Evolution of The Apoidea

A MONOGRAPH OF
THE BALTIC AMBER BEES

AND EVOLUTION OF THE
APOIDEA (HYMENOPTERA)
MICHAEL S. ENGEL
Research Scientist, Division of Invertebrate Zoology,
Presently: Research Associate, Division of Invertebrate Zoology,
American Museum of Natural History;
Fellow, the Linnean Society of London;
Assistant Curator, Division of Entomology,
Natural History Museum and Biodiversity Research Center;
Assistant Professor, Department of Ecology and Evolutionary Biology,
1460 Jayhawk Boulevard, Snow Hall, University of Kansas,
Lawrence, Kansas 66045-7523
BULLETIN OF THE AMERICAN MUSEUM OF NATURAL HISTORY

Number 259, 192 pp., 123 figures, 7 color plates, 15 tables
Issued April 16, 2001
Price: $17.00 a copy
Copyright q American Museum of Natural History 2001
ISSN 0003-0090
Photomicrograph of the holotype (IMGP-K72) of Glyptapis mirabilis Cockerell.
CONTENTS
Abstract . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 5
Zusammenfassung . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 6
Introduction . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 7
Historical Review . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 13
Age and Origin of Amber . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 15
Materials and Methods . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 17
Specimen Preparation and Study . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 18
Format . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 18
General Morphology . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 19
Prosoma . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 20
Mesosoma . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 23
Metasoma . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 32
Systematic Paleontology . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 33
Superfamily Apoidea Latreille . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 34
Division Apiformes . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 35
Short-Tongued Versus Long-Tongued Bees . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 35
Short-Tongued Bees . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 36
Family Halictidae Thomson . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 36
Subfamily Halictinae Thomson . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 36
Tribe Halictini Thomson . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 37
Genus Electrolictus, new genus . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 38
Family Paleomelittidae, new family . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 41
Genus Paleomelitta, new genus . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 42
Family Melittidae Schenck . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 44
Subfamily Macropidinae Robertson . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 46
Tribe Eomacropidini, new tribe . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 46
Genus Eomacropis, new genus . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 46
Long-Tongued Bees . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 49
Family Megachilidae Latreille . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 49
Subfamily Lithurginae Newman . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 50
Tribe Protolithurgini, new tribe . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 51
Genus Protolithurgus, new genus . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 51
Subfamily Megachilinae Latreille . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 53
Tribe Osmiini Newman . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 54
Genus Glyptapis Cockerell . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 56
Genus Ctenoplectrella Cockerell . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 68
Genus Glaesosmia, new genus . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 75
Family Apidae Latreille . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 76
Subfamily Xylocopinae Latreille . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 77
Tribe Boreallodapini, new tribe . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 77
Genus Boreallodape, new genus . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 78
Subfamily Apinae Latreille . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 86
Tribe Electrobombini, new tribe . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 88
Genus Electrobombus, new genus . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 88
Tribe Electrapini Engel . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 90
Genus Electrapis Cockerell . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 92
Genus Protobombus Cockerell . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 99
Genus Thaumastobombus, new genus . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 109
Tribe Melikertini, new tribe . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 112
Genus Succinapis, new genus . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 115
Genus Melikertes Engel . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 123
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Genus Melissites, new genus . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 129

Genus Roussyana Manning . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 131
Tribe Meliponini Lepeletier de Saint Fargeau . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 133
Genus Kelneriapis Sakagami . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 134
Genus Liotrigonopsis, new genus . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 135
Apoidea Incertae Sedis . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 137
Genus and Species Indeterminate . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 137
Missing Taxa of Uncertain Identity . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 138
Cladistic Analyses . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 139
Lithurginae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 139
Xylocopinae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 140
Corbiculate Apinae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 142
Discussion . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 150
Origin of Bees . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 150
The Eocene Bee Fauna . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 159
Epilogue . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 161
Acknowledgments . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 161
References . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 162
Appendix 1: Bees in Amber and Copal . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 175
Appendix 2: Tribal Classification and Geological Distribution of Bees . . . . . . . . . . . . . . 177
Appendix 3: Catalog of Family- and Genus-Group Names for Fossil Bees . . . . . . . . . . . 180
Appendix 4: Full Titles of Journals Abbreviated in References . . . . . . . . . . . . . . . . . . . . . 182
Index of Taxonomic Names . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 185
Color Plates . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 186192
2001
ENGEL: BALTIC AMBER BEES
ABSTRACT
The bees found as Baltic amber inclusions are revised and the history of studies on these
fossils is briefly reviewed. In total this subtropical Eocene fauna contains 36 species and 18
genera, all extinct. These are classified here into nine tribes, six subfamilies, and five families
of which six tribes and one family are unknown in the modern fauna. The following taxa are
described as new to science: PALEOMELITTIDAE, new family; Boreallodapini, Electrobombini, Eomacropidini, Melikertini, Protolithurgini, new tribes; Ctenoplectrellina, new
subtribe; Boreallodape, Electrobombus, Electrolictus, Eomacropis, Glaesosmia, Liotrigonopsis, Melissites, Paleomelitta, Protolithurgus, Succinapis, Thaumastobombus, new genera;
Boreallodape baltica, B. mollyae, B. striebichi, Ctenoplectrella cockerelli, C. grimaldii, Electrapis krishnorum, Electrobombus samlandensis, Electrolictus antiquus, Eomacropis glaesaria, Glaesosmia genalis, Glyptapis densopunctata, G. disareolata, Liotrigonopsis rozeni,
Melikertes clypeatus, Melissites trigona, Paleomelitta nigripennis, Protobombus basilaris,
Protolithurgus ditomeus, Succinapis goeleti, S. micheneri, S. proboscidea, Thaumastobombus andreniformis, new species (seven new family-, 11 new genus-, and 22 new speciesgroup taxa). The genus Electrapis is found to be paraphyletic and the subgenera Melikertes,
Roussyana, and Protobombus are given generic status outside of Electrapis. The subtribe
Electrapina is elevated to tribal rank among the corbiculate Apinae and the subfamily Glyptapinae of Cockerell is reduced to subtribal rank within Osmiini. The genera Chalcobombus
and Sophrobombus are newly synonymized with Protobombus. Glyptapis reducta Cockerell
is synonymized with G. fuscula Cockerell, Electrapis minuta Kelner-Pillault with Apis palmnickenensis Roussy, Ctenoplectrella splendens Kelner-Pillault and C. dentata Salt both with
C. viridiceps Cockerell, Electrapis apoides Manning and Chalcobombus humilis Cockerell
both with Protobombus indecisus Cockerell, and Electrapis bombusoides Kelner-Pillault with
E. tornquisti Cockerell (new synonymies). The following new combinations are proposed:
Electrapis martialis (Cockerell), Melikertes proavus (Menge), M. stilbonotus (Engel), Kelneriapis eocenica (Kelner-Pillault), Protobombus fatalis (Cockerell), P. hirsutus (Cockerell), and
Electrapis martialis (Cockerell) (new combinations). A lectotype is designated for Electrapis
minuta Kelner-Pillault and neotypes designated for Apis meliponoides Buttel-Reepen, A. palmnickenensis Roussy, Chalcobombus humilis Cockerell, C. hirsutus Cockerell, C. martialis
Cockerell, Ctenoplectrella dentata Salt, C. viridiceps Cockerell, Electrapis tornquisti Cockerell, Glyptapis reticulata Cockerell, G. neglecta Salt, Protobombus indecisus Cockerell, P.
tristellus Cockerell, and Sophrobombus fatalis Cockerell. The subfamily Xylocopinae is recorded for the first time from amber, and the families Halictidae and Melittidae are confirmed
as occurring in Baltic amber. The oldest fossils of the Halictidae, Megachilidae (Lithurginae
and Megachilinae), Melittidae, and Xylocopinae are reported and described herein. Keys are
presented for the identification of the Baltic amber bees. Three new recent taxa are also
proposed: Penapini, Redivivini, and Meliturgulini (new tribes).
Cladistic analyses of the Lithurginae, Xylocopinae, and corbiculate Apinae are presented.
Preliminary investigation of lithurgine relationships demonstrates that the fossil, Protolithurgus, is sister to extant genera of the subfamily and that Lithurgus s.s. is likely paraphyletic
with respect to Lithurgopsis and Microthurge. Xylocopine relationships are generally in accord
with those previously proposed: Xylocopini sister to all other tribes and Manueliini sister to
the abruptly narrowed mandible tribes (i.e., Ceratinini, Allodapini, and Boreallodapini). Boreallodapini, despite some apparently apomorphic similarities with Ceratinini, is supported as
sister to Allodapini. The paleontological evidence for corbiculate bee phylogeny reinforces
traditional concepts over relationships of these tribes and contradicts recent molecular studies.
Two extinct corbiculate tribes are based on specimens that exhibit worker morphologies indicative of advanced eusocial behavior and are related to the living, advanced eusocial tribes
Apini and Meliponini, supporting a hypothesis of a single origin for this behavioral characteristic.
The composition of the Eocene bee fauna of Europe is discussed. The fauna is predominantly composed of long-tongued bees, but some short-tongued bees are represented in the
families Halictidae, Paleomelittidae, and Melittidae. Bees in Baltic amber are generally allied
with groups currently inhabiting sub-Saharan Africa and southeast Asia, a biogeographic pattern common to many Baltic amber groups.
The phylogeny and origin of bees are discussed. The bees are a derived, monophyletic
NO. 259
group of the spheciform wasps and presumably arose sometime in the earliest mid-Cretaceous
after the origin of angiosperms. Reports of bees from Jurassic strata or earlier are all refuted.
All available evidence supports the idea that bees originated shortly after flowering plants and
diversified into higher lineages contemporaneously with the radiations of angiosperms. Major
bee lineages (i.e., families) were thus presumably established by the Late Cretaceous.
A catalog of bees presently known in amber and copal is appended as is a catalog of familyand genus-group names for fossil bees (both amber inclusions and compression fossils). A
preliminary outline of the tribal classification of Recent and fossil bees with their geological
distribution is presented (with three new family-group taxa for living bees).
Z U S A M M E N FA S S U N G
Die als Inklusen des Baltischen Bernsteins nachgewiesenen Bienen werden revidiert und
ihre Erforschungsgeschichte kurz dargestellt. Insgesamt umfasst diese subtropische Fauna des
Eozans 36 Arten und 18 Gattungen, die alle ausgestorben sind. Diese werden in neun Tribus,
sechs Unterfamilien, und funf Familien klassifiziert, von denen sechs Tribus und eine Familie
in der heutigen Fauna unbekannt sind. Die folgenden Taxa sind neu fur die Wissenschaft:
PALEOMELITTIDAE, fam. nov.; Boreallodapini, Electrobombini, Eomacropidini, Melikertini, Protolithurgini, tribus nov.; Ctenoplectrellina, subtribus nov.; Boreallodape, Electrobombus, Electrolictus, Eomacropis, Glaesosmia, Liotrigonopsis, Melissites, Paleomelitta,
Protolithurgus, Succinapis, Thaumastobombus, gen. nov.; Boreallodape baltica, B. mollyae,
B. striebichi, Ctenoplectrella cockerelli, C. grimaldii, Electrapis krishnorum, Electrobombus samlandensis, Electrolictus antiquus, Eomacropis glaesaria, Glaesosmia genalis, Glyptapis densopunctata, G. disareolata, Liotrigonopsis rozeni, Melikertes clypeatus, Melissites
trigona, Paleomelitta nigripennis, Protobombus basilaris, Protolithurgus ditomeus, Succinapis goeleti, S. micheneri, S. proboscidea, Thaumastobombus andreniformis, spec. nov.
(sieben neue Familien-, 11 neue Gattungs-, und 22 neue Artengruppentaxa). Die Gattung
Electrapis ist paraphyletisch, und die Untergattungen Melikertes, Roussyana, und Protobombus werden unabhangig von Electrapis als Gattungen akzeptiert. Die Untertribus Electrapina
erhalt den Rang einer Tribus innerhalb der corbiculaten Apinae, und die Unterfamilie Glyptapinae von Cockerell wird als Untertribus innerhalb der Osmiini interpretiert. Die Gattungen
Chalcobombus und Sophrobombus werden neu mit Protobombus synonymisiert. Folgende
Arten werden miteinander synonymisiert: Glyptapis reducta Cockerell mit G. fuscula Cockerell, Electrapis minuta Kelner-Pillault mit Apis palmnickenensis Roussy, Ctenoplectrella
splendens Kelner-Pillault und C. dentata Salt beide mit C. viridiceps Cockerell, Electrapis
apoides Manning und Chalcobombus humilis Cockerell beide mit Protobombus indecisus
Cockerell, Electrapis bombusoides Kelner-Pillault mit E. tornquisti Cockerell (neue Synonyme). Die folgenden Kombinationen werden neu eingefuhrt: Electrapis martialis (Cockerell),
Melikertes proavus (Menge), M. stilbonotus (Engel), Kelneriapis eocenica (Kelner-Pillault),
Protobombus fatalis (Cockerell), P. hirsutus (Cockerell), und Electrapis martialis (Cockerell)
(neue Kombinationen). Ein Lectotypus wird fur Electrapis minuta Kelner-Pillault festgelegt.
Fur folgende Arten werden Neotypen designiert: Apis meliponoides Buttel-Reepen, A. palmnickenensis Roussy, Chalcobombus humilis Cockerell, C. hirsutus Cockerell, C. martialis
Cockerell, Ctenoplectrella dentata Salt, C. viridiceps Cockerell, Electrapis tornquisti Cockerell, Glyptapis reticulata Cockerell, G. neglecta Salt, Protobombus indecisus Cockerell, P.
tristellus Cockerell, und Sophrobombus fatalis Cockerell. Die Unterfamilie Xylocopinae wird
erstmals in Bernstein nachgewiesen, und das Vorkommen der Familien Halictidae und Melittidae in Baltischem Bernstein wird bestatigt. Die altesten Fossilien der Halictidae, Megachilidae (Lithurginae und Megachilinae), Melittidae, und Xylocopinae werden nachgewiesen
und beschrieben. Bestimmungsschlussel fur die Bienen des Baltischen Bernsteins werden
erstellt. Drei rezenten Taxa sind neu: Penapini, Redivivini, und Meliturgulini (tribus nov.).
Kladistische Analysen der Lithurginae, Xylocopinae, und der corbiculaten Apinae werden
vorgestellt. Vorlaufige Untersuchungen der verwandtschaftlichen Beziehungen der Lithurginae zeigen, dass die fossile Protolithurgus Schwestergruppe der rezenten Gattungen der Lithurginae ist. Auerdem ist Lithurgus s.s. wahrscheinlich paraphyletisch in Bezug auf Lithurgopsis und Microthurge. Die Verwandtschaftsbeziehungen der Xylocopinae stimmen
grundsatzlich mit den bisher angenommenen uberein: Die Xylocopini sind die Schwester-
2001
gruppe aller ubrigen Tribus, wahrend die Manueliini die Schwestergruppe aller Tribus mit
Arten mit abrupt verschmalerten Mandibeln ist (z.B. Ceratinini, Allodapini, und Boreallodapini). Boreallodapini wird als Schwestergruppe der Allodapini begrundet, abgesehen von
hnlichkeiten mit den Ceratinini. Die palaontologischen
einigen anscheinend apomorphen A
Indizien stutzen traditionelle Konzepte der phylogenetischen Beziehungen der corbiculaten
Bienen und widersprechen neueren molekularen Untersuchungen. Die Arbeiter zweier fossiler, corbiculater Tribus zeigen morphologische Merkmale, die auf stark abgeleitetes eusoziales Verhalten hindeuten: Diese Tribus sind offenbar mit den rezenten, hoch-eusozialen
Tribus Apini und Meliponini verwandt, was die Hypothese der einmaligen Entstehung dieses
Verhaltensmerkmals unterstutzt.
Die Zusammensetzung der eozanen Bienenfauna wird diskutiert. Sie besteht vorwiegend
aus langzungigen Bienenarten, auch wenn einige kurzzungige Arten der Familien Halictidae,
Paleomelittidae, und Melittidae ebenfalls vertreten sind. Im Allgemeinen sind die Bienen des
Baltischen Bernsteins mit rezenten Bienen des Afrikas sudlich der Sahara und Sudostasiens
verwandt; ein bei Organismen des Baltischen Bernsteins haufiges biogeographisches Muster.
Die Phylogenie und der Ursprung der Bienen werden diskutiert. Die Bienen sind eine
abgeleitete, monophyletische Gruppe der spheciformen Wespen und entstanden irgendwann
in der fruhesten Mittelkreide nach der Entstehung der Angiospermen. Nachweise von Bienen
aus dem Jura oder noch fruher werden widerlegt. Alle verfugbaren Indizien stutzen die Annahme, dass die Bienen kurz nach den Blutenpflanzen entstanden und in ihre grundsatzliche
Diversifikation parallel zur Angiospermenradiation erfolgte. Die groen Linien der Bienen
(z.B. Familien) waren daher in der spaten Kreide bereits vorhanden.
Ein Katalog der in Bernstein und Kopal nachgewiesenen Bienen und ein Katalog der
Familien- und Gattungsgruppennamen fossiler Bienen (sowohl aus Bernstein wie aus Sedimenten) wird vorgestellt. Eine vorlaufige Idee einer Klassifikation der rezenten wie fossilen
Bienen auf Tribus-Niveau mit ihrer geologischen Geschichte wird entwickelt (mit drei neuen
Familiengruppennamen fur rezente Bienen).
Et latet, et lucet Phaetontide condita gutta,

