gonocoxite 39. 41. M M (0^1). short M (0^1). Spine on posteroventral margin of gonocoxite M M M M 49. on outer apex of hind tibia Metapostnotum F NODES 66—>65: 16. 34. Tibial spine F (1^0). NODES 52-^51 tooth 4. Medial carina of metasomal sterna F (0->l ). 25. (0=>2). Margins of maculations absent (0—>1). . 23. 36. Apex of mandible tridentate F (0—>1). 32. NODES 94—>93: 11. 32. NODES 90->89: NODES 89^88: 30. M {l->2). 36. M (0=>1). Mesosomal (0=>1). 1. 43. Shape of tegula M (0=>1). venter of gonocoxite NODES 91^90: 19. 11. : gonostylus M M (1=>0). Apex of mandible bidentate F and preapical tooth (l->0). Medially projecting lobe on apex M venter of gonocoxite M (0— >1). (0=>1). 29. 32. 47. Mediallv projecting lobe on apex of gonostylus M M NODES 56—^53: 26. 40. metasomal 24. 31. 27. 51 Spine on posteroventral 31 Distribution of tergal graduli M M M M M M . 51. M (0=>2). Medially projecting lobe on apex of gonostylus NODES 67-^66: 11. Ventromedial margin of (0=>1). Appressed setae near metasomal spiracles lations absent M M M M M : M M (0=>1). lobe on apex of gonostylus NODES 86^83: 1. 5. Vestiture {0=>1). 19. 52. Pregradular areas of sterna spiracles M (0=>1). 49. 44. 49. Apical 2b. Graduh on NODES Tl-3 F (0=>2). 21. 37. M Gonostvlus slender. 28. 37. 16. Mesosomal gland reservoir emarginate B (0=>2). 49. 25. 52. 51. posteroventral margin of gonocoxite NODES 88^87: 22. 26. M NODES 58—>57: NODES 57—>56: (1^0). Setae on apex of gonostylus (1—>2). Appressed setae near metasomal spi- M (0=>1). 39. Number 17. (0=>1). Upper rutellar NODES 90-^78. NODES 75^74: 21. Penis valves NODES 77—>76: 22. Marginal spines of pygidial plate F (1=>0). 28. M (0^1). Subapical tooth on mandible (0—>1).1). 52. Inner basal margin of Vestiture M (0=>1). 45. One spine on outer apex of hind tibia Basitihial plate apex bifid F (0—»1). 25. Distribution of ter6. Medial carina of S6 M (1=>0). 55. 51 Spine Divergence of ventromedial margin of gonocoxites M (1— on posteroventral margin of gonocoxite M (1^0). M (0=>2). 6. 50. Epistomal suture F Setae on apex of M (1=>0). Spines on posterior margin T7 M (0=>1). Subapical tooth on upper carina of mandible F C0=>1). Spine on (0=>1). NODES 62->61: apex bifid Margins of basiHbial plate F (0=>1). 26. 38. NODES 80-»79: F (1=>0). 53. 36. 33. ventral surface of hind femur gal graduli Metapostnotum not defined F (0=>1). Apical margin of SI B(2=>1). Mesosomal gland reservoir M (1^2). plate of gonocoxite NODES 75—>73: 12. of gonostvlus M (0=>1). Supraocellar pit F (0=>1). Shape of gradulus on Tl (0=>1). 12. Ventromedial margin of gonocoxites (0—>1). 25. 24. Gonostylus (2—>0). 5. 8. Penis valves 81->80: 26. Mesosomal gland reservoir (0=>5). tooth on upper carina of mandible F (0-»l). NODES 65—>64: 6. Unmodified (2^0). Beginning of thoracic declivity M M (0=>1). 17. 50. 49. Tibial spine F (0—>1). 41. Mesosomal gland reservoir >0). 52. M (0=>1). Length of NODES 59->58: 13. gonocoxites (1=>0). (0=>1 ). Medially (0—>2). Distribution of tergal graduli NODES 85^84: 34. 40. NODES 86^85: 31. NODES 70—>69: Metapostnotum not defined M (0=>1). NODES 92-^91: 48. Extent of basitibial plate F {1=>0). Metapostnotum not defined margin of gonocoxite M (1=>0). Supraocellar pit F (1=>0). Subapical tooth on mandible (0=>1). projecting lobe on apex of gonostylus NODES 55^54: 32. NODES 87-^86: 3. 7. 26. Apex of penis valves M M M (0->l). Appressed setae near metasomal spiracles (1—>2). NODES 63^62: 4. Anterior face of Tl F (0=*2). 