Ut videatur apis nectare clausa suo;
Dignum tantorum pretium tulit illa laborum,
Credible est ipsam sic voluisse mori.1
Bees occupy a unique role in the world of

arthropods. They are entwined into most aspects of human culture and mythology, not
to mention our agriculture, economy, and
general ecology. Mention of their name instantly recalls a suite of images and senses
tying together warm days, sunny skies, soft
breezes, and fields of fragrant and colorful
flowers teeming with the gentle murmur of

benevolent pollen harvesters at their labors.
Bees have instilled within us ideal notions of
industriousness, loyalty, obedience, and sacrifice not through denigration but through almost holy reverence to a greater good. Even
the famous parable from the Bible (Proverbs
6) teaching us to go to the ant originally
included reference to the labors of bees2.
Such emotions and concepts connected to
bees flow through our common natural heritage; even the great Mother Goddess uncovered at the root of most Western religions
was symbolized by a bee.
More significant, perhaps, than their pervasiveness in human society is that bees
serve as the most significant pollinators of
flowering plants, helping to sustain and underpin major ecological systems worldwide
1 Translation: Now dim, now bright, trapped in its

amber tear, / A bee seems sealed in its own nectar clear;
/ For a life of endless toil, most fitting pay /Surely a
bee would wish to die this way! // Martials Phaethontis
or amber tear is a reference to the myth of Phaeton,
who died when he tried to drive the chariot of the sun
across the sky. Phaetons sisters wept and their tears
turned to amber. Translation courtesy V. Krishna.
2 Go to the bee, and learn how diligent she is, and

what a noble work she produces, whose labors kings and
private men use for their use, she is desired and honored
by all, and though weak in strength she values wisdom
and prevails. Proverbs, Chapter 6, Septuagint version
(i.e., the oldest Greek version and perhaps the oldest
extant version) of the Holy Bible (see also Engel, 2000a:
2).
M. V. Martialis, Epigrammata,
ca. 89 anno domini
MONOGRAPHIA APUM SUCCINI BALTICI

INTRODUCTION
(e.g., Moldenke, 1976; Arroyo et al., 1982;

Bawa et al., 1985; Bawa, 1990; Petanidou
and Vokou, 1990; Buchmann and Nabhan,
1996; Proctor et al., 1996). The bees are also
arguably one of the more diverse groups of
pollinators, with a standing diversity of ca.
20,000 species. A phylogenetic study of the
fossil record is crucial to understanding how
this modern diversity came to be, as well as
for identifying the periods of diversification
and extinction, reconstructing historical zoogeography, and uncovering the sequence and
origin of synapomorphies (e.g., Patterson,
1981; Grande, 1985; Eldredge and Novacek,
1985; Donoghue et al., 1989; Novacek and
Wheeler, 1992; Grimaldi, 1992). The study
of fossil bees, or paleomelittology3, is still in
its infancy and a complete synthesis of fossil
and living taxa has yet to be achieved. The
present work is one of a series of attempts to
unite paleomelittology with the systematics
of living bees into a meaningful whole. Fossils of bees are rare compared to some other
insect groups; the few known specimens are
typically discovered in isolation, and scattered in sites around the world. Three deposits, however, have produced sizeable bee paleofaunas.
F LORISSANT , C OLORADO : Fossils from
Florissant, Colorado, are compressions with
virtually no relief, and were formed in fine
volcanic ash over several millions of years
of repeated eruptions. Preservation at Florissant can be quite fine, but those bees discovered to date typically preserve only the
wing venation or some structures of the legs,
so meaningful comparisons with other deposits or living taxa are not possible. Bees
have been described from the families Halictidae, Andrenidae, Melittidae, Megachilidae, and Apidae (Cockerell, 1906, 1908a,
1908b, 1908c, 1909d, 1909e, 1911a, 1911b,
1913a, 1913b, 1914, 1917, 1923, 1925), although those of the Melittidae, Andrenidae,
and some Apidae are dubiously assigned to
3 The word paleomelittology, admittedly of my own
creation, is introduced here for the first time. It is derived
from the Greek words palaioz (palaios, meaning ancient), melitta (melitta, meaning bee), and logoz
(logos, suffix denoting knowledge of). It can be defined as: (noun) the science of fossil bees; a branch of
knowledge that deals with bees of past geological periods.
NO. 259
family. These deposits have been dated at approximately 34.1 Ma (Epis and Chapin,
1974), placing them near the Eocene-Oligocene boundary, and are conservatively treated as early Oligocene herein. Work on the
bees of these deposits is ongoing.
DOMINICAN AMBER: Bees in Dominican
amber can be quite common; worker specimens of Proplebeia dominicana (Wille and
Chandler) number into the thousands and can
be frequently purchased in gem and mineral
shops at no great expense. All other bee
groups from these deposits are known on the
basis of but a few individuals, often only
one. Through a series of recent papers this
fauna has been essentially revised. Table 1
summarizes the Miocene bee fauna of Hispaniola as it is presently understood. This paleofauna is quite modern in character. Most
species are representative of extant genera,
although several are today locally extinct
from the West Indies (e.g., Heterosarus, Neocorynura, Chilicola, Meliponini) or from
Hispaniola (e.g., Euglossa, Eufriesea). Those
few Dominican amber genera that are extinct
are remarkably similar to living forms. All
groups are easily placed in living subtribes,
tribes, and other higher categories.
The age of the Dominican amber mines
has been of some controversy. Lambert et al.
(1985) suggested a range of ages from Eocene to Miocene based on NMR analyses.
Grimaldi (1995), however, argued that Dominican amber had to be younger than the
Eocene and Oligocene dates, while IturraldeVinent and MacPhee (1996, 1999) later provided a strong stratigraphic basis for an early
Miocene age. This geological evidence is
also consistent with the inclusions in Dominican amber that are modern not only in general appearance but also in cladistic position
(e.g., bees discussed briefly above; for flies
see Grimaldi, 1995).
BALTIC AMBER: The middle Eocene amber of northern Europe was previously believed to contain merely a handful of specimens of rather rare and enigmatic bee species
(e.g., Cockerell, 1908b, 1909b, 1909c; Zeuner and Manning, 1976). Although the few
bees were recognized to be of phylogenetic
import, the general paucity of material and
the loss of several type specimens deterred
meaningful work on the deposit. Through
2001
TABLE 1
Bee Fauna of Miocene Dominican Amber

(Updated from Engel, 1999d)
10
diligent efforts over the past three years I

have managed to increase the number of
known specimens by nearly sevenfold and,
as will be shown in the following pages of
this monograph, the Baltic amber bee fauna
is in actuality the most diverse extinct bee
fauna known (summarized in table 2). Although I have increased the number of
known individuals and species significantly,
bees are still quite rare in Baltic amber, with
only one specimen appearing among approximately every 5000 inclusions. Naturally, as
work continues the number of specimens will
grow steadily. I will be interested in examining new material as it becomes available
for the production of supplements to this
work. Since this monograph concerns itself
expressly with the Baltic fauna, a discussion
of the age of this amber is presented below
(refer to Age and Origin of Amber).
Various other Cenozoic sites have produced bees (e.g., Late Cretaceous amber
from New Jersey, Oligocene-Miocene amber
of Mexico, Miocene compression from Japan, sundry Eocene through Miocene compression fossils in Europe and North America), and although work is continuing on
these and other deposits (e.g., Engel, 1998c,
2000b, unpubl. data), none have at present
revealed the number of individuals or diversity of bees exhibited by the above three locales. Moreover, most of these are compression fossils. If we seek to explore the implications of fossil taxa on our knowledge of
bee evolution in general, then direct comparisons are required between often tiny or obscure morphological traits; such features are
rarely preserved in even the finest of compression fossils. Amber, however, is unrivaled in the paleontological realm for its
ability to preserve fine details (including internal tissues; e.g., Grimaldi et al., 1994) and
so from at least those fossils of the Dominican Republic and the Baltic region we can
begin to seek the necessary character information for comparison and eventual cladistic
study. A further limitation arises, however;
only one of the two major amber sites (i.e.,
Baltic amber; not surprisingly the older of
the two deposits) has revealed taxa that are
not essentially modern. As briefly mentioned
above, the bees of the lower Miocene found
as inclusions from the Dominican Republic
NO. 259
TABLE 2
Hierarchical Supraspecific Classification of

Baltic Amber Bees
(Number of named species in
each genus-group indicated)
2001
fall nicely into extant genera or into genera

closely allied to living groups. Thus, although providing valuable information on local West Indian paleomelittology, extinctions, and patterns of historical biogeography, Dominican taxa contribute little to our
knowledge of bee phylogeny. In fact, a general comparison (of all deposits) of bees from
the Oligocene to the present day shows very
little difference in the fauna at the generic or
higher levels (refer to Discussion, below).
Early statements pertaining to Baltic amber bees were tantalizing: With the general
form of Apis [honey bees], it has venation
nearer to that of Bombus [bumble bees]. . .
(Cockerell, 1909b: 8). Observations like
these, combined with the much older age of
Baltic amber, make a study of those taxa all
the more enticing and critical. Now, with so
many more Baltic specimens at hand, it is
possible to confirm the observations of earlier authors, to add numerous newly discovered lineages to the list of known taxa, to
carefully and systematically describe such
peculiar groups, to explore the phylogenetic
implications of these taxa, and to make more
general observations and hypotheses on the
origin, diversification, and evolution of the
bees. As the following pages demonstrate,
the Baltic amber bee fauna is the best documented paleofauna for bees, the most diverse (in both numbers of species, genera,
and other higher categories), and the most
critical for phylogenetically important taxa.
The Baltic amber is the oldest deposit in the
world with a bee fauna of any sizeable significance.
This work is, in essence, a faunal revision,
the area under consideration being the Baltic
region of 45 million years ago. As with any
faunal revision there is concern over whether
or not the author took a global view when
delimiting groups and thereby avoided proposing synonyms for taxa well characterized
outside of the limited geographic scope of
the work. I hope to eliminate this potential
criticism at the start by stating that, although
dealing with a bee fauna that has long since
vanished from the Earth, I took an exhaustive
geographic and temporal perspective when
analyzing these fossils. Not only were fossils
studied from all existing deposits containing
bees (of all ages, Cretaceous to Pleistocene!),
11
but the living fauna was considered on a

global scale. Moreover, many of the fossils
have been included in cladistic studies with
living groups so as to better define their affinities to known lineages and the validity of
basing taxa on their peculiar character combinations. This has manifested itself most notably for the corbiculate Apinae. Numerous
fossils of this clade occur in Baltic amber,
and owing to the controversy concerning relationships among Recent corbiculates (e.g.,
Schultz et al., 1999), it has been particularly
critical to document the morphology and cladistic implications of the extinct corbiculate
taxa (Engel, 2000c). As a result of this body
of work, the classification of Baltic amber
bees has been radically altered (table 3).
A recent classification of the bees (Michener, 2000a) recognized seven extant families: Colletidae, Stenotritidae, Halictidae,
Andrenidae, Melittidae, Megachilidae, and
Apidae. The classification presented here
also recognizes seven families in total but
differs in two major groupings from that proposed by Michener (2000a). The family
Stenotritidae is a clade of Colletidae and renders the latter paraphyletic; as mentioned
elsewhere (e.g., Engel, 2000a: 5), I consider
Stenotritus and Ctenocolletes to form a subfamily of Colletidae, although stenotritines
should perhaps be further demoted to tribal
rank within Colletinae (or perhaps more appropriately, the latter group should be broken
into several subfamilies). Furthermore, I
have recognized a new family from the fossil
record (described below) that is intermediate
in character between the basic short-tongued
families and a clade consisting of Melittidae
and the long-tongued bees. Lastly, with regard to the familial classification, I believe
sufficient evidence exists for Melittidae
monophyly so as to retain it as a single family, thereby avoiding the proliferation of
melittoid families suggested by Alexander
and Michener (1995: Melittidae s.s., Dasypodaidae4, and Meganomiidae). Appendix 2
along with figures 122 and 123 summarize
my suprageneric classification of bees as it
4 The original spelling for this subfamily, Dasypodinae, has been emended in order to remove homonymy
with the family-group name for armadillos (Mammalia:
Xenarthra: Dasypodidae) (Alexander et al., 1998; ICZN,
1999a).
12
TABLE 3
Comparison of Present Classification with Earlier Usage
NO. 259
2001
presently stands; more detailed studies and

further refinements will be presented elsewhere.
HISTORICAL REVIEW
Baltic amber has had a long association
with humans, perhaps more so than any other
amber deposit. The first Baltic amber bees
were simply mentioned rather than described
or studied in any detail. Authors of antiquity
regularly referred to amber and insect inclusions and, owing to the already tight bond
between bees and man by the Hellenic and
Roman eras, it is little wonder that even fossil bees would appear in their literature (e.g.,
the poem at the opening of this monograph;
the work of Ausonius).
Burmeister (1831, 1832) seems to have
been the first researcher to comment on the
presence of bees in Baltic amber after the
standardization of binomial nomenclature by
Linnaeus (1758). In his works he refers to a
Trigona similar to South American species. None of his material has survived to the
present day; however, individuals of the melikertine bees have a general habitus strongly
reminiscent of Trigona species and it is possible Burmeister had one of these bees before
him. Hope (1836) also mentioned bees in his
private collection of Baltic amber insects as
well as in the collections of his correspondents. Hope referred in his synopsis of amber
insects to specimens of Andrena, Apis,
and Trigona (the latter two perhaps included specimens of Electrapini and Melikertini, respectively, or were actually living
taxa preserved in copal5).
Menge (1856) was the first to describe
some of the forms he had before him as new
species of extant genera, naming species of
what he believed to be Apis and Bombus. In
addition, Menge referred to individuals of
Anthophora, Dasypoda, and Osmia, although
he did not give them specific names. After a
5 Examination of surviving material from Hopes collection in the University Museum, University of Oxford,
has revealed only specimens of extant Hypotrigona and
one Heriades in African copal (mistakenly identified as
succinite). It is possible that all of Hopes original material was actually copal, although that material he reports on from his contemporaries collections certainly
included true Baltic amber. I am grateful to C. OToole
for a loan of this interesting material.
13
diligent search it appears that only one of his

specimens has survived to the modern day,
this being his Apis proava (treated below as
Melikertes proavus). Owing to the misplacement of this species, even following the taxonomic conventions of his time, Menges remaining species must be considered dubiously assigned as to genus. Motschulsky
(1856), working at the same time, recorded
a specimen of Andrena and proposed the
genus and species Bombusoides mengei; the
latter becoming the first new genus-group
proposed for the fauna and the first recognition of an extinct supraspecific lineage
from the Baltic amber. Brischke (1886) later
identified what he considered to be Andrena,
Chalicodoma, Anthophora, Melipona, and
Bombus in Baltic amber, but none were given
names. Like much of Menges material, the
specimens examined by Burmeister, Brischke, and Motschulsky have been presumably lost. All of these authors, except, of
course, Motschulsky, placed the specimens
before them into Recent genera and species
groups. None of the identifications by these
authors can be considered entirely accurate
and should not be relied upon for determining the presence of a particular living bee
genus in Baltic amber nor for placing a middle Eocene age on any of these same genera.
Buttel-Reepen (1906) was the first researcher of the last century to systematically
study a Baltic amber bee as part of his larger
monograph on the honey bees of the world.
In his study, Buttel-Reepen described a Baltic amber fossil as A. meliponoides and considered it as intermediate between the honey
bees (tribe Apini) and their sisters, the stingless bees (tribe Meliponini); hence his choice
of a specific epithet. Apis meliponoides, despite Buttel-Reepens contention, does not fit
into this position (see below), although there
are indeed Baltic amber fossils that appear
intermediate in some characters between Apini and Meliponini and strengthen their sister-group relationship supported by numerous authors (e.g., Engel, 1998d; Noll, 1998;
Schultz et al., 1999, and references therein).
Cockerell was the next individual to undertake a study of the Baltic amber bees and
in far greater depth than Buttel-Reepen, who
was primarily interested in Apis. In a series
of papers published in 1908 and 1909, Cock-
14
erell described the majority of the Baltic amber bees known until the completion of the
present study. Cockerell was intimately familiar with the world bee fauna and was the
first to recognize the significance of the Baltic amber bees, attempting, in a very rudimentary way, to explore the implications of
these taxa on apid evolution (e.g., Cockerell,
1908b) (fig. 120).
Subsequent to Cockerell only a few workers have attempted to study the Baltic amber
bees, none in great detail. Only two small
papers appeared between the work of Cockerell and a small outburst of activity in the
1970s. Salt (1931) proposed new species of
Glyptapis and Ctenoplectrella as well as a
species he believed to be an Andrena (see
Apoidea Incertae Sedis, below), while Manning (1960) described a new species of what
he believed to be Electrapis along with a
new subgenus. Outside of these two works,
systematic study of bee inclusions from the
Baltic stagnated for about one-half of a century. It was not until the papers of KelnerPillault, late curator of Hymenoptera at the
Museum National dHistoire Naturelle, that
scientific study of this fauna resumed, although for only a very short while. KelnerPillault engaged herself in five short studies
of bees from these deposits. Although her
work was adequate, she did not attempt to
question previous studies nor to compare her
material to prior descriptions, thereby forcing
her specimens into new taxa within groups
as defined by earlier authors. Curiously, Kelner-Pillault published erroneous information
for the type depositories of species she described (e.g., Kelner-Pillault, 1970a, 1970b).
In her papers she regularly refers her material
to the Institut und Museum fur Geologie und
Palaontologie in Gottingen, when nearly all
of her material is in the Institut fur Palaontologie, Museum fur Naturkunde, in Berlin.
It cannot be that the specimens were simply
sent to the wrong institution after publication, since each is accompanied by older labels that catalog them as belonging to Berlin
(and these are often quoted by Kelner-Pillault
in her papers). This unfortunate confusion is
the reason it has been difficult for other
workers to locate and study her material
[e.g., Michener (1990) records that he was
unable to locate Trigona eocenica in Gottin-
NO. 259
gen as indicated in Kelner-Pillaults paper;