26. Mesosomal gland reservoir (0^2). 32. 38. Extent of basitibial plate F (0^1). F (0=>2). Metapostnotum riot evident (0=>1). 47. Propodeal tooth B (0=>1). 52. Penis M (0->l). character weights equal) NODES 93->92: NODES 95—>94: (0—^1). Epistomal suture raised F (0=>1). 52. Apical margin of SI B (0^2). Beginning of thoracic declivity F (0=>1). Epistomal suture F (1=>0). Facial macu(0=>1). 18. 49. (0->l). Apex of mandible tridentate F (0=>1). 31. Pregradular areas of sterna vi'ithout setal areas NODES 56—>55: 2. Metapostnotum defined (1=>0). Paired margnial spines of pygidial plate F (0^1). Beginning of thoracic declivity (0=>1). Malar space F (0=>1). 36. 23. Two spines on outer apex of hind tibia M (1 ^2). Tubercle on ventral surface of male hind femur (0=>1). Anterior face of Tl NODES 51^50: NODES 50^49: (1—>0). 7. Apical margin of SI entire B (2=>0). Basitibial plate apex F (1^0). 17. of tergal graduli F NODES 72^71: NODES 71—>70: setae near Margins of basitibial plate (0—>1). Mesosomal gland reservoir M (2—>3). Medial carina of M (0^1). Metapostnotum not defined F (0=»1). Graduli Epistomal suture F (0=>1). Medially projecting (0—>1). 26. 16. 55. Xylocopini 45 APPENDIX 2 Character changes and node support for tree shown Abbreviations and conventions as in Appendix 1 in Fig. 54. Beginning of thoracic dechvity F (1^2). (0=>1). Ventroapical plate of gonocoxite (0=>1). Pregradular areas of sterna apex of hind racles and preapical tooth fused F (0—>1). 41. Length of gonostylus NODES 87^81 14. M (1—>2). Divergence of ventromedial margin of gland reservoir (0=>1). No spines (1=>0). 23. declivity F (0=>1). 40. 47. Metapostnotum defined Propodeal tooth absent B (1=>0). Beginning of thoracic on Tl-3 F (0=>2). Upper rutellar tooth 4. 40. 42.

Mesosomal gland Medially M NODES 56^55: 2. 51 Spine on posteroventral margin of gonocoxite (1=>0). NODES 82^81 10. 49. M M M M Marginal spines of pygidial plate F (1=>0). Mesosomal gland reservoir M (2—>3). character weights equal). 32. Two spines on outer apex of hind tibia F (0—>I). 47. Subapical tooth on upper carina of mandible F (0=>1). 35. 6. Epistomal suture F (1=>0). (0=*1). 32. NODES Metapostnotum not defined M (0=>1). NODES 90->89: NODES 89-^88: 16. 40. on apex of gonostylus M (0-^1). Medially projecting lobe on apex of gonostylus NODES 70^69: 6. Basitibial plate 1. NODES 77^76: 33. Ventromedial (0=>1). Triangular hind trochanter F (0=3 1). 52. 44. 52. 23. 39. Long malar space F (0=>1). 49. M M . M M M M ning of thoracic dechvity F (1—>2). 55. 25. Inner basal margin of venter of gonocoxite M (0=il). NODES 51^50: Number of apical teeth of mandible F (0=>I). Medial groove of Tl F (0=>1). Subapical tooth on mandible Appressed setae near metasomal spiracles M (0=>1). 49. Extent of basitibial plate F (1^0). 32. Natural History Museum. gland reservoir M (0=>2). Mesosomal gland reservoir M (2—>5). 23. Ventromedial margin of gonocoxites diverge M (0=>1). Ventroapical plate of gonocoxite M (1—>2). Subapical tooth on upper carina of mandible F (0=>1 ). Spine on ventral surface of hind femur (0^2). 23. Metapostnotum defined F (0— 26. 10. 38. NODES 80^64: 12. on ventroposterior margin of gonocoxite tion of tergal graduli >1). 53. 50. Number of tergal graduh F (0^2). 37. surface of hind femur NODES 74—>73: 3. NODES 94-^93: 17. One spine on outer apex of hind tibia M (0^1). 38. Pregradular areas of sterna M lobe on apex of gonostylus M (0-^1). Marginal spines of pygidial plate F (1—>2). Ventroapical plate (1—>2). Appressed setae near metasomal spiracles M (0=>1). 27. Two spines on outer apex of hind tibia M (1^2). Anterior face of Tl (0^1). Margins of basitibial plate F (0=>2). One spine on outer apex of hind tibia ). {0=>I). NODES 72^71 34. M (0=>2). Divergence of ventromedial margin of gonocoxites (1 ^0). M NODES 55->54: NODES 63^62: la tions 32. M (1^0). Propodeal tooth B (0=>1). Distribu- (0=*1). Inner basal margin of venter of gonocoxite M M (0—>1). (0=>1). 36. Ventroapical plate of gonocoxite M (0^1). NODES 49. Setae on apex of (1=>0). M M (2->0). Supraocellar pit F (1=>0). M (0— M M . M (0=>1). 22. 12. Ventroapical plate of gonocoxite valve modified NODES 59^58: 13. 25. NODES 92^91 24. 18. 36. Epistomal suture F (0=>1). One spine on outer apex of hind tibia F (1-^0). NODES 91->90: 49. Mesosomal gland reservoir M (0— 36. Metapostnotum not defined (0—>1 ). M NODES 67^66: 6. 43. 5. Apical margin of SI B (2^0). 49. 16. 40. 54. 40. 2. 52. projecting lobe Subapical tooth on mandible (0—>2). One spine on outer apex of gonostylus hind tibia F (0=>1). 32. Pregradular areas of sterna M (0=>1). Anterior face of Tl F (0=>2). Metapostnotum not defined F (0=>1). Subapical tooth on upper carina of mandible F (0->l). Basitibial plate 1 ^0). Propodeal tooth B (1=>0). Vestiture M (0=>1). 51. Beginning of thoracic declivity F (0=>1). Metapostnotum defined F (1=>0). Basitibial plate NODES 77^65: 19. Apical margin of SI B (2=>0). 23. (0—>1). Penis valves reservoir M (0=>1). 4. 24. Facial Apex of mandible tridentate F (0=>1). 1 7. NODES 85^84: 11. Beginning of metasomal spiracles M (0=>1). Distribution of tergal graduli M =>2). NODES 84^82: 48. 47. Inner basal margin of M M M NODES NODES 58—>57: 57—>56: 51. Graduli on Tl-3 F (0=>2). Spine on ventroposterior margin of gonocoxite M M venter of gonocoxite M {] —>0). NODES 85->63: 26. Setae on apex of gonostylus M (1=>0). 55. of gonocoxite pressed setae near metasomal spiracles M (0=>1). Margins of basitibial plate F (0=>1 ). Beginning Medial carina of S6 (1=>0). Margins of basidbial plate F (0=>1). Ap- apex bifid F {0—>1). (5^1). Facial maculations (0=>1 ). 42. NODES 78^77: 22. Anterior face of Tl F {2=>1 ). Spines on posterior margin T7 M (0=>1). Mesosomal M (0-^1). Apical margin of SI (0=>2). 29. 48. 51 Spine (1—>0). Length of gonostylus M =>0). 40. Length of gonostylus NODES 74->68: 14. NODES 76^75: 7. Medial carina of metasomal sterna F (0—>1). NODES 73^72: 31. ERNST MAYR LIBRARY iii'iiii'iiiiii'ini 2044 . 21. margin of gonocoxites diverge NODES 60^59: 3. 28. M of thoracic declivity NODES (0=>1). 8. 84->83: 24. 39. 30. Extent of basitibial plate F (0—>1). 25. NODES 79^78: 47. One outer tibial spine F (1 ->0). 53. The University of Kansas APPENDIX 3 Character changes and node support for tree shown in Figure 13 A (Analysis 3. 25. Ventroapical plate of gonocoxite NODES 93^92: setae near 11. Mesosomal gland M Medially projecting lobe on apex of gonostylus (1-^0). 68^67: 26. Beginning of thoracic declivity M (0^2). NODES 82—>80: 50. 34. Penis M (0=>I). NODES Abbreviations and conventions as in Appendix 1 95^94: IT One spine on outer apex of hind tibia M (0->I). Medial carina of S6 {0=>1). (0=>1). Pregradular areas of sterna with setal patches M (0^1 Appressed Apex of penis valves M (0=>1). 28. M M : : M M (0=>1). Beginning of thoracic declivity F (1=>0). Up- per rutellar and preapical tooth fused F (3—>8). projecting lobe on apex of gonostylus NODES 63^60: 13. Medially projecting lobe on apex of gonostylus M NODES 91^87: (0=>1). thoracic declivity (1 >1). 