the type was actually in Berlin]. All of this
information has been corrected below.
The last major work on the Baltic amber
fauna was as part of a larger attempt to
monograph all fossil bees (Zeuner and Manning, 1976). This work was compiled from
accumulated notes more than a decade after
the deaths of the authors and as such unfortunately suffers from a number of errors.
Most of their monograph does not present
original information and often the sections
for a given taxon are merely taken from the
original descriptions. This is particularly true
for the Baltic amber bees. In several instances it is all too apparent that neither had ever
seen specimens of particular taxa (e.g., see
sections on Ctenoplectrella and Glyptapis,
below). Moreover, in their work some taxa
have been incorrectly diagnosed owing to a
failure to examine types (e.g., A. henshawi;
see Engel, 1998c), and the authorship attributed to numerous groups is wrong. Researchers should be mindful of these difficulties and proceed with caution when attempting to use their monograph. Had both
Zeuner and Manning lived longer, the study
doubtless would have been a more accurate
and beneficial work.
Since the 1970s almost nothing has appeared on the Baltic amber bee fauna. Of the
minor references to Baltic amber bees in recent literature only two papers deserve mention. Both Gerlach (1989) and Engel (1998a)
presented short notes on small collections of
specimens. Gerlach (1989) reported three
specimens misidentified as Dasypoda, Megachile, and Apis (identifications corrected
herein as Glyptapis, Ctenoplectrella?, and
Electrapis, respectively), while Engel
(1998a) described a new species and subgenus in Electrapis and commented on the classification of the genus as it was understood
at that time (that classification is radically altered herein). Although reference to the enigmatic nature of Baltic amber bees has often
been made (e.g., Winston and Michener,
1977; Michener, 1990; Engel, 1998a), as well
as the desperate need for a critical revision
and reexamination of the fauna (e.g., Lutz,
1993), the field has remained essentially untouched.
2001
The present work is an attempt to monograph this unique fauna of bees, to explore
its implications for understanding higher-level affinities of apoid taxa, and to consider
general evolutionary patterns of the bees inclusive of their early origins. This is the first
comprehensive treatment of a fossil bee fauna and is part of a series of papers by the
present author to monograph the fossil bees
of the world, ultimately aimed at unifying
paleomelittology with our neontological
knowledge of bees. Future revisions of this
monograph will undoubtedly be needed as
new or more perfectly preserved specimens
are discovered. For the present, however, I
hope that this work will provide a foundation
for paleoentomological studies of bees. A
catalog and bibliography of the fossil bee
species of the world (both amber inclusions
and compression fossils) is currently in preparation (Engel, in prep.), while a catalog of
family- and genus-group names for fossil
bees is appended at the end of this work (appendix 3).
AGE
AND
ORIGIN
OF
AMBER
Although pockets of Baltic amber can be

found throughout northern Europe (e.g.,
Denmark, Sweden, Germany, Poland, and
Lithuania, among other countries), the greatest concentration, and where most material
originates, occurs on the Samland Peninsula
(historically part of Prussia but today occupied by Russia). The city of Konigsberg (today Kaliningrad), situated near the base of
the Samland Peninsula, once held the most
significant and largest collection of Baltic
amber inclusions owing to the proximity of
the richest deposits. The Albertus Universitat
in Konigsberg was estimated to hold some
100,000 inclusions at one time. This material
was actively studied for a number of years,
particularly by the Koniglichen Physikalisch konomischen Gesellschaft (Royal Physical
O
Economy Society), which began publishing
special works on Baltic amber in 1860 and
continued until 1938. Significant works on
insect inclusions were produced for ants
(Wheeler, 1914), apterygotes (Olfers, 1907;
Silvestri, 1912), and, of course, the studies
by Cockerell (1909b, 1909c) on bees, among
15
others. Tragically, Albertus Universitat suffered, alongside millions of people and other
cultural institutions, during the second World
War, and the collection was mostly destroyed, although some surviving specimens
were scattered for safekeeping. Today, the
largest surviving portion from Konigsbergs
collection is located in the Institut und Museum fur Geologie und Palaontologie in Gottingen, although it is still only a fraction of
the original collections size.
Baltic amber occurs in the blau Erde
(blue Earth), which, as mentioned above,
occurs throughout northern Europe approximately 45 m below the surface and runs ca.
5 m below sea level. Thus, exposed deposits
can erode out and wash amber up on shores
not only of the Baltic countries but as far
away as the eastern coasts of England. The
formation continues under the Baltic sea
floor. Stratigraphic studies of the blau Erde
indicate it to be middle Eocene in age (Kosmowska-Ceranowicz, 1987; KosmowskaCeranowicz and Muller, 1985). The stratigraphic dates correlate perfectly with recent
K-Ar radiometric studies (Ritzkowski, 1997),
all of which indicate that Baltic amber is
44.161.1 Ma, placing it in the middle of the
Lutetian stage of the Eocene Epoch. The Bitterfeld amber (or Saxonian amber) from Germany is actually contemporaneous with Baltic amber and similar in both chemical composition and inclusions of the fauna and flora
(e.g., Schumann and Wendt, 1989; Roschmann and Mohrig, 1995). This Eocene amber
occurs in Miocene strata and has therefore
been considered to be much younger than
true Baltic amber; however, it has been demonstrated that Bitterfeld amber has been redeposited at least twice during its geological
history (Weitschat, 1997). Bitterfeld amber
is, in essence, Baltic amber that has been
eroded and redeposited in Miocene formations. A few of the inclusions studied herein
are in Bitterfeld amber and are identical to
taxa from true Baltic amber (see below).
Pieces of Baltic amber can be readily identified by a number of diagnostic features. The
succinic acid content of most Baltic amber is
relatively high (ca. 8%); this is the reason it
is commonly referred to as succinite. Some
more rare deposits, however, either lack suc-
16
cinic acid (e.g., glessite, gedanite), or possess

it in only trace amounts (e.g., beckerite). The
chemical composition of Baltic amber, however, is relatively uniform and diagnostic by
comparison with all other amber deposits.
Analysis using pyrolysis gas chromatography
of Baltic amber produces a diagnostic set of
peaks (Shedrinsky et al., 1991) and, similarly, infrared spectroscopy analysis produces a
characteristic absorption spectrum with the
unique Baltic shoulder between 1250
1175 cm21 (Beck et al., 1964, 1965; Beck,
1986). Pyrolysis GC analysis of Bitterfeld
amber is identical to that of true Baltic amber
(Shedrinsky et al., 1991). Additionally, Baltic amber frequently includes numerous stellate trichomes (plant hairs), and inclusions
are sometimes covered, either partially or entirely, in Schimmel (a whitish froth of microscopic bubbles resembling mold: e.g., pl. 7a).
Although it has been understood since antiquity that amber (from northern Europe in
particular) is the petrified resin of ancient
trees, the botanical origin of Baltic amber is
still debated (although a final solution is perhaps on the horizon). Both Pliny the Elder
(in his Natural History) and Tacitus (in his
Germania), writing around the same time
during the 1st century, record the ancients
understanding of the resinous origin of amber. Tacitus noted that, The Germans gather
amber but they have not investigated the natural cause or process. . . It is sap [Tacituss
term] from trees, which can be inferred from
the fact that you can see creeping things and
winged insects which were trapped when the
substance gradually hardened, while Pliny
further elaborated on the processes producing
amber when he wrote Amber is formed of
resin seeping from the interior of pine
trees. . . The exudation was hardened by
cold or heat or by the sea. . . That amber
originated as sap is proved by the visible
presence of gnats, ants, and lizards trapped
inside as it hardened. Most authors, like
Pliny, have attributed these resins to a member of the pine family (Pinaceae) and to the
extinct species Pinites succinifer Goppert
(1836). Although arguments have been made
for Araucariaceae (e.g., Langenheim, 1969;
Poinar and Haverkamp, 1985), the pinaceous
hypothesis retains significantly stronger sup-
NO. 259
port. Among the thousands of inclusions in

Baltic amber are numerous cones and needles of pines (e.g., Weitschat and Wichard,
1998) as well as wood fragments with microstructural characters indicative of Pinaceae (Pielinska, 1997; Turkin, 1997). There
are no araucariaceous inclusions in Baltic
amber, and all studies of the botanical inclusions, particularly those of wood fragments,
have not identified any features distinctive to
Araucariaceae (Agathis, in particular). Enigmatically, most pines today do not produce
succinic acid and Baltic amber does not contain the abietic acid that today distinguishes
pine resin; neither does it contain agathic
acid. Recently, however, it has been discovered that both extant and extinct species of
the pine genus Pseudolarix produce succinic
acid and its resin is chemically very similar
to succinite (Anderson and LePage, 1995).
The present day distribution of this genus is
in Asia but included at least the Arctic during
the Eocene. It may be highly significant that
many Baltic amber inclusions have phylogenetic affinities with taxa today living in
Asia (e.g., see summaries in Larsson, 1978).
Baltic amber was likely produced by taxa related to Pseudolarix or even by an extinct
species of this genus.
The Baltic amber forest was, obviously,
not similar to those forests presently occurring in northern Europe. During the middle
Eocene northern Europe was a single landmass. The remainder of the continent was
composed of series of large central and
southern islands or archipelagos in an expanded warm sea. This sea led northward
into the Turgai Straits and Obik Sea that at
the time separated Europe from northern
Asia. Africa and India were still well separated from Europe and Asia, so most biotic
exchange was between the latter two regions,
although some connections were perhaps
possible with North America earlier in the
Paleocene and earliest Eocene (e.g., McKenna, 1975). Global climates in the Paleocene
and Eocene were significantly warmer than
in later Tertiary epochs. The climate in Europe in the Eocene was subtropical or tropical, probably very similar to present day pine
forests in the South Atlantic Coastal Plain of
2001
the United States (i.e., North Carolina to

Florida).
During an era when human understanding
wrestled vigorously with the idea of biotic
and global changes, Rev. Hope (1836) had a
profound insight into the biogeographical association of Baltic amber insects and the climate of the amber forest. Toward the conclusion of his paper summarizing, at the generic
level, those taxa known to him in succinite
he stated:
From the above discrepancies I abstain at present
from entering more largely into the geographical investigation of amber-insects, and reserve it for some
future paper; but from what has already been adduced, may we not conclude that the climate and temperature of Europe have undergone considerable
change? The above examples of tropical insects sufficiently testify that the amber-tree did not vegetate
under a climate such as Prussia now enjoys, but in a
warm region. I trust the above notices will afford a
sufficient stimulus to induce others to take in hand a
subject replete with interest; a subject embracing in
its scope not only the changes of temperature and
climate which our globe has undergone, but also the
consideration of the geographical distribution of insects and plants, by which alone we can arrive at any
satisfactory conclusion respecting them. (Hope, 1836:
137).
It is not of minor significance that Hope

was able to make such broad-reaching conclusions concerning global change and zoogeography from even the most cursory study
of Baltic amber inclusions.
The transition between the Eocene and Oligocene Epochs is a well-documented episode of extinction (e.g., Prothero, 1994). At
this time global temperatures dropped precipitously and the geography of the world was
significantly altered, particularly for Europe.
As temperatures cooled dramatically toward
the end of this epoch the tropical flora and
fauna were gradually extinguished. Likewise,
ocean levels and seaways were altered,
bringing the European and Asian continents
together and significantly shrinking the
southern European ocean to create the Mediterranean Sea we are more familiar with today. Thus, those areas that were once tropical
islands or subtropical mainland forests became part of a colder, more temperate, continual landmass and the amber forest disappeared.
17
MATERIALS AND METHODS

Many institutions throughout the world
maintaining amber collections were contacted or visited regarding the presence of unstudied amber bees among their holdings and
the possible location of lost types. Of the museums and individuals either contacted or
visited I was only able to locate material in
the following: AMNH, American Museum
of Natural History, Division of Invertebrate
Zoology, New York, New York, David A.
Grimaldi; BMNH, The Natural History Museum (British Museum), Department of Palaeontology, London, United Kingdom, Andrew J. Ross; BSPG, Bayerische Staatssammlung fur Palaontologie und Historische
Geologie, Munich, Germany, Helmut Mayr;
GPUH, Geologisch-Palaontologisches Institut und Museum, Universita t Hamburg,
Hamburg, Germany, Wolfgang Weitschat;
IMGP, Institut und Museum fur Geologie
und Palaontologie, Gottingen, Germany,
Hans Jahnke; PMUZ, Palaontologisches
Museum, Universitat Zurich, Zurich, Switzerland; SAMH, Swedish Amber Museum,
Hollviken, Sweden, Leif Brost; ZMHB, Museum fur Naturkunde, Institut fur Palaontologie, Humboldt-Universitat, Berlin, Germany, Erika Pietrzeniuk; ZMPA, Muzeum Ziemi Pan, Polska Akademica Nauk, Warsaw,
Poland, Barbara Kosmowska-Ceranowicz;
ZMUC, Zoological Museum, University of
Copenhagen, Copenhagen, Denmark, Rudolf
Meier. A single type specimen is deposited
in Paris. Unfortunately, this material was not
made available to me for study. The specimen is discussed below and the type depository abbreviated as follows: MNHN, Laboratoire dEntomologie, Museum National
dHistoire Naturelle, Paris, France, Andre
Nel.
In addition, specimens from the following
personal collections were examined (material
couriered through W. Weitschat, GeologischPalaontologisches Institut und Museum,
Hamburg): CCGG, Collection of Carsten
Grohn, Glinde, Germany; CDTG, Collection
of Dirk Teuber, Gutersloh, Germany; CFEG,
Collection of Friedhelm Eichmann, Hannover, Germany; CGHG, Collection of Gunther Herrling, Bramsche-Engter, Germany;
CFKG, Collection of F. Kernegger, Ham-
18
burg, Germany; CHFG, Collection of C. and