52. 41. Island 5. Apical margin of SI B (0=>1). Mesosomal gland reservoir (0^2). Tubercle on ventral surface of hind femur Epistomal suture F (0=>1). Mesosomal gland reservoir (0=>1). 26. NODES 62-^61 apex bifid : 17. 45. Penis valves M (0=>1). 16. NODES 87^86: 3. Medially (0=>1). M M (0=>I). Distribution of tergal graduh (0^2). M Beginning of thoracic declivity F (2—>3). Graduh on TI-3 F (0=>2). Epistomal suture F (1=>0). 31. 26. 31 Distribution of tergal graduli {0=>1). 26. 37. Number of tergal graduli F (0^1). NODES 73^70: 4. Medially projecting NODES 51->49: 1. 53. NODES 52^51: 12. (1—>2). macu- absent M (0=>1). short NODES 94^85: 36. NODES 56—>53: 26. M M 6. Spine on ventroposterior margin of gonocoxite M (0=>1). 39.46 Scientific Papers. Appressed setae near metasomal spiracles (0=>1). Two outer tibial spines F (0^1). Paired marginal spines of pygidial plate F (0->I). gonostylus M (0=>1). 41. 31. 40. Gonostylus slender. Graduli on Tl-3 F (0=>2). Lateral edge of penis 24. 11. 52. 21. Metapostnotum not defined : M (0=>1). Beginning of thoracic declivity M (1^0). Metapostnotum not defined F (0=>1). Medial carina of metasomal sterna F (0—>1). Supraocellar pit F (0=*1 ). 50. Vestiture (0=>1). Unmodified ventral (1 53^52: 6. Begin41. NODES 75^74: 22. Shape of tegula (0=>1). ( NODES 80-^79: 19. Epistomal suture F (0=>1). reservoir (0=>1). Metapostnotum defined M {0->l). 26. 7. apex entire F >2). Shape of gradulus on Tl F (0=>1 ).

ventroposterior margin of gonocoxite NODES 85^84: 22. M NODES plate F (0^1). 27. 34. 17. Inner basal margin of venter of gonocoxite M M M M NODES 56^53: Mesosomal gland reservoir M (2->3). 4. Length of (1—>2). 25. 37. Subapical tooth on upper carina of mandible F {0=>1). Spines on posterior margin T7 NODES 62^61: 17. 51. Metapostnotum not defined (l-»0). 41 Paired marginal spines of pygidial . Propodeal M (1=>0). declivity F (0=>1). Supraocellar pit F ( 1 =>0). 55. Epistomal suture F Beginning of thoracic 33. Mediallv projecting lobe on apex of gonostylus 26. NODES 78->77: 6. M (0=>1). NODES 90->89: 24. Subapical tooth on upper carina of mandible F (0->l). 13. 50->49: 5. 21. sterna (0=>1). Graduli tooth and preapical tooth fused F (0->l). NODES 91->90: 36. M (0=>1). 21 Extent of basitibial plate F (0^1). Ventroapical plate ). 51^50: 32. gonostylus (1=>0). (0—>1). NODES NODES 39. Beginning Setae on apex of NODES NODES 87^75: 8. Mesosomal gland (0—>1). Apex of mandible bidentate F (l->0). 32. NODES 65^64: 6. 25. Gonostylus slender. Appressed setae near metasomal spiracles (1—>2). Metapostnotum defined F (1=>0). NODES 71->70: setae near 39. Medial carina of margin of venter of NODES 55^54: 32. Appressed setae near metasomal spiracles (0=>1). NODES 80^79: 26. 50. . 55. Setae maculations absent (0=»1). Upper rutellar 12. Beginning of thoracic declivity (0=>1). 40. Mesosomal gland reservoir M (1^2). Spine on ventroposterior margin of gonocoxite 57-»56: 49. Graduli on Tl-3 F (0=>2). 38. Graduli on Tl-3 F (0=>2). 49. 48. Basitibial plate apex F reservoir Mesosomal gland (0=>1). 52. 41. 40. 40. Anterior face of Tl F (0=>1). (l->0). 26. 36. Epistomal suture F (1=>0). M M NODES 78-»76: 16. NODES 92-»91. Apical margin of SI B (0=>1). of hind tibia Two spines on outer apex M (0->l). Margins of basitibial plate F (0=>2). Xylocopini 47 APPENDIX 4 Character changes and node suitort for tree shown Abbreviations and conventions as in Appendix 95^*^4: 1. Medial groove of Tl F (0=»1). NODES NODES 58->57: 51. M 4. 36. Long malar space F (0=>1). M One spine on M Two spines on outer apex of hind tibia (1^2). Facial macu- gonocoxite M Cl->0). Apical margin of SI M (2=s0). Apical margin of SI B (2^0). 87^86: 6. Spine on ventral surface of hind femur M (0=>2). Ventroapical plate of gonocoxite M One spine on outer apex of hind tibia (0—>1). Metapostnotum not defined M (0=>1). 23. NODES 94^93. in Figure 15 (Analysis 3. Ventromedial margin of gonocoxites diverge M (0=>1). F (1^0). Beginning of thoracic declivity F (0=>I): 26. metasomal sterna F (0=>1). Vestiture (0=>1). Number of tergal graduli F (0^1). 18. 31. 7. Mesosomal gland reservoir M M M M \'al\-e modified M (0=>1). Appressed setae near metasomal spiracles (0=>1). Medial carina of S6 M (1^0). 54. on Tl-3 F (0->2). Setae on apex of gonostylus M (0=>1). Epistomal suture F (0=>I). M M (0^1). M (1^0). Ventromedial mar{0=>1). 26. NODES 89^87: 19. Basitibial plate (0=>1). Supraocellar pit F (0=>1). 49. 11. 28. 52. Medial carina of S6 M (0^1 ). 50. One spine on outer apex of hind tibia NODES Basitibial plate apex bifid F (0^1). Margins of F (0=>1). 32. 43. Propodeal tooth B (0=>1 ). M M NODES 67->66: NODES 83^82: 31. Medially projecting lobe on apex of gonostylus M (0—>1). Shape of gradulus on Tl F {0-»l). Epistomal suture F gonostylus M (1=>0). NODES 90->74: 22. Mesosomal gland reservoir M (0->2). lations absent metasomal sterna F 1 0. Distribution of tergal graduli {1=>0). Beginning of thoracic declivity M (0=>2). Ventroapical plate of gonocoxite M (1 ->2). Subapical tooth on mandible (0=>1). 24. Margins of basitibial plate F (0=>1). Appressed metasomal M (0^1). Island 6. NODES 66^65: 16. 52. Inner basal Shape of M tegula M (0=>I). 25. 52. 51. 29. character weights equal). In(0=>1). M(0=>1). 28. NODES 59^58: 13. 52. 17. NODES 70^69: Metapostnotum not defined (0=>1). NODES 63->62: 4. Distribution of tergal graduh M (0=>2). (0=>1). 41. 32. M Metapostnotum not defined F (0=>1). Penis valves M (0=>1). 45. 44. 19. Apical margin of SI B (2^0). (0=>1). 50. Medially (0—>2). Apex of mandible tridentate F (0=>1). (1^2). Beginning of thoracic declivity F (2^3). Subapical tooth on mandible (0^1). 42.Subgenera and Genera of Carpenter Bees. Two spines on outer apex of hind tibia F (0^1). Extent of basitibial plate F (1=>0). 53. 49. M Medially projecting lobe on apex of gonostylus Epistomal suture F (1 =>0). 11. NODES 89^88: 48. 12. reservoir (0->l). (0=>1). NODES 52-^51 Apex of mandible tridentate F (0->l). NODES 69-^68: NODES 68->67: (0^1). Pregradular areas of of thoracic declivity M (0=*1). Penis M(O^l). Marginal spines of pygidial plate F (1=>0). Medially projecting (0—>1). Penis valves ner basal margin of venter of gonocoxite 91^73: 10. 25. 40. 49. 23. apex bifid NODES 72^71. Spine on 31. outer apex of hind tibia F CO—>1). 26. lobe on apex of gonostylus NODES 83^80: 1. Ventroapical plate of gonocoxite (0=>1). 39. projecting lobe on apex of gonostylus Length of gonostylus M (0=>1). (1^5). 47. 37. Tubercle on ventral surface of hind femur M (0=>1). Distribution of tergal graduh M (0=>2). NODES 82->81: 34. Anterior face of Tl F (0=>2). Pregradular areas of sterna NODES 56^55: 2. NODES 60^59: 3. NODES 63^60. 11. 26. 36. Anterior face of Tl F (2=>1). ventroposterior margin of gonocoxite 6. (0=>1). Facial M M M (0=>1). 47. NODES 84->83: 23. Ventroapical plate of gin of gonocoxites diverge gonocoxite (0->l). Beginning of thoracic declivity F (1^2). 