H. Hoffeins, Hamburg, Germany; CHMG,
Collection of Hans Mendl, Kempten, Germany; CJDL, Collection of Jonas Damzen,
Vilnius, Lithuania; CJVG, Collection of Jurgen Velten, Idstein, Germany; CMGG, Collection of Matthias Glink, Buxtehude, Germany; CMSE, Collection of Michael S. Engel, Lawrence, Kansas, United States;
CSUL, Collection of Saulius Urbonas and
Juozas Veilandas, Klaipeda, Lithuania. Although his material did not contain any bees
(instead, spheciform and chalcidoid wasps),
I am grateful to Horst Wegner (Hannover,
Germany) for allowing me to examine specimens from his collections. To each of these
collectors I am greatly indebted, particularly
for their willingness to donate name-bearing
types to museums (and therefore listed as belonging to particular museums herein).
SPECIMEN PREPARATION
AND
STUDY
When possible, specimens were vacuumembedded in epoxy following the procedure

of Nascimbene and Silverstein (2000). Water-fed trimming and polishing was then undertaken on most specimens so as to make
clear windows for the observation of particular structures and photography. Amber
pieces were then examined using a Zeiss Stereomicroscope SV-8 and measurements made
using an ocular micrometer. Scanning electron microscopy was done on uncoated specimens using a Hitachi S4700 Field Emission
Scanning Electron Microscope for partial inclusions of Boreallodape striebichi preserved at the amber surface (see below). I
prepared a photographic slide of each specimen [a few of which are presented here as
either color plates or black-and-white images
(e.g., the frontispiece)], thereby creating a
photographic library of bees in Baltic amber.
FORMAT
In preparing this work the recently introduced fourth edition of the International
Code of Zoological Nomenclature has been
followed (ICZN, 1999b). Standard formats
are used for taxonomic histories. References
in each line refer only to taxonomic works
and thus if a species was simply cited or listed by an author such a reference is excluded.
NO. 259
In presenting taxonomic summaries for

family-group names only those names applicable to groups under consideration are presented. For example, under the subfamily
Megachilinae the nominate tribe is not considered in the present monograph owing to
the absence of this group from the Baltic amber fauna. The taxonomic history under Megachilinae therefore presents only the derivation of the family-group name based on
Megachile and not all of the other names for
valid tribes (e.g., Anthidiini, Dioxyini). Nor
are synonyms of the nominate tribe included
(in this particular example, synonyms of Megachilini, like Coelioxynae), since such synonyms are probably not equivalent with the
group at a higher rank (e.g., in this work the
family-group name based on Coelioxys
should not be listed as synonymous with Megachilinae, since the former is not equivalent
to a group inclusive of Anthidiini or other
tribes of the subfamily but only with the Megachilini). The taxonomic history is presented only at its lowest rank within the familygroup so as to avoid extensive repetition
(e.g., the taxonomic history of the familygroup name based on Halictus is presented
under tribe Halictini and not also under Halictinae and Halictidae). For extant bees complete details on family-group names and their
synonyms are presented by Michener (1986)
with additions and corrections by Michener
(1997) and Engel (1999e). A catalog of family- and genus-group names for fossil bees is
presented in appendix 3, while a separate
species catalog is in preparation (Engel, in
prep.).
I have used the term nomen translatum for
family-group names that have been changed
in hierarchical rank (e.g., tribe elevated to
subfamily) and required an alteration of suffix. The term status novus is reserved for
changes in rank whereby the name retains its
suffix from the former rank without alteration (e.g., subspecies elevated to species,
subgenus elevated to genus). As stipulated by
the Principle of Coordination (ICZN, 1999b:
Art. 36), names altered in either manner retain their original authorship and date regardless of rank.
In the Material sections I have presented
detailed label information from each specimen as well as a summary of the total num-
2001
ber of specimens (immediately following the

heading Material for each species). In these
summaries a double slash (//) separates text
from individual labels. The material sections
are broken into paragraphs, with each paragraph representing a single amber piece except when the holotype and paratypes occur
in the same piece; otherwise multiple individuals in a single amber block are treated
within the same paragraph. Each paragraph
is headed by a boldface indication as to
whether this material is a type or not. When
two or more taxa are synonymized there are
two paragraphs with the boldface heading
holotype, lectotype, or neotype, but
each is followed in parentheses by an indication as to which synonymous name it belongs. When the holotype for one species is
the same specimen designated as the neotype
for another, these begin separate paragraphs
but are cross-referenced to avoid repeating
label data.
As a result of attempting to forge a new
path, I have had to make difficult decisions
regarding previously described taxa for
which specimens have since become lost or
destroyed, mostly through the ravages of
world war. I therefore found it necessary to
designate neotypes and in so doing to carefully attempt to associate newer material with
often inadequate and incomplete original descriptions. This has been done expressly for
the purpose of stabilizing the nomenclature
of fossil bees (in accordance with ICZN,
1999b: Art. 75.3.1) and thereby allowing and
encouraging meaningful comparison of these
taxa with Recent bees. For all of the taxa
below I have provided necessary information
for both the recognition of the specimens
chosen as neotypes, my reasons for believing
them to be conspecific with the original material, and diagnoses of all of these species
(meeting the requirements of ICZN, 1999b:
Art. 75.3.2, 75.3.3, and 75.3.5). Article
75.3.6 (ICZN, 1999b) states that the neotype
must come . . . as nearly as practicable
from the original type locality.; since all of
the original fossils, as well as my own, originate from middle Eocene strata of the Baltic
region, I believe this criterion to have been
sufficiently met. All neotypes are either in
the AMNH, ZMHB, GPUH, or BMNH as
indicated below, making them accessible for
19
future study (satisfying ICZN, 1999b: Art.

75.3.7). The more difficult criterion to meet
is Article 75.3.4, which requires reasons for
. . . believing the name-bearing type specimen(s). . . to be lost or destroyed, and the
steps that had been taken to trace it or them.
All of the species for which neotypes were
required were originally located in the Albertus Universitat in Konigsberg. During
World War II the collections of this university were destroyed by fire. I contacted numerous institutions in search of described or
undescribed material. The bulk of Konigsbergs surviving material found its way into
the collection of the Institut und Museum fur
Geologie und Palaontologie, Gottingen. I
personally visited this institution during the
summer of 1999 (as well as several other European collections) and inspected the entire
amber collection. From this material I was
only able to locate three original specimens
(with Cockerells own handwritten labels). I
was therefore left with the conclusion that
the remaining taxa, which could also not be
traced in any other museum, must have been
lost or destroyed during the tumultuous years
of the last world war. For these reasons I believe to have adequately satisfied all of the
ICZN (1999b) requirements, inclusive of Article 75.3.4, for the designation of neotypes.
Metrics are provided in the descriptions
for the name-bearing type specimen. All
measurements should be considered approximate, since it was not always possible to
achieve the optimal angle for making a given
measure. In the description of higher-rank
groups, italicized characters indicate features
that could not be confirmed for the fossils
described within that particular group. The
following descriptive abbreviations are used:
F, flagellomere; OD, ocellar diameter (always
based on the diameter of the median ocellus);
S, metasomal sternum; T, metasomal tergum.
Descriptions are of females (of the worker
caste where applicable in the corbiculate Apinae) except for Ctenoplectrella viridiceps
where a description of the male is included;
otherwise males are unknown.
GENERAL MORPHOLOGY
Numerous accounts discuss the general external morphology of bees (e.g., Michener,
20
1944, 1965, 2000a; Snodgrass, 1956; Camargo et al., 1967; Urban, 1967; Eickwort,
1969; Pesenko, 1983; Engel, 2000a) and the
ensuing section is not meant to entirely supplant those. Instead, I have attempted to summarize standard terms for the external morphology of bees, to relate each characteristic
to gross variations found among the Baltic
amber fauna, and to provide new terms for
features not previously considered. This section is also intended to provide a working
terminology for researchers and amateurs
collecting, organizing, and studying Baltic
amber inclusions. It is hoped that this overview of bee morphology will make the subsequent systematic section and its included
descriptions and dichotomous keys more
useful.
Morphological terminology generally follows that proposed by Michener (1944,
2000a) with additions for mandibular structure by Michener and Fraser (1978), coxal
structure by Michener (1981a), and glossal
morphology by Michener and Brooks (1984).
From these accounts, however, I differ in
some terminologies for wing veins, mouthpart structures, and leg structures most notably in terms used for types of pollen-carrying apparati (see specific instances below).
The following sections provide greater detail
into the precise morphology of bees in general and the Baltic amber bees in particular,
defining terms frequently used throughout
the text.
PROSOMA
The head, or prosoma, is the first body
tagma and is hypognathous. The compound
eyes (figs. 1, 4) are situated laterally on the
head and are composed of numerous minute
facets representing the individual lenses of
the ommatidia. At the top of the face, arranged in the pattern of an inverted triangle,
are three simple eyes termed ocelli (fig. 1).
At the extreme lower apex of the head is the
labrum (figs. 1, 3), a freely movable sclerite
that at its base broadly articulates with the
remainder of the prosoma. The labrum is
sometimes composed of two sections: the
distal process and the basal area. The distal
process (fig. 1) is a medioapical extension
that extends apicad from the transverse basal
NO. 259
area (fig. 1) and is slightly set below the

plane of the basal area when viewed in profile; the basal area is the portion that articulates with the head capsule. When the distal
process is present it is variously modified
with either lateral teeth or serrations, a dorsal
ridge or keel, or tubercles of differing morphologies. Similarly, the basal area can bear
tubercles and/or ridges, and is found in a variety of gross morphological shapes, none of
which depend on the presence or absence of
the distal process. The labrum broadly articulates at the apical margin of the clypeus
(figs. 1, 3), the lowermost, immovable sclerite of the head capsule. The basal attachment
of the clypeus to the remainder of the head
is delimited by the epistomal sulcus (fig. 1)
(sometimes called the epistomal suture),
which laterally forms an angle of various degrees of arc; typically the angle is measured
between the two portions of the sulcus opening toward the compound eye. Similarly the
dorsalmost, transverse portion of the epistomal sulcus ranges from convex (i.e., bending
upward and extending the clypeus dorsally),
straight, to concave (i.e., bending downward
and shortening the clypeus). Almost universally (except in some living genera, such as
Nomioides), the anterior tentorial pits are set
into the epistomal sulcus, frequently along its
lateral parts. Extending dorsally from the epistomal sulcus to the antennal sockets are the
subantennal sutures (fig. 1). Typically there
is a single subantennal suture reaching each
antennal socket, although in some groups
(e.g., Andrenidae) there are two subantennal
sutures reaching each antennal socket and
defining between them a small subantennal
area. Frequently, the subantennal suture
meets the antennal socket along its lower
margin, although in the Lithurginae and Megachilinae the single subantennal suture runs
to the outer, lateral margin (i.e., that side
nearest the compound eye) of the antennal
socket. The antennal sockets (or alveoli) (fig.
1) are the points at which the antennae articulate and, together with the subantennal sutures, serve to demarcate a poorly defined region of the head referred to as the supraclypeal area (figs. 1, 3). The face above the
level of the antennal sockets to the level of
the ocelli and between is the frons (fig. 1).
The ill-defined region of the face running
2001
21
Fig. 1. Diagrammatic representation of a bees head (5 prosoma) with major morphological structures labeled. Left image is a frontal view of the head with the antennae and mandibles removed; right
image is a ventral view of the head.
along the inner margin of the compound eye

and outside of the frons is the paraocular
area, which is sometimes equipped with a
carina running dorsoventrally along the compound eye (i.e., the paraocular carina: fig.
1). In some bee lineages (e.g., Andrenidae)
the paraocular area near the upper, inner margin of the compound eye bears a shallow integumental impression or facial fovea (fig.
1), sometimes covered with dense, microscopic setae. At the lower extremity of the
compound eye is the malar area (fig. 1), a
small region of integument separating the
lower border of the compound eye from the
mandibular articulation (see below). In some
bee taxa there is an interocellar furrow
formed of an impressed line running between
and just posterior to the lateral ocelli (Engel,
1998b, 2000a). On the top of the head and
immediately behind the ocelli is the vertex
(figs. 1, 4), which runs to the posterior margin of the head where the head once again
angles or curves ventrally to its posterior surface, or occiput (fig. 1). The ridge separating
the vertex from the occiput is the preoccipital ridge (fig. 3) and surrounds the concave,
posterior surface of the head (i.e., the occiput), which itself finally gives rise centrally
to the postocciput that is surrounded by distinct postoccipital sulcus (or suture). The
preoccipital ridge can be either rounded, angled, carinate, or even lamellate. The postoc-
ciput bears a large opening, the foramen

magnum, where the prosoma and mesosoma
connect. To the sides of the vertex the lateral
portion of the head behind the eyes and ocelli
and anterior to the preoccipital ridge is the
gena (fig. 3), which ventrally gives way to
the postgena (fig. 1) on the ventral surface
of the head. The postgena borders a longitudinal concavity composed of the hypostoma on the undersurface of the head and is
referred to as either the hypostomal fossa or
the proboscidial fossa, since the proboscis
(i.e., the labium and maxilla) rests within this
region when in repose. The ridge separating
the hypostoma from the postgena is the hypostomal ridge (fig. 1) and, like the preoccipital ridge, can be rounded, carinate, or
even lamellate and frequently projects slightly beyond the occiput posteriorly. Anteriorly
the hypostomal ridge curves laterally (fig. 1)
and runs behind the mandibular articulation.
Articulating with the head are a number of
appendages representing ancestral remnants
of a once segmented, limbed tagma of a debated total number of somites. The appendages discussed here are not presented in order from anterior to posterior of their presumed relative position on ancestral segments. At the lower, apicolateral extremity of
the head are the mandibles (figs. 1, 3). The
mandibles of bees are variably structured and
typically sexually dimorphic. Numerous an-
22
atomical details concerning mandibular

structure were covered by Michener and Fraser (1978) and it is not necessary to repeat
that material here. Instead, I shall focus on
only those gross structural features that have
been observed in the fossils discussed herein
and require some explanation of their associated terms. The mandibles, as in all pterygote and some apterygote insects, are dicondylic, having two points of articulation
and therefore moving in a single plane. The
articulation is formed of an anterior mandibular acetabulum and a posterior condyle each
corresponding to an anterior condyle and
posterior acetablum on the head capsule, respectively. The apical margin of the mandible is frequently beset with a series of incisions defining apical teeth. Along the upper
margin the subapical tooth (figs. 1, 3) and a
groove (termed the acetabular groove) running from the mandibular acetabulum demarcate a region referred to as the mandibular pollex, while the remaining area of the
mandible is the rutellum. The mandibular
pollex is in some groups (e.g., the corbiculate
Apinae) greatly expanded to form a single,
elongate cutting margin, or is divided into
additional teeth. On the outer margin of the
rutellum are a series of ridges that delimit an
associated number of interspaces. Along the
lower margin of the mandile runs the condylar ridge (below it is the condylar groove)
and between this ridge and the acetabular
groove is the outer surface of the mandible
with its outer grooves, the number and presence of which can vary among bee lineages
(e.g., completely absent in some corbiculate
Apinae, such as the honey bees).
The appendages of the next two head segments are intimately tied together and form
the primary mouthparts of the bee. Frequently referred to as the proboscis or the labiomaxillary complex, these structures represent
the maxillae, from the segment immediately
behind that of the mandibles, and the labium,
from immediately behind the maxillary segment, united basally by membranous connections which are typically strengthened anteriorly by thin, sclerotic conjuctival thickenings. The overall structure of the labiomaxillary complex as viewed from the front of a
bee where the proboscis is fully extended
(e.g., fig. 2) can be thought of as the lateral,
NO. 259
paired maxillae and the central labium. The

bases of the maxillae are the site of attachment and articulation of the labiomaxillary
complex to the head capsule. The basalmost
segment of the maxilla is the cardo (fig. 2),
an elongate sclerite that attaches to the head
capsule. At the apex of the cardo is the stipes
(fig. 2), the next prepalpal segment of the
maxilla, as well as the submentum (or lorum), the basalmost sclerite of the labium
(see below). The ventral (lateral surface in
fig. 2) margins of the stipes is sometimes
concave with a distinct comb of stiff setae
set within the concavity and termed the stipital comb (e.g., present in many Apidae). At
the apex of the stipes is the frequently sixsegmented maxillary palpus (fig. 2). Two
sclerites form the postpalpal portion of the
maxilla and are the elongate galea and the
small lacinia (fig. 2). Like the stipes, the galea is frequently beset (in the short-tongued
bee families) with a comb of stiff setae on
its inner surface, which when present is referred to as the galeal comb. As mentioned
above the basalmost sclerite of the labium is
the submentum (fig. 2), which is often termed
the lorum to indicate its dubious homology
(Snodgrass, 1935) with the true submentum
of other insects. As discussed elsewhere (Engel, 1999b), I prefer the term submentum as
employed by Michener (1944) in place of lorum. The submentum is shaped like an inverted V in many bees (except in groups
such as some Colletidae it is more of a broad
inverted triangle, or in Halictidae, rectangular or membranous) with the apices of the
submental arms meeting the cardines of the
maxillae. Along the inner, subapical surface
of the submental arm is sometimes a small
spine referred to as the submental spine (fig.
2), most notably found in the corbiculate Apinae. Medially the submentum attaches to the
mentum, a structure, sometimes absent or
membranous, that at its opposite end meets
the base of the prementum, an elongate sclerite (fig. 2). The apex of the prementum is
trilobed with the lateral processes supporting
the frequently four-segmented labial palpi
and a more broad, medioapical (or subligular) process that varies in length (being either shorter or longer than the lateral processes) (fig. 2). Postpalpally on the labium
are the lateral paraglossae and the medial
2001
glossa (fig. 2). As is discussed briefly below

(under Systematic Paleontology) the mouthparts of bees can be grossly classified into
two structural categories: short-tongued and
long-tongued. Most notably, in short-tongued
bees the first two segments of the labial palpus are cylindrical and usually more or less
equal in length to the distal two segments,
while in long-tongued bees the basal two
segments are greatly elongate and flattened.
In short-tongued bees the glossa is frequently
short (and, in the Colletidae, broadened or
usually bilobed at its apex as in the spheciform wasps), while it is pointed and almost
universally elongate in the long-tongued
bees. The premental fragmentum is a detached basal portion of the prementum. This
structure is involved in the articulation of the
prementum with the mentum in melittid and
other bees. In long-tongued bees the apex of
the glossa bears a minute additional structure
termed the flabellum (fig. 2). Numerous other
characters of the labiomaxillary complex
could be discussed at some length, but since
they are hidden in most bee fossils, I will not
treat them further here (but see Michener,
1944, 2000a; Snodgrass, 1956; Eickwort,
1969).
Lastly, the antennae are somewhat geniculate, bending at the pedicel, and, as mentioned above, articulate with the face above
the level of the clypeus (figs. 3, 4). The antennae are composed of three units: a basalmost scape that articulates via a basal bulb
with the antennal socket; a short, intermediary pedicel; and the typically filiform flagellum (rarely serrate in some odd, living genera), which is itself divided into a series of
individual, cylindrical flagellomeres (figs. 3,
4), sometimes called flagellar segments. In
bees the number of flagellomeres is dimorphic: females almost always having 10 and
males 11. The relative lengths of the basal
flagellomeres can be useful characters for
separating groups of bees.
MESOSOMA
The mesosoma is a combination of the true
thoracic tagma with the first abdominal segment (5 propodeum). It is thus composed of
four primary segments: prothorax, mesothorax, metathorax, and propodeum. The meso-
23
and metathoracic segments bear the wing articulations and are sometimes together referred to as the pterothorax. The major segmental structures are prefixed by pro,
meso, or meta as a reference to which
thoracic segment they relate (e.g., mesocoxa,
metatibial spurs).
The pronotum, or dorsal surface of the first
thoracic segment is expanded laterally to
form lateral surfaces and processes (the postcoxal processes) that extend along the sides
and behind the procoxae to form a complete
or nearly complete ring tightly adjoining the
anterior margin of the mesothorax (figs. 3,
4). Dorsally the pronotum extends somewhat
laterally and is posteriorly broadly U-shaped;
the shape of the dorsolateral angle (fig. 4),
where the pronotum bends posteriorly, is a
useful taxonomic character. Along the posterior border of the pronotal lateral surface
and extending toward and just below the level of the mesothoracic wing base is a distinct
pronotal lobe (fig. 3) that covers the first thoracic spiracle. The presence of this lobe is a
synapomorphic trait of the superfamily Apoidea. As a result of this overall pronotal morphology the propleura face ventrally and lie
anterior to the procoxae (fig. 3). The prosternum is mostly obscured from view between the propleura but can be viewed internally. The anterior, triangle-shaped apex of
the prosternum has been termed by Snodgrass (1956) the basisternum, while the
broad, posteriorly-projecting portion is the
furcasternum. The overall shape of the prosternum varies among higher bee lineages
(e.g., tribes). The lateral processes and apical
extension of the basisternum are of varying
lengths and the furcasternum is similarly variable, sometimes with a more or less developed longitudinal, medial groove. Most notable is the variation in construction exhibited among the corbiculate Apinae (e.g.,
Prentice, 1991) where, in the Apini and Meliponini, there is a strong constriction immediately posterior to the basisternum (i.e.,
between the basisternum and furcasternum).
The mesonotum, or dorsal surface of the
mesothoracic segment, is divided into two
primary units: the mesoscutum (or scutum)
and the mesoscutellum (or scutellum). The
mesoscutum is the larger, anterior sclerite the
mesonotum (figs. 3, 4). It is laterally bor-
24
NO. 259
Fig. 2. Diagrammatic representation of a long-tongued bees labiomaxillary complex with major

structures labeled.
dered by small movable sclerites termed tegulae that cover the mesothoracic wing bases
(figs. 3, 4). Borne upon the mesoscutum are
a series of longitudinal, impressed lines. The
median line (fig. 4), as the name implies,
runs medially from the anterior border frequently to about the midpoint of the mesos-
cutum. The notauli are short, submedial lines

that run from the anterior border to about
one-third of the mesoscutal length in their
most extreme case (fig. 4). The parapsidal
lines are also parallel to the median line, although closer to the lateral, mesoscutal borders (fig. 4). Posterior to the mesoscutum is
Fig. 3. Diagrammatic representation of a left lateral view of a bee with major morphological features labeled. NOTE: The figure combines
characters not found in combination in any known bee (living or fossil); graduli not illustrated for all terga or sterna.
2001
25
26
the scutellum, the second sclerite of the mesonotum (figs. 3, 4). The scutellum is frequently dorsally gently convex, while laterally it is concave and forms regions referred
to as axillae (figs. 3, 4). The axillae are immediately posterior to the mesothoracic wing
bases. The mesepisternum (figs. 3, 4) is the
large, lateral portion of the mesothoracic segment. A series of ridges and grooves run
across this surface and these structures are
important characters for the recognition of
suprageneric groups within bees. Running
from just below the mesothoracic wing base
toward the ventral margin is the preepisternal groove, which serves to define the posterior border of a region termed the preepisternal area or preepisternum (fig. 3). If the
mesepisternum is strongly bent, then the
preepisternal area sometimes includes an anterior-facing surface. The preepisternal
groove does not define the ridge separating
this anterior-facing surface from the lateral
surface of the mesepisternum; this ridge is
termed the omaular ridge and can be rounded, carinate, lamellate, or rarely entirely absent. Running between the preepisternal
groove and the posterior border of the mesepisternum is a short, longitudinal, impressed line that crosses a pit at about its
midpoint. This pit is termed the scrobe or
episternal scrobe, while the impressed line is
termed the scrobal groove (fig. 3). Above the
scrobal groove is a usually convex region of
frequently distinctive sculpturing referred to
as the hypoepimeral area (fig. 3). The subpleural signum is a small, impunctate, frequently raised spot at the separation of the
pleural and ventral portions of the mespisternum. The precoxal triangle is a distinctive
area of sculpturing on the mesepisternum
bordering the mesocoxa in glyptapine bees
(fig. 3). The precoxal triangle is delimited by
a carina on all sides (posteriorly this carina
is the precoxal carina). The integumental
sculpturing within the precoxal triangle or
the structure of its separation from the remainder of the mesepisternum is quite useful
for separating species of the genus Glyptapis.
The metathoracic segment is greatly shortened longitudinally by comparison to the mesothorax. Dorsally it is represented by the
metanotum (figs. 3, 4) immediately posterior
to the scutellum, which may sometimes pro-
NO. 259
ject over its surface to varying degrees. Laterally the metepisternum (fig. 3) runs from
the metathoracic wing base (fig. 4) to the
metacoxa and typically narrows ventrally. As
mentioned previously, the hindmost portion
of the mesosoma is the propodeum (figs. 3,
4), which is ancestrally the first segment of
the abdomen. The propodeum is frequently
angled basally to form a dorsal-facing basal
area and a posterior surface (figs. 3, 4). Typically, the sculpturing of the integument on
the basal area of the propodeum is diagnostic
for species or groups of species. Similarly,
the angle between the basal area and the posterior surface can be useful for the definition
of generic groups among bees. The propodeal triangle (5 metapostnotum of Brothers,
1976) is a distinctive region of integument
including the basal area or most of it and
extending posteroventrally toward the propodeal pit (fig. 4) on the posterior surface.
The mesosomal appendages are all born
on the true thorax. The legs are composed of
the typical series of segments (from base to
apex): coxa, trochanter, femur, tibia, tarsus,
and pretarsus; the tarsus is further subdivided into the basitarsus (basalmost subsegment
or tarsomere), mediotarsus (subsegments two
through four), and distitarsus (distalmost
subsegment) (fig. 3). The procoxae, like the
propleura, are situated next to one another on
the anteroventral surface of the mesosoma.
The meso- and metacoxae are typically more
well separated from one another, situated on
the ventral surface of the posterior half of the
mesosoma and closer to one another than either is to the procoxae. In most short-tongued
bees families (i.e., Colletidae, Halictidae,
Andrenidae) the mesocoxa is hemicryptic,
that is to say its dorsal articulation is partially
hidden inside of the mesosoma. All other bee
families (i.e., Paleomelittidae, Melittidae,
Megachilidae, and Apidae) have a fully exposed mesocoxa. The metacoxa is fully exposed and ventral to the metepisternum and
anterior portion of the propodeum. The protrochanters are slightly laterally compressed
segments that articulate in a longitudinal
plane (rather than vertical plane of movement as in the meso- and metatrochanters).
The femora and tibiae are elongate, tubular
segments, typically widest at their midpoint.
On the ventral, mediobasal surface of the
2001
27
Fig. 4. Diagrammatic representation of a bees prosomal and mesosomal dorsum with major anatomical structures labeled.
mesofemora is a dense, longitudinal series of

short setae that form the mesofemoral comb.
Similarly, on the inner, medioapical surface
of the mesotibia is a dense, longitudinal series of setae termed the mesotibial comb. The
metathoracic legs are typically modified for
the transport of pollen (although this is by no
means universal for bees; e.g., in Megachilidae pollen is carried on the metasoma). The
scopa is developed to varying degrees and
consists of elongate, frequently plumose setae on a variety of metathoracic leg segments
of females except in parasitic species and the
queen caste of some eusocial species. The leg

segments bearing the scopal setae are useful
characters for the separation of higher lineages among bees (e.g., in the extreme case
the complete absence of scopal setae on the
legs helps to characterize numerous parasitic
taxa as well as groups such as the megachilids, where the scopal setae are born on the
metasomal sterna). Thus, among bees there
are metasomal scopae, femoral scopae, tibial
scopae, &c. Perhaps the most distinctive pollen-carrying apparatus is the corbicula or
pollen basket (fig. 5). The corbicula is
28
formed by an expansion of the metatibia toward its apex; it is somewhat flattened and
the outer surface concave to varying degrees.
The concave region on the outer surface
(varying in length, although typically comprising only the apical half of the metatibia)
is bordered anteriorly and posteriorly by
elongate scopal setae. Michener (1999) has,
as have some other authors (e.g., Thorp,
1979), used the term corbicula more inclusively for any glabrous, frequently concave
surface bordered by fringes of elongate scopal setae and used for the transport of pollen.
Thus some structures found in other bee
groups (e.g., andrenines, halictines) may
have what these authors refer to as femoral
corbiculae or propodeal corbiculae. In
attempting to denote the uniqueness of the
actual structure found only in the corbiculate
Apinae, i.e., those taxa not only exhibiting
the corbicula as defined by Michener (1999)
but also the associated modifications of the
metatibia, I prefer to restrict the term corbicula to this structure. Those other pollen-carrying apparati termed corbiculae by Michener (op. cit.), are here referred to as either
corbiculate scopae or more appropriately as
fiscinae (sing. fiscina: L. meaning flower or
bread basket). Thus corbiculae s.s. are homologous structures found in a single clade
of apine bees. Both scopae and fiscinae are
more general terms for other pollen-carrying
structures and are not necessarily homologous across taxa. Thus, it may prove at a
later time to be useful to indicate such homologies by their own terms owing to the
diversity of fiscinae among bees (e.g., the
term canistra for a propodeal fascina).
The inner surface of the metatibia of many
bees and wasps is covered by a dense field
of minute, blunt setae termed keirotrichiae.
The region occupied by the keirotrichiae is
termed the keirotrichiate field (fig. 5) and its
separation from the remainder of the metatibial inner surface is a useful systematic
character among Meliponini and Electrapini.
Articulated on the inner apex of the tibiae are
spurs, the number and general structure of
which vary on legs of differing segments;
there is a single protibial spur and a single
mesotibial spur, while the inner apex of the
metatibia ordinarily has two metatibial spurs
(figs. 3, 5). The protibial spur is modified
NO. 259
into part of an antenna cleaner or strigilis

(fig. 3). The thickened, spur backbone of
the strigilis is the malus, while along its inner
basal margin is a broad lamella termed the
velum (fig. 3). The overall shape of the velum does not vary greatly (typically it is
roughly rectangular, but sometimes the outer
margin of the lamella is slightly concave),
although some corbiculate Apinae (e.g.,
Bombini, Electrobombini) there is a second
anterior velum formed from a thickening
along the anterior border of the malus and
approximately orthogonal to the true velum;
the anterior velum is not always developed
into a lamella like the true velum. The distal
portion of the malus that extends beyond the
velum is either serrate, ciliate, or pectinate
with a series of long teeth gradually decreasing in size apically. The mesotibial spur,
while varying to some degree in length, is
almost universally serrate or minutely ciliate.
Among some living bee lineages there are
other minor modifications of its apex, e.g.,
within Ericrocidini. The presence/absence or
shape of the metatibial spurs varies dramatically among bee groups. In some lineages
(e.g., the Meliponini and Apini) the spurs are
entirely absent, while in several other groups
a single metatibial spur is present (e.g., Electrapini, Melikertini). The spurs are typically
of one of the three morphologies: serrate, ciliate, or pectinate; when two metatibial spurs
are present the inner and outer spurs are not
necessarily of the same morphological type.
Along the inner, apical margin of the metatibia of corbiculate apines is a tight series of
short, stiff setae known as the rastellum (fig.
5). In the Meliponini the outer surface of the
corbicula near the posteroapical corner there
is a penicillum (fig. 5), or dense patch of
modified setae. As mentioned above, the tarsi
are composed of five subsegments, the first
of which is always greatly elongate by comparison to the distal four, and is referred to
as the basitarsus. On the inner, basal surface
of the probasitarsus is a deep concavity corresponding in relative position to the velum
of the strigilis and serves as part of the antenna cleaning apparatus. This concavity is
the strigilar concavity (fig. 3) and is frequently equipped with a linear series of fine
setae to form a strigilar comb. The probasitarsus sometimes also bears a tight, longitu-
2001
dinal series of setae referred to as a probasitarsal comb; non-homologous combs occur

in a variety of positions and serve differing
functions (e.g., Augochlorini, Tapinotaspidini). The mesobasitarsus is little different
from the probasitarsus except that it lacks the
stigilar concavity. The metabasitarsus is
broader than the metamediotarsus and metadistitarsus, although the degree of its lengthto-breadth ratio ranges dramatically across
bee lineages. In some tribes of the corbiculate Apinae (i.e., Euglossini, Bombini, Electrobombini, Electrapini, Melikertini, Apini)
the base of the metabasitarsus is modified
into an auricle or pollen press (fig. 5). The
auricle is formed of a slightly concave surface bordered by a carina or lip that is particularly broadly developed along the outer
margin. The microsculpturing of this concavity is typically equipped with rows of microscopic, broadly flattened, pointed spicules directed toward the outer margin. The auricle,
which is used to compress and move pollen
into the corbicula for transport, is sometimes
bordered along its outer margin by a series
of more or less stiff setae here referred to as
the auricular basket. The posterior apical or
outer apical corner of the metabasitarsus is
sometimes extended apically into a distal
process beyond the articulation of the metabasitarsus to the metamediotarsus. This distal
process sometimes bears a penicillus (not to
be confused with the penicillum on the metatibia of Meliponini) or brush of dense,
modified setae. The distatarsus is frequently
slightly more elongate than the subsegments
of the mediotarsus. The pretarsi bear the
paired claws, which themselves sometimes
bear an inner notch of varying depths to form
an inner tooth (fig. 3). Toothed claws occur
in most bees, although some lineages (e.g.,
Megachilini) have simple claws. Between the
claws is a membranous lobe or arolium (fig.
3), the presence or absence of which is an
important character for distinguishing taxa in
a variety of lineages. A number of other associated structures are present in the protarsus [these are covered in greater depth by
Michener (1944)]; none of these structures
were observed for the Baltic amber bees
(owing to their minute size and optical limitations).
The wings are borne on the meso- and
29
metathoracic segments and are joined to the

mesosoma via a series of complex, minute
axillary sclerites. For further details on the
morphology of the axillary sclerites in bees
refer to Michener (1944) and Snodgrass
(1956). The wings are formed of a membrane
braced by longitudinal, sclerotic veins (abbreviated by capital letters) that are themselves, when not fused, connected at various
points by transverse crossveins (abbreviated
by lowercase letters). At the wing base the
longitudinal veins in the forewing are (from
anterior to posterior: fig. 6) the costa (C), the
fused subcosta and radius (Sc1R), the media
fused with the cubitus (M1Cu), and the first
anal (A); the radius further out in the wing
splits to form a posterior branch termed the
radial sector (Rs). At roughly one-third of
the total forewing length from the base, the
media and cubitus separate and follow separate courses, the former eventually fuses for
a short distance with the radial sector before
separating once again shortly thereafter (fig.
6); the first abscissa of M (the free portion
of this vein immediately after its separation
from Cu) is termed the basal vein. In the
hind wing the longitudinal veins are Sc1R
(the costa is lost in the hind wing, leaving an
unbraced, membranous leading edge),
M1Cu, and A. Along the leading edge of the
forewing there is an enlarged, sclerotized region (that weighs the leading edge of the
wing) termed the pterostigma and formed of
an expansion in R. Immediately basal to the
pterostigma is a region formed of an expansion of Sc1R termed the prestigma. The basalmost edge of the prestigma is defined by
the point at which the first abscissa of Rs
originates from Sc1R. The leading edge of
the hind wing is equipped with a series of
minute hooks termed hamuli (fig. 7), which
link the two sets of wings in flight. The number and pattern of arrangement of the hamuli
is variable across bee taxa and is a useful
taxonomic character. Crossveins are denoted
by two letters indicating which longitudinal
veins they connect; anterior vein first, posterior second (i.e., rs-m runs between Rs and
M). Sections of the longitudinal veins running between two crossveins are termed abscissae and are numbered from base to apex;
similarly, when more than one crossvein occur between a pair of longitudinal veins they
30
NO. 259
Fig. 5. Diagrammatic, corbiculate metatibiae with structures labeled. Left metatibia depicts an outer
surface; right metatibia depicts an inner surface. NOTE: The figured metatibiae are not meant to represent outer and inner surfaces of the same leg or a single species.
are numbered consecutively from base to

apex and this number precedes the crossvein
abbreviation (e.g., 2m-cu). At specific locations certain veins are crossed by minute
clearings of sclerotization termed alar fenestrae that indicate flexion lines where the
wing will bend or fold during the motion of
flight. Some of these lines are indicated in
wing illustrations by dotted lines crossing the
submarginal cells and their associated veins.
Fusion and confusion would be an appropriate descriptor for the pattern of veins in the
Hymenoptera forewing and its changes
through the evolutionary history of the order.
It would be excessive to discuss at length
every point at which two veins merge, separate, remerge, and eventually terminate.
Thus, for the sake of brevity, I refrain from
such a discussion here and refer the reader
to figure 6 where all of the veins are labeled.
It must be noted that some of the vein terms
used here differ from other works on bees. I
have attempted to standardize the wing vein
terms of the Apoidea with general work on
the order so as to indicate the homologies of

these veins not only among superfamilies of
Hymenoptera but, as best as can be done,
across insect orders. A few of these differences are as follows: 1. the anal vein is often
referred to as the vannal vein (abbreviated as
V) in other works on bees; 2. those veins
delimiting the apical edge of each submarginal cell and often referred to as the transverse cubitals (despite having no association
with the cubitus!), or more appropriately the
submarginal veins (referred to as transverse
submarginals in Michener, 2000a), are considered to be herein, from basal to apical, the
second abscissa of Rs, 1rs-m, and 2rs-m
(when only two submarginal cells are present
these veins are second abscissa of Rs and
2rs-m; see below), respectively; in other accounts where the vernacular terms are not
employed the indication that the latter two
crossveins arise from Rs is dropped and they
are termed 1r-m and 2r-m; 3. in a similar
fashion the small crossvein running between
the pterostigma (R) and the posterior border
2001
of the marginal cell (formed of Rs) just beyond the first submarginal cell is actually the
r-rs crossvein but is frequently and enigmatically referred to as simply r, presumably
owing to the fact that this crossvein runs between two portions of the radial system; 4.
in the hind wing the crossvein r-m is changed
to rs-m to regain, once again, more information pertaining to which section of the radial system this vein connects to the medial
system. Thus, although these are minor alterations to the terminology of bee wing
veins, they more accurately reflect the identity of the veins as they are presently understood and refer to homologies outside of the
Apoidea. The membranous regions of the
wing running between the veins are termed
cells when bounded on all sides by veins.
The cells of the forewing and hind wing are
indicated in figure 7. I will mention only two
sets of cells that play a particularly important
role in the following descriptions and keys.
The marginal cell is the large membranous
region apicad the pterostigma and bordered
by R anteriorly and r-rs and Rs posteriorly.
This cell is variously modified, particularly
at its apex, which can be set along the leading edge of the wing, bent away from the
leading edge of the wing to different degrees,
become broadly truncated (e.g., Perditini,
some Electrapini), sometimes equipped with
a minute abscissa of Rs beyond the cell (the
cell is referred to as appendiculate when this
abscissa is present: e.g., fig. 7), or in the most
extreme case be entirely open (e.g., the stingless bees: figs. 111, 113). The submarginal
cells are a longitudinal series of two or, more
frequently, three cells immediately posterior
to the pterostigma and marginal cell; when
only two cells are present it is typically owing to a loss of 1rs-m, thereby effectively
uniting the second and third submarginal
cells. The shapes and and relative sizes of
the submarginal cells are useful for distinguishing species and genera of bees. In one
of the most extreme instances (i.e., the forewing of some Meliponini) the loss of the distal wing veins has left not only the marginal
cell open at its apex, but the submarginal
cells are completely lost (e.g., figs. 111, 113).
The posterior edge of the membrane of the
wings is typically incised at specific sites that
indicate the endpoints of particular folds in
31
the wing. These incisions and furrows demarcate specific regions of the insect wing
that appear as lobes along the posterior edge
of the wings. The basalmost lobe is the jugal
lobe (fig. 7), which defines the anterior border of that region of the wing termed the jugum. The absence, presence, and size of the
jugal lobe in the hind wing is a useful character for recognizing groups of bees. This
structure, however, can be easily misinterpreted even in living bees since the jugal
lobe frequently folds under the remainder of
the hind wing and therefore appears absent.
In fossil bees this is even more problematic
since the amber often makes it difficult to
discern the margins of such a folded lobe
when attempting to look through the membrane of the wing; this is further complicated
by the fact that amber sometimes has a
clearing effect on wing veins and membrane, making them appear more faint. The
next more apical lobe is the vannal lobe (fig.
7) and is defined by a fold running near to
the anal vein; the region between the jugal
fold and the vannal fold is the vannus. The
vannus is the posterior part of a region typically termed the clavus among other groups
of Hymenoptera and other insects; the associated structures are referred to in those instances as the claval lobe and claval furrow
(the latter of which typically runs just posterior to Cu). Thus, in other groups of Hymenoptera what is referred to in bees as the
vannal lobe is sometimes termed the claval
lobe. The remainder of the wing membrane
apical from the axillary region (bearing the
axillary sclerites) is the remigium. The lobes
of the hind wing are particularly useful for
classifying bees. The relative sizes of the jugal and vannal lobes as well as the presence
or absence of the former are useful phylogenetic and taxonomic characters. Because
they are rather uniform, those lobes of the
forewing are not typically referred to for
classificatory purposes, but the vannal lobe,
at the minimum, is present. The membrane
of the wings is typically covered by minute
setae, although in some bee groups (e.g., Xylocopa, Bombus, Electrobombus) the apical
portion of the wing beyond the veins is studded with minute, darkened swellings termed
alar papillae (labeled as such in fig. 6). The
32
NO. 259
Fig. 6. Diagrammatic wing with longitudinal and crossveins labeled as well as a single membrane
feature (i.e., alar papillae). Upper image is a forewing; lower image is a hind wing.
wing membrane is usually hyaline but can be

infuscated or patterned to varying degrees.
METASOMA
The metasoma consists of the true abdomen minus the first segment that is fused to
the thorax. Each of the metasomal terga and
sterna has a suite of morphological structures
associated with them, but are not discussed
further here since they are almost universally
unobserved for the known fossil bees. For a
more complete discussion of the exoskeletal
morphology of the metasoma refer to Michener (1944, 2000a), Snodgrass (1956), and
Eickwort (1969), among other authors. The
first metasomal tergum is constricted anteriorly to join the mesosoma and is therefore
typically composed of an anterior-facing surface as well as a dorsal-facing surface. In the
Lithurginae the first tergum is shortened such
that there is no dorsal-facing surface. Running transversely across the basal half or the
middle (or basal part of the visible portion
of terga and sterna) of each tergum and sternum at about the level of the spiracle is a
distinct line that is frequently carinate or minutely lamellate, referred to as the gradulus
(fig. 3). The gradulus divides the terga and
sterna into pregradular and postgradular regions.
2001
33
Fig. 7. Diagrammatic wing with membranous cells and gross anatomical structures labeled. Upper
image is a forewing; lower image is a hind wing.
The terminalia (i.e., sting apparatus, male

genitalia) of bees is not considered in the
present work simply owing to the fact that in
no Baltic amber bee specimen are these
structures visibly preserved (except the sting
apex in many individuals and with less frequency the worker gonostyli). The sting is
the modified ovipositor of female Aculeata
(fig. 3), while the female gonostyli, or sting
sheaths, are what are typically referred to as
the third valvulae (which themselves are the
apical accessory lobes or styli of the second
valvifer: see Snodgrass, 1935; Michener,
1944). For information on terminal struc-
tures, particularly those of the male, refer to

Snodgrass (1935, 1941, 1956), Michener
(1944, 1965, 2000a), or Eickwort (1969),
among other publications.
SYSTEMATIC PALEONTOLOGY
This section covers the descriptive morphology and taxonomy of all bees presently
known in Baltic amber. Numerous keys are
presented, all of which apply only to those
groups presently known from Baltic amber.
Therefore, in each case the reader should not
be alarmed at the exclusion of certain taxa
34
(e.g., the absence of Colletidae from the key

to families of bees, for this family is not currently known from the middle Eocene of Europe). Likewise, characters are employed that
may only apply to certain groups of a particular family if those particular groups are the
only ones presently known in Baltic amber.
For example, the strongly arcuate basal vein
is given as a character to separate Halictidae
in the key to families, because only the subfamily Halictinae is presently known in Baltic amber, even though the subfamilies Rophitinae and Nomiinae lack this feature. Similarly, within Apidae, only corbiculate apines
are presently known in Baltic amber and so
the key to subfamilies of Apidae uses the
presence of a corbicula to separate Apinae
from Xylocopinae, even though numerous
living tribes of Apinae also lack a corbicula.
Thus, future researchers discovering bees in
Baltic amber should be careful when using
these keys, since new specimens may represent higher rank groups (e.g., the aforementioned Colletidae) that would be new records
for Baltic amber and were unknown to me at
the time of this study. Naturally, I would be
interested in examining any newly discovered fossils for the preparation of future supplements to this monograph.
SUPERFAMILY APOIDEA LATREILLE
DIAGNOSIS: Aculeate Hymenoptera macropterous in both sexes; sexual dimorphism
slight. Antenna with 10 flagellomeres in females and 11 in males (except in a few, apomorphic instances). Posterior border of pronotal lateral surface extended just below level
of wing base to form a distinct pronotal lobe;
postcoxal process of pronotum greatly produced mesad and approaching counterpart
(processes well separated in most spheciforms, while being much closer, if not in contact, in bees); posterodorsal margin of pronotum broadly U-shaped. Prepectus transverse
and fused midventrally to mesopleurosternum. Metapostnotum (propodeal triangle)
expanded, fused to propodeum. Forewing
with well-developed venation, frequently
with 9 to 10 closed cells; hind wing frequently with jugal lobe. Metasomal S1 and S2 not
separated by a strong constriction. Ovipositor
concealed at rest and modified as sting.
NO. 259
TABLE 4
Familial Classification of Superfamily

Apoidea
COMMENTS: The superfamily Apoidea consists of two primary divisions: the paraphyletic assemblage of spheciform wasps (families Angarosphecidae, Ampulicidae, Crabronidae, Heterogynaidae, and Sphecidae),
and the monophyletic bees (Apiformes). The
Spheciformes is not treated further herein.
Table 4 summarizes the familial classification
of the superfamily.
It has been observed that the name Spheciformes is a hybrid name (composed of
Greek sphekos and Latin forma) and should
therefore be replaced by appropriately
formed names; namely Sphecomorpha (5
Spheciformes) and Melissomorpha (5 Apiformes). I have chosen to follow Brothers
(1975) for these informal groups rather than
employ those terms of wholly Greek origin.
Key to Divisions of Apoidea
1. Body setae simple; metabasitarsus not broader than remaining tarsal segments, frequently with strigil; female T7 not divided
into hemitergites . . . . . . Spheciformes
At least some setae, particularly those of legs
and surrounding propodeal spiracle,
branched or plumose; metabasitarsus flattened, wider than remaining tarsal segments, without strigil; female T7 divided
into hemitergites (bees) . . . . . Apiformes
2001
DIVISION APIFORMES
(THE BEES)
DIAGNOSIS: Some setae branched or plumose; subantennal sutures internally connected to fan-shaped sheet of tentorium; labrum thickened basally and attached at clypeal margin; proboscidial fossa opened onto
posterior surface of head capsule; cardo not
broadened apically, approximately parallelsided; suspensorium of paraglossa with
sparse setae or bristles; mesotibial comb present; dorsoventral length of mesocoxa about
equal to distance from summit of mesocoxa
to hind wing base; metabasitarsus broader
than following tarsal segments, without concavity or strigil; cu-a of hind wing shorter
than second abscissa of M1Cu; T7 of female
divided into two hemitergites.
COMMENTS: The bees are a monophyletic
group among the Apoidea. Although I have
not included it in the list above, an additional
character diagnosing the bees includes the
vegetarian diet of the larvae. Aside from a
few, apomorphic exceptions bee larvae consume pollen, nectar, and plant oils.
McKenna and Bell (1997: 513) presented
a series of ranks and suffixes for familygroup names (adapted from Bour and Dubois, 1984) that could be applied to provide
more hierarchy to the classification of Apoidea. In particular the rank of epifamily (intermediate between superfamily and family
and with suffix -oidae) could be applied to
refer to the bees as a whole. This would result in what I have called division Apiformes
being replaced by epifamily Apoidae. Since
this name would be quite similar to the superfamilial name (i.e., Apoidea), I feel that it
would merely lead to confusion with the
more inclusive group consisting of the spheciforms and bees. I have, therefore, used informal names without official ranks for these
intercalary entities.
SHORT-TONGUED VERSUS LONG-TONGUED
BEES
Bees have been segregated into major
groups; or the short- and long-tongued bees,
a division originally recognized by Kirby
(1802) and used ever since. Although these
vernacular names are useful, they are a bit of
a misnomer as these characters are not uni-
35
versal (also, one is a paraphyletic group).

The long-tongued bee families (Megachilidae and Apidae) have the first and second
segments of the labial palpus sheath-like
(i.e., elongate and flattened), while shorttongued families (Colletidae, Halictidae, Andrenidae, Paleomelittidae, and Melittidae)
have these same segments cylindrical and
more or less similar to the following segments. The long-tongued families have been
repeatedly demonstrated to be a monophyletic group; the short-tongued families are
not. The short-tongued family Melittidae, in
particular, has been demonstrated to be more
closely related to the Megachilidae 1 Apidae
clade owing to the shortened jugal lobe, fully
exposed mesocoxa, and V-shaped submentum, among other characters. Michener and
Greenberg (1980), for the purposes of a discussion on the higher-level phylogeny of
bees, subdivided these groups so as to reflect
the special position of Melittidae among
short-tongued bees. Most notably these authors distinguished basic short-tongued
bees apart from short-tongued bees (as
well as other phenetic groupings not employed here). In the following text I regularly
refer to these entities for comparative purposes. It therefore seems worthwhile to briefly provide a definition of these terms.
LONG-TONGUED BEES (5 Megachilidae
and Apidae): families with elongate and flattened labial palp segments one and two; jugal
lobe shortened; mesocoxa fully exposed;
submentum V-shaped; preepisternal groove
absent at least below level of scrobe.
SHORT-TONGUED BEES (5 Colletidae, Halictidae, Andrenidae, Paleomelittidae, and
Melittidae): families with cylindrical labial
palp segments one and two. Among this
group the Melittidae stand out by having the
short jugal lobe, V-shaped submentum, absence of the preepisternal groove; and fully
exposed mesocoxa typical of the longtongued bees. Paleomelittidae is also noteworthy in that it has the characters of the
basic short-tongued bees (defined below) except this family has a fully exposed mesocoxa and lacks a preepisternal suture (like
Melittidae and long-tongued bees) but retains
the elongate jugal lobe.
B ASIC S HORT -T ONGUED B EES (sensu
Michener and Greenberg, 1980: 5 Colleti-
36
dae, Halictidae, and Andrenidae): families

with cylindrical labial palp segments one and
two; jugal lobe elongate; mesocoxa hemicryptic; submentum plate-like; preepisternal
groove complete.
Key to Families of Bees in Baltic Amber
(Based on Females)
1. Scopa strongly developed on metasomal sterna; subantennal sutures directed to outer
margins of antennal sockets (figs. 26, 32,
40); long-tongued bees . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . Megachilidae Latreille
Scopa not developed on metasomal sterna,
variously developed on hind legs; subantennal sutures attaching to antennal sockets
on inner or lower margins (e.g., figs. 12,
17, 21, 49, 59, 64); short- or long-tongued
bees . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2
2. Preepisternal groove absent below level of
scrobe; mesocoxa fully exposed; basal vein
variously shaped; short- or long-tongued
bees . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3
Preepisternal groove present and complete;
mesocoxa hemicryptic; basal vein strongly
arcuate (fig. 13); basic short-tongued bees
. . . . . . . . . . . . . . . . . Halictidae Thomson
3. Jugal lobe of hind wing short (e.g., figs. 63,
76); flabellum absent or present; preepisternal groove either present or absent;
short- or long-tongued bees . . . . . . . . . . 4
Jugal lobe of hind wing elongate; flabellum
absent; preepisternal groove absent below
level of scrobe; short-tongued bees . . . .
. . . . . . . . . . . . . . . Paleomelittidae n. fam.
4. Short-tongued bees (i.e., labial palp segments
one and two cylindrical and similar in
length to distal segments); flabellum absent; galea not elongate or if elongate then
prepalpally, galeal comb present . . . . . .
. . . . . . . . . . . . . . . . . . Melittidae Schenck
Long-tongued bees (i.e., labial palp segments
one and two elongate and flattened); flabellum present; galea elongate postpalpally,
galeal comb absent . . . . Apidae Latreille
SHORT-TONGUED BEES
Family HALICTIDAE Thomson
DIAGNOSIS: The halictids comprise three
homogeneous subfamilies of basic shorttongued bees (i.e., cylindrical labial palp segments, elongate jugal lobe, hemicryptic mesocoxae, &c.; see above). Halictidae is most
notable for the loss of the mentum and submentum.
NO. 259
DESCRIPTION: Subantennal suture meeting

inner, lower margin of antennal socket, infrequently lower margin; two subantennal sutures in some Rophitinae otherwise only one
subantennal suture present (only one in the
known fossils). Facial foveae typically absent. Galea elongate prepalpally. Mentum
and submentum absent; segments one and
two of labial palpi similar to distal segments,
cylindrical, rarely elongate; glossa acute, typically short, without flabellum. Preepisternal
groove complete (except in Nomiinae groove
weak or absent below scrobe). Mesocoxa
with exposed area shorter than distance from
summit to metathoracic wing base; metabasibitial plate frequently present (except in
parasitic forms); scopa developed on hind leg
in nonparasitic females. Jugal lobe of hind
wing much more than half as long as vannal
lobe. Pygidial plate present in females. Volsellae present.
COMMENTS: The family presently contains
three subfamilies: Rophitinae, Nomiinae, and
Halictinae. The monophyly of the rophitines
remains to be tested and the subfamily is
possibly paraphyletic. The Nomioidinae sensu Alexander and Michener (1995) and
Michener (2000a) is considered here as the
basal tribe of Halictinae (see also Engel,
2000a; Pesenko, 1999; Pesenko et al., 2000).
Of these subfamilies, only the Halictinae is
presently known from the fossil record.
Subfamily HALICTINAE Thomson
DIAGNOSIS: This subfamily is most closely
related to the Nomiinae, both sharing the development of the scopa on the metafemur
(rather than the metatibia in Rophitinae), the
distal process of the labrum (absent in Rophitinae), and strong distal process with a
penicillus on the metabasitarsus (either completely absent or under-developed in the basalmost subfamily, Rophitinae). The halictines
can be separated from the nomiines most notably by the third submarginal cell being
shorter than the first submarginal cell, by the
prepygidial fimbria being typically divided
by a longitudinal median line or area, and by
the complete and strong preepisternal
groove.
DESCRIPTION: Labrum of female with distal
process. Clypeus longer than labrum. Pree-
2001
pisternal groove strongly impressed below

level of scrobe. Basal vein strongly arcuate;
first submarginal cell longer than third submarginal cell. Scopa developed on metacoxa,
metatrochanter, metafemur, and to varying
degrees on metatibia in nonparasitic females;
metabasitarusus with distal process and penicillus. Prepygidial fimbria of female distinctly divided by pseudopygidial area.
COMMENTS: Traditionally this subfamily
includes three tribes: Augochlorini, Halictini,
and Nomioidini; however, see Engel (2000a)
for an alternative classification dividing the
paraphyletic Halictini into at least five tribes.
The classification of Engel (2000a) is adopted here and summarized with slight modification in table 5.
The halictines are the only subfamily of
Halictidae recorded from the fossil record.
The tribes Caenohalictini (Eickwortapis) and
Augochlorini (Augochlora, Neocorynura,
and Oligochlora) are present in Miocene amber from the Dominican Republic (table 1).
Numerous compression fossils of the Halictini are recorded from the Eocene-Oligocene
boundary of Florissant, Colorado (Engel,
1996, unpubl. data), as well as some specimens in scattered, younger deposits throughout the world (e.g., Bachmayer et al., 1971;
Engel, 1996; Arillo et al., 1996) but the tribal
placement of these fossils is slightly uncertain (all are likely to be Halictini s.s.). Bachofen-Echt (1949) reported a Halictus from
Baltic amber, but I have been unable to locate the material from which he made his
determination. Oddly, Bachofen-Echt (1949)
referred his fossil Halictus to the subfamily Apinae (a gross misplacement even allowing for the taxonomic conventions of his
time; even the most radical of classifications
never classified Halictus with the apines!).
Below I describe the only definitive halictine
in Baltic amber.
Tribe HALICTINI Thomson
Sphecodidae Schenck, 1869: 316. Type genus:
Sphecodes Latreille, 1804. Suppressed in favor
of Halictina Thomson (Michener, 1991; ICZN,
1993).
Halictina Thomson, 1869: 8. Type genus: Halictus
Latreille, 1804.
Thrinchostomini Sakagami, 1974: 258. Type genus: Thrinchostoma Saussure, 1890.
37
TABLE 5
Hierarchical Suprageneric Classification of

Halictidae
DIAGNOSIS: This tribe is most similar to the

Gastrohalictini (5 Lasioglossum group) but
differs most notably by the strong distal wing
venation.
DESCRIPTION: Anterior tentorial pits set in
epistomal sulcus (not in clypeus and connected to epistomal sulcus by a distinct sulcus as in Nomioidini); distal wing venation
strong; pseudopygidial area of T5 in females
lacking a median cleft; males with a pygidial
plate present on the apex of T7; males lacking a spiculum; first and second metatarsal
segments of male not fused.
COMMENTS: I have used here the narrowed
concept of Halictini as presented by Engel
(2000a: 67) excluding the weakened wing
vein groups, the agapostemonine genera, the
caenohalictine genera, and the augochlorine
genera. Such a narrowing of the tribe to exclude the latter groups makes the Halictini
putatively monophyletic. Table 5 outlines the
classification of Halictidae adopted herein.
38
Fig. 8.
1 mm.
NO. 259
Right lateral habitus of holotype female of Electrolictus antiquus, new species. Scale bar 5
Electrolictus, new genus

TYPE SPECIES: Electrolictus antiquus Engel, new species.
DIAGNOSIS: This group appears in some respects to be similar to the principally African
genus Patellapis (sensu Michener, 1978)
combining features of the subgenera Patellapis s.s. and Chaetalictus. From Chaetalictus the fossil differs by the strongly bordered
metabasitibial plate that is not obsolescent
anteriorly, while from Patellapis s.s. the fossil differs by the absence of metasomal setal
bands. From the whole genus Patellapis,
Electrolictus can be distinguished by the absence of tomentum, absence of metasomal
banding, reception of both m-cu crossveins
by the third submarginal cell, and sparse and
simple mesosomal pubescence. The reception of both 1m-cu and 2m-cu in the third
submarginal cell is reminiscent of Thrinchostoma but Electrolictus differs from this
genus by the absence of tergal setal bands
(in Thrinchostoma these are formed by laterally directed setae along the apical margins), strong preepisternal groove, and extremely short malar space. Furthermore, most
Thrinchostoma (subgenera Thrinchostoma
s.s. and Diagonozus) have a distinctive,

dense setal patch near 1rs-m in the forewing
which Electrolictus lacks.
DESCRIPTION: Mandible with strong subapical tooth, mandible almost bidentate (fig.
11). Malar space much shorter than basal
mandibular width, base of mandible nearly
touching lower margin of compound eye (fig.
10). Basal area of labrum with a low apical,
suborbicular elevation; distal process narrowly triangular and laterally serrate with
dense, long, lateral fimbria (fig. 11); distal
keel of dorsal process strong without basal
expansion, slightly wider at apex. Hypostomal ridge carinate (fig. 11), anterior angle
rounded, ridges parallel (not diverging anteriorly). Proboscis not greatly narrowed
(length less than six times width); glossa relatively short; maxillary palpal segments
about equal in length, total length of palpus
about 90% of prementum length. Clypeal
truncation without short, lateral processes
(fig. 12), lower half of clypeus below lower
tangent of compound eyes (fig. 12); clypeus
gently convex, not protuberant in lateral view
(fig. 10). Epistomal sulcus forming a greatly
obtuse angle, nearly linear (fig. 12). Supra-
2001
clypeal area not protuberant in lateral view.

F1 about as long F2 (figs. 9, 10). Gena slightly narrower than compound eye in lateral
view. Compound eyes without pubescence;
inner margins slightly emarginate above level of antennal sockets; eyes slightly converging below. Ocelli not enlarged; interocellar
furrow absent (fig. 9). Preoccipital area
rounded. Pronotal dorsal ridge carinate; lateral ridge rounded; dorsolateral angle slightly
obtuse (fig. 8). Anterior border of mesoscutum broadly rounded (fig. 8), anterior extremity rising well above pronotum but not
overhanging, anterior vertical surface curving gently onto dorsal surface; tegula oval;
metanotum not tomentose. pree pisternal
groove present and strong below level of
scrobal groove. Probasitibial comb absent;
malus short, slightly shorter than velum, with
long, thin pectinate teeth; claws with short
inner tooth (fig. 16); arolium present; metabasitibial plate with strong borders, narrowly
triangular (fig. 14); inner hind tibial spur pectinate, five long teeth (not including apex)
(fig. 15); metabasitarsus with distal process
and penicillus. Basal vein strongly arcuate
(fig. 13); marginal cell apex feebly truncate
and appendiculate; three submarginal cells
(fig. 13); both m-cu crossveins entering third
submarginal cell (fig. 13); first submarginal
cell about as long as combined lengths of
second and third submarginal cells; distal
wing veins strong. Basal area of propodeum
with strong striae radiating to apical margin
(fig. 8), not strongly declivitous (nearly horizontal); ridge between basal and posterior
surfaces rounded; no carinae on lateral margins of posterior surface. Metasoma with tomentum; T1 broader than long; terga without
setal bands (basal or apical); tergal graduli
slightly lamellate; metasoma without integumental banding; pseudopygidial area not
cleft.
ETYMOLOGY: The new genus-group name
is a combination of electrum (L. meaning
amber) and -lictus (root of the type genus
of the Halictini, Halictus). The name is masculine.
Electrolictus antiquus, new species
Figures 816
Plate 1a,b
DIAGNOSIS: As for the genus.

DESCRIPTION: Female. Total body length
39
8.17 mm; forewing length 5.60 mm. Head

slightly longer than wide (length 2.08 mm,
width 1.68 mm). Face relatively flat in lateral
view. Upper interorbital distance 1.28 mm;
lower interorbital distance 1.16 mm. Vertex
unmodified, relatively short (ca. 1 OD in
length: measured from posterior margin of
lateral ocellus to preoccipital area. Interocellar distance 0.36 mm; ocellocular distance
0.28 mm; median ocellus to lateral ocellus
0.12 mm. Median and parapsidal lines relatively strong. Basal vein distad cu-a by about
two times vein width; 1m-cu distad 1rs-m by
about vein width; 2rs-m distad 2m-cu by
about four times vein width, 2rs-m arched
(not straight); first submarginal cell about as
long as combined lengths of second and third
submarginal cells; second submarginal cell
slightly narrowed anteriorly; anterior border
of second submarginal cell equal to that of
third submarginal cell; posterior border of
third submarginal cell approximately 1.5
times longer than anterior border.
Clypeus with coarse, faint punctures,
punctures separated by a puncture width or
less and integument between faintly imbricate on apical third, remainder of clypeus
with smaller, well-defined punctures separated by a puncture width or less and integument between smooth. Supraclypeal area
sculptured as on upper two-thirds of clypeus.
Face with similar punctures as those on supraclypeal area except separated by less than
a puncture width, in most places punctures
nearly contiguous; integument between
punctures smooth; punctures slightly smaller
near ocelli. Vertex and gena with minute
slightly more, integument between smooth.
Postgena faintly imbricate with a few widely
scattered coarse punctures. Pronotum
smooth; mesoscutum with well-developed
punctures separated by 12 times a puncture
width except around median line punctures
slightly more sparse, separated by 23 times
a puncture width, integument between
smooth; tegula smooth except inner margin
bordering mesoscutum with minute punctures separated by less than a puncture width;
scutellum sculptured as on mesoscutum; metanotum rugulose. Preepisternum with coarse
punctures, punctures nearly contiguous, integument between smooth when evident;
40
NO. 259
Figs. 912. Head of holotype female of Electrolictus antiquus, new species. 9. Dorsofrontal view.
10. Left lateral view. 11. Ventral view. 12. Frontal view. Scale bars 5 1 mm (a 5 figs. 9, 10, 12; b 5
fig. 11).
mesepisternum sculptured as on preepisternum except hypoepimeral area with smaller

less; metepisternum sculptured as on hypoepimeral area although punctures slightly
transversely elongate. Basal area of propodeum with strong striae radiating from basal
margin, striae reaching to apical margin of
dorsal surface, integument between smooth
and shining; lateral surface minutely punc-
tured and faintly imbricate along posterior

half. Terga and sterna imbricate.
Head and mesosoma dark brown to black
with weak metallic highlights; tegula dark
brown. Legs dark brown. Wing membrane
hyaline; veins strong and dark brown. Metasoma uniformly dark brown, without integumental banding.
Mandible with moderate length, simple setae scattered along lower margin and on out-
2001
41
Fig. 13. Forewing of holotype female of Electrolictus antiquus, new species. Scale bar 5 1
mm.
er surface. Clypeus with apical fimbria of

simple setae; face, supraclypeal area, and
surface of clypeus with short to moderate
length, subappressed to suberect, scattered
setae each with a few minute branches; setae
more sparse on vertex. Gena with subappressed short, scattered setae each with a few minute branches. Postgena with widely scattered, simple setae of moderate length. Mesoscutum with widely scattered, erect, moderate-length setae, each with minute
branches; tegula with minute, appressed,
simple setae along inner half; scutellum with
scattered, long setae, each with numerous
minute branches; metanotum with setae as on
scutellum. Mesepisternum with scattered,
moderate-length, suberect setae, each with
several minute branches; metepisternum with
sparse, minute, simple, erect setae. Scopa of
metafemur with long, plumose setae; similar
setae on metatibia although more dense. Lateral and posterior surface of propodeum with
sparse, long, simple setae. Terga without tomentum, instead, with sparse, simple, subappressed, short setae; sterna with similar pubescence except setae slightly longer and restricted to apical halves.
MATERIAL: One specimen. Holotype. Female, MB.I.1952 (ZMHB): labeled Holotype, Electrolictus antiquus Engel // Museum fur Naturkunde Berlin, Palaontologisches Museum, Slg. Kuhl 23 (3/12), Inv.
Nr. MB.I.1952 // Halictidae: Halictinae,
det. M. S. Engel, 1999.
ETYMOLOGY: The specific epithet is the
Latin word antiquus meaning ancient and
is a reference to this being the oldest record
of an amber halictid.
PALEOMELITTIDAE, New Family
TYPE GENUS: Paleomelitta Engel, new genus.
Figs. 1416. Leg structures of holotype female of Electrolictus antiquus, new species. 14.
Metabasitibial plate. 15. Inner metatibial spur. 16.
Claw, arolium, distitarsus, and apical two segments of mediotarsus. Scale bars 5 0.5 mm (a 5
fig. 14; b 5 figs. 15, 16).
DIAGNOSIS: This family is a typical shorttongued bee lineage with the cylindrical labial palpus segments and elongate jugal lobe;
the latter, plesiomorphic character distinguishing the group from the Melittidae. Paleomelittidae also differs from the melittids
by the entirely complete scrobal groove (present only posterior to episternal scrobe and
reaching to anterior suture of metepisternum
in Melittidae). Unlike the basic shorttongued families, however, Paleomelittidae
has a fully exposed mesocoxa.
DESCRIPTION: Single subantennal suture
(i.e., subantennal areas absent) meeting inner,
lower margin of antennal socket. Facial foveae absent. Segments one and two of labial
palpi similar to distal segments, cylindrical,
distal segments in line with basal segments;
glossa acute, short, without flabellum; stipital
concavity and comb absent; galeal comb absent. Preepisternal groove incomplete, absent
below level of scrobe; scrobal groove present
and entire (i.e., present both in front of as
well as posterior to episternal scrobe). Mesocoxa fully exposed, exposed area slightly
longer than distance from summit to metathoracic wing base; metabasibitial plate present; scopa developed on metafemur (more
weakly than on following leg segments), metatibia, and metabasitarsus, scopa strongest
42
NO. 259
on metatibia. Jugal lobe of hind wing much

more than half as long as vannal lobe.
COMMENTS: Paleomelittidae occupies an
interesting position intermediate between the
basic short-tongued families (Colletidae,
Halictidae, and Andrenidae) and the Melittidae 1 long-tongued families complex.
The family has the fully exposed mesocoxa
typical of melittids and long-tongued bees
and is therefore closer to this group than to
other basic short-tongued families. It could
be retained as the basalmost subfamily of
Melittidae but this would render the melittids
paraphyletic and I have therefore accorded
the group family rank.
Paleomelitta, new genus
TYPE SPECIES : Paleomelitta nigripennis
Engel, new species.
DIAGNOSIS: As for the family (see above).
Paleomelitta has the general habitus of Dasypoda species and the forewing venation of
the two genera is somewhat similar in many
respects (except in dasypodaines 1m-cu is
not strongly apicad the second abscissa of
Rs). Paleomelitta can be separated from Dasypoda by the above familial characters as
well as those presented below in the generic
description (most notably the absence of a
metabasitibial plate in Dasypoda is significantly different from Paleomelitta where this
structure is large and well developed).
DESCRIPTION: As for the family with the
following additions: Labrum with two short,
medial tubercles on basal surface (fig. 17),
about 2.5 times wider than long, without distal process. Mandible broad at base and tapering rapidly to apex, strongly bent inward
near midpoint, with strong subapical tooth
(fig. 17). Malar space short, much shorter
than basal mandibular width. Clypeus slightly concave medially on central disc, apical
margin with distinct medial tubercle laterally
bordered by tufts of elongate, erect setae
(tufts composed of three setae each: fig. 17);
epistomal sulcus forming an obtuse angle;
apical quarter of clypeus set below lower tangent of compound eyes. Inner margins of
compound eyes approximately parallel. F1
longer than F2 (fig. 17); F3 slightly longer
than F2. Vertex short, ca. 1.5 OD in length
(fig. 18). Posterior margin of scutellum with
Fig. 17. Frontal view of head of holotype female of Paleomelitta nigripennis, new species.
Scale bar 5 1 mm.
fringe of elongate, simple setae (fig. 18); scutellum rising well above metanotum then
abruptly curving behind posterior fringe to
meet metanotal basal border, not projecting
over metanotum. Metabasitibial plate present, large, and strongly bordered on all edges; scopal setae simple; claws with short inner tooth (fig. 19); arolium present and large
(fig. 19). Marginal cell apex set away from
wing margin by approximately three-quarters
of pterostigma width, feebly appendiculate
(fig. 20). Margin of pterostigma inside of
marginal cell slightly convex; pterostigma
longer than wide; r-rs arising in apical half
of pterostigma. Two submarginal cells; second abscissa of Rs approximately orthogonal
to M, strongly basad of 1m-cu (fig. 20).
is a combination of palaios (Greek, meaning
ancient) and melitta (Greek, meaning
bee). The name is feminine.
Paleomelitta nigripennis, new species
Figures 1720
Plates 1c,d, 2a

2001
43
Fig. 18. Dorsal view of mesosoma, posterior border of prosoma, and anterior border of metasoma
of holotype female of Paleomelitta nigripennis, new species. NOTE: The depicted incision in the hind
wing is the jugal fold demarcating the apical margin of the jugal lobe; the vannal fold is positioned just
under the posterior edge of the forewing. Scale bar 5 1 mm.
wider than long (length 3.25 mm, width 4.03

mm). Compound eye width approximately
twice that of gena. Interocellar distance 0.60
mm; ocellocular distance 0.50 mm; median
to lateral ocellus 0.15 mm. Median and parapsidal lines strongly impressed (fig. 18). Intertegular distance 3.05 mm; mesoscutum
length 1.05 mm; scutellum 2.5 times longer
than metanotum; basal area of propodeum
about as long as metanotum. Basal vein basad cu-a by vein width; second abscissa of
Rs basad 1m-cu by four times vein width,
straight, approximately orthogonal to M (fig.
20); 2rs-m distad 2m-cu by two times vein
width; first submarginal cell subequal to second submarginal cell; posterior border of
second submarginal cell approximately 1.25
times length of anterior border.
Labrum with small punctures separated by
a puncture width or less, integument between
smooth. Clypeus with small punctures separated by 12 times a puncture width on apical half, basal half punctures smaller, faint
and sparse, integument between smooth. Su-
praclypeal area, face, vertex, and gena sculptured as on basal half of clypeus. Pronotum
smooth and impunctate. Mesoscutum with
small punctures separated by 12 times a
puncture width except around median line
separated by a puncture width or less, integument between smooth. Scutellum sculptured
as on mesoscutum except punctures separated by 23 times a puncture width. Tegula
with minute punctures separated by less than
a puncture width on inner half, outer half impunctate, integument between finely imbricate. Metanotum imbricate. Pleura with
sparse, minute punctures, integument between smooth. Propodeal lateral and posterior surfaces finely imbricate; basal area of
propodeum strongly imbricate. Metasomal
terga with minute punctures separated by a
puncture width, integument between finely
imbricate; sterna imbricate.
Coloration dark brown. Wing membranes
exceedingly infumated, dark brown; veins
black.
Pubescence fuscous. Labrum with scat-
44
NO. 259
Fig. 20. Forewing of holotype female of Paleomelitta nigripennis, new species. Scale bar 5
1 mm.
Fig. 19. Distitarsus, claw, and arolium of holotype female of Paleomelitta nigripennis, new
species. Upper image is an outer view; lower image is an inner view. Scale bar 5 0.25 mm.
tered, short (ca. 1 OD or less), erect, simple

setae. Clypeus and supraclypeal area with
scattered, short, suberect, simple setae; clypeus with two submedial patches of three (2
OD in length) simple setae (fig. 17). Face,
vertex, and gena with scattered, simple, erect
to suberect, short (1 OD in length) setae; setae of gena progressively longer toward postgena (reaching 3 OD in length). Pronotum
with sparse, minute, suberect, simple setae
except surface at dorsolateral angle and pronotal lobe with dense, plumose setae (1 OD).
Mesoscutum with scattered, minute, erect,
simple setae, setae sparse on central disc but
more noticeable along borders. Scutellar pubescence as described for mesoscutum except posteriorly fringed with dense, plumose
setae; axilla with more elongate (1.52 OD),
simple, erect setae. Metanotum with scattered, short, suberect to erect, simple setae.
Pleura with scattered, simple, erect setae
(1.52 OD) except hypoepimeral area without setae. Lateral and posterior surfaces of
propodeum with scattered, short, simple se-
tae; basal area of propodeum without setae.

Metafemur without scopal setae; metatibia
and metabasitarsus with scopae composed of
dense, elongate, simple setae, setae longest
on inner surfaces (34.5 OD). Anterior-facing surface of T1 with widely spaced, erect
setae (2 OD), each with a few minute branches; remaining terga and apical margin of T1
with scattered, minute, simple, appressed setae; sterna with scattered, simple, erect, postgradular setae.
MATERIAL: Two specimens. Holotype. Female, B-JH 101 (AMNH) labeled: Baltic
amber: Eocene, Kaliningrad, Yantarny //
Holotype, Paleomelitta nigripennis Engel.
The block of amber containing the two bees
is an impressive size (pl. 2a) and measures
approximately 13.5 cm in length, 5.5 cm in
width, and 3.75 cm in height.
Paratype. Female, in same piece of amber
as holotype (see above).
ETYMOLOGY: The specific epithet is a combination of the Latin words nigra (meaning
black) and penna (meaning wing) and
is a reference to the darkened membrane of
the forewings.
Family MELITTIDAE Schenck
Melittidae Schenck, 1860: 136. Type genus: Melitta Kirby, 1802.
DIAGNOSIS: This family consists of shorttongued bees with the fully exposed mesocoxa, V-shaped submentum, and short jugal
lobe typical of the long-tongued bees while
retaining the plesiomorphic short-tongued
labial structure, absence of a stipital comb
and concavity, and absence of a flabellum.
Melittids also lack a preepisternal groove and
2001
the scopa is limited to the metatibia and metabasitarsus.

D ESCRIPTION : Labrum frequently much
wider than long, apical margin fringed with
bristles. Single subantennal suture; suture
meets lower margin of antennal socket frequently at midpoint, although sometimes
slightly laterad midpoint (either outer or inner side of lower margin), never meeting outer margin of antennal socket. Facial foveae
absent. Lower lateral margin of clypeus not
bent posteriorly on either side of labrum. Flabellum absent; glossa acute; glossal rod absent; paraglossa frequently reduced, sometimes absent; labial palpus with first two segments cylindrical, not sheath-like, third and
fourth segments in line with preceding segments, not projecting laterally. Mentum elongate, tapering basally, curved over submentum to its articulation; apex of mentum articulated with prementum via fragmentum;
submentum strongly V-shaped and sclerotized, submental arms articulating with cardines slightly above cardo-stipital articulations. Galeal blade equal or shorter than stipes, frequently longer than prepalpal section
of galea; galeal comb present; stipital comb
and concavity absent; basistipital process
elongate. Preepisternal groove absent. Mesocoxa entirely exposed. Metabasitibial plate
frequently distinct. Scopa restricted to metatibia and metabasitarsus. Jugal lobe of hind
wing one quarter to one half as long as vannal lobe. Metapostnotum without pubescence. Pygidial plate and fimbria of female
present; pygidial plate of male frequently absent. Volsella present; penis valves do not
form a distinct spatha.
C OMMENTS : The subfamily Melittinae
(sensu Michener, 1981b) contains three distinct groups that are not closely affiliated;
each has numerous characters separating
them from one another as much as the remaining subfamilies of the family. Herein I
have separated Macropis from the subfamily
Melittinae and resurrected the subfamily Macropidinae; I have here added a second genus
to this subfamily (described below). Likewise, the African genera Rediviva and Redivivoides, while retained in Melittinae, are restricted to their own tribe. The Melittinae is
therefore reduced to two groups: tribe Melittini with Melitta s.l. and Dolichochile, and
45
TABLE 6
Hierarchical Supraspecific Classification of

Melittidae
(Currently recognized genera included
but subgenera excluded.)
tribe Redivivini (new tribe; type genus: Rediviva Friese) with Rediviva and Redivivoides. The Redivivini can be separated from
Melittini by the reduced, shining, and smooth
propodeal triangle and males with S7 bifid
(Michener, 1981b). Table 6 summarizes a
new higher classification of Melittidae.
The separation of Macropis from the Melittinae implies that the oil-collecting structures of the Macropidini and the Redivivini
arose independently. This seems to be a valid
conclusion since macropidines appear to
share features with basal dasypodaines (e.g.,
Promelitta) particularly in the presence of
only two submarginal cells and, in fact, the
structure of the second abscissa Rs in basal
46
dasypodaines is difficult to distinguish from

the state seen in Macropidinae.
Subfamily MACROPIDINAE Robertson
Macropididae Robertson, 1904: 42. Type genus:
Macropis Panzer, 1809.
DIAGNOSIS: This subfamily resembles the

Dasypodainae by the presence of only two
submarginal cells in the forewing (i.e., loss
of 1rs-m). In dasypodaines, however, the second abscissa of Rs is not widely separated
from 1m-cu and is essentially orthogonal to
M. From Melittinae, in the restricted sense
that I have used it here, macropidines differ
by the reduced number of submarginal cells,
the presence of yellow maculations on the
face of males, and the presence of a pygidial
plate in males.
DESCRIPTION: Mandible with strong subapical tooth. Two submarginal cells, cells
subequal in length; second abscissa of Rs
slightly oblique, widely separated from 1mcu. Pygidial and prepygidial fimbriae strong.
Males with yellow maculations on the clypeus; males with pygidial plate present.
COMMENTS: The subfamily as presently defined contains two genera (Macropis and
Eomacropis), which I have segregated into
two monobasic tribes. This is owing to the
numerous, significant differences between
the two genera, each with several apomorphic traits supporting their respective monophyly. They are sufficiently different to warrant
placement in different tribes.
EOMACROPIDINI, New Tribe
TYPE GENUS: Eomacropis Engel, new genus.
DIAGNOSIS: This tribe differs from Macropidini most notably by the complete absence
of the suite of oil-collecting adaptations seen
in Macropis (e.g., velvety setae on thickened
tarsi). Eomacropidini has an elongate metabasitarsus and vein r arises distad the middle
of the pterostigma. The apomorphic presence
of a strongly convex and medially carinate
metanotum and V-shaped propodeum is immediately diagnostic.
D ESCRIPTION : Labrum without median
transverse ridge, with subapical elevation
(fig. 21). Anterior border of scutellum gently
NO. 259
biconcave (straight in Macropidini), posterior

border rising well above metanotum (fig. 22)
(not gently curving over to meet metanotum
as in Macropidini). Metanotum strongly convex, with median longitudinal carina (fig.
22). Protarsi and mesotarsi not thickened,
without dense, velvety setae; metatibia and
metabasitarsus thin and elongate (not broad
as in Macropidini), metabasitarsus at least
three times longer than wide (fig. 24); metatibia without oil-collecting setae; metabasitibial plate strong, broadly rounded (fig. 25);
metabasitarsus with setae present on distal
process; metabasitarsal setae unmodified.
Vein r-rs arising distad middle of pterostigma
(fig. 23). Propodeum strongly V-shaped, dorsal-facing surface not transverse.
Eomacropis, new genus
TYPE SPECIES: Eomacropis glaesaria Engel, new species.
DIAGNOSIS: As for the tribe (see above).
DESCRIPTION: Galea with long setae at
apex; maxillary palpus six-segmented (basal
segment not visible in figure 21 but is visible
from a lateral view of the head). Width of
labrum three times greater than length; surface impunctate; short, median, distal process
with long setae along margins. Malar space
linear, length much less than basal mandibular width, base of mandible nearly touching
lower margin of compound eye. Mandible
with strong subapical tooth. Epistomal sulcus
forming approximately orthogonal angle (fig.
21). F1 longer than F2 (fig. 21). Compound
eye weakly emarginate above level of antennal sockets (fig. 21); compound eyes without
setae. Gena slightly broader than compound
eye in lateral view. Preoccipital area sharply
angled but not carinate. Scrobal groove present. Claw with inner tooth, arolium present
(fig. 24); metatibial and metabasitarsal scopa
composed of densely plumose setae; setae of
apical truncation of metabasitarsus not parted
to expose narrow strip of integument. Basal
vein straight, basad cu-a, basal vein twice as
long as first abscissa of Rs (fig. 23); prestigma shorter than margin of pterostigma inside
of first submarginal cell, prestigma about as
wide as pterostigma; pterostigma inside of
marginal cell slightly convex (fig. 23). Basal
area of propodeum impunctate, granular and
2001
47
Figs. 2122. Holotype female of Eomacropis glaesaria, new species. 21. Frontal view of head. 22.
Right dorsolateral oblique view of mesosoma, posterior border of prosoma, and anterior border of
metasoma. NOTE: Staphyliniform beetle larva on mesoscutum of the holotype. Scale bars 5 1 mm (a
5 fig. 22; b 5 fig. 21).
without pubescence (plesiomorphically similar in this respect to Macropis s.s.).

is a combination of eo- (a reference to the
epoch name, Eocene) and Macropis, type genus of the Macropidinae. The name is feminine.
Eomacropis glaesaria, new species
Figures 2125
Plate 1e,f
DIAGNOSIS: As for the genus (see above).

slightly wider than long (length 2.65 mm,
width 2.85 mm). Distal half of clypeus extending below lower tangent of compound
eyes (fig. 21). Frontal line carinate from an-
tennal sockets to approximately 1 OD from

median ocellus. Upper interorbital distance
1.65 mm; lower interorbital distance 1.55
mm. Interocellar distance 0.35 mm; ocellocular distance 0.4 mm; median to lateral
ocellus 0.15 mm. Scutellum slightly less than
twice length of metanotum; metanotum
slightly longer than basal area of propodeum.
Malus serrate; distal section of protibial spur
beyond velum slightly shorter than section of
spur bordering velum. Intertegular distance
1.9 mm. Second abscissa of Rs strongly basad 1m-cu (fig. 23); 2rs-m distad 2m-cu by
three times vein width; first submarginal cell
slightly shorter than second submarginal cell;
eight distal hamuli, arranged in a single
evenly spaced series.
Labrum with minute, scattered punctures,
48
NO. 259
2001
integument between smooth. Clypeus with

sparse, faint, coarse punctures. Face with
small punctures separated by a puncture
width, integument between smooth. Vertex
with punctures becoming faint, coarse, and
sparsely distributed. Gena sculptured like
face. Postgena imbricate and impunctate.
Mesoscutum and scutellum with small punctures separated by a puncture width or less,
integument between smooth. Tegula impunctate and smooth. Metanotum rugulose. Pleura
with small punctures separated by a puncture
width or less, integument between smooth.
Terga and sterna imbricate.
Head dark brown to black with light
brown to yellowish marks lateral to clypeus