53. Medially projecting lobe on apex of gonostylus M M M M M M (0^1). Ventro- M . Spine on (0=>1). M (0=>1). 5. M NODES 53^52: : 6. 26. Medial carina of Vestiture M (0=>1). short M (0->l). Metapostnotum defined F (1=>0). 36. Apical margin of SI (11^2). Upper rutellar tooth and preapical tooth separate F 7. M (0=>1 40. Apex of penis valves (0=>1). 53. Graduh on Tl-3 F (0=>2). M (0=>1). M NODES 84^78: 14. 2. Two spines on outer apex of hind tibia M (1=*0). 47. basitibial plate on apex of gonostylus M (0=>1). Appressed setae near metasomal spiracles 38. >2). 49. M tooth Mesosomal gland reservoir M (5->l). Metapostnotum not defined F Mesosomal gland reservoir 3. NODES 86-^85: 30. 1. medial margin of gonocoxites diverge (1-^2). 23. Pregradular areas of sterna spiracles M (0=>I). Triangular hind trochanter F (0->l). of gonocoxite M (0— 24. Marginal spines of pygidial plate F Gonostylus (0=>1). 35. Lateral edge of penis NODES 92->72: NODES 72-^63: 48.

Natural History Museum. and Special Publications of the Natural History Museum can be purchased from the Office of Publications. THE UNIVERSITY OF KANSAS The University of Kansas Publications. Series Editor: William E. fax (913) 8645335. Duellman Printed by The University of Kansas Printing Services Lawrence. USA. The University of Kansas Libraries. Available back issues of The University of Kansas Science Bulletin may be purchased from the Library Sales Section. Publications are composed using Microsoft Word® and Adobe PageMaker® on a Macintosh computer and are printed by The University of Kansas Printing Services. The publication is printed on acid-free paper. final acceptance is 1 in 1997. The University of Kansas Libraries. The piration date. Scientific Papers. beginning with Volume 1 in 1902. The University of Kansas Science Bulletin. Kansas . Available issues of former publication series. The University of Kansas. Special Publications at the discretion of the editor. or e-mail (kunhm@ukans. Lawrence. by addressing the Exchange Librarian. be- gun with Number began in 1976 and continue as an outlet for longer contributions and are available by purchase only. Purchasing information can be obtained by calling (913) 864-4450. include exInstitutional libraries interested in exchanging publications entific Papers. USA. Museum of Natural History. Retrieval Services Department. may obtain the SciUniversity of Kansas. Shorter research papers formerly published in the above series were published as The University of Kansas Natural History Museum Occasional Papers until Number 180 in December 1996. Kansas 66045-2454. Monographs of The University of Kansas Natural History Museum were initiated in 1970 and discontinued with Number 8 in 1992. The University of Kansas. VISA and MasterCard accepted. The Miscellaneous Publications of The University of Kansas Natural History Museum began with Number 1 in 1946 and ended with Number 68 in February 1996. Natural History Museum.edu).PUBLICATIONS OF THE NATURAL HISTORY MUSEUM. Kansas 66045-2800. USA. Natural History Museum. Lawrence. All manuscripts are subject to critical review by intra. Kansas 660452800. The foregoing publication series are now combined in a new series entitled Scientific Papers.and extramural specialists. Lawrence. was discontinued with Volume 20 in 1971. beginning with Volume 1 in 1946. was discontinued with Vol- ume 55 in 1996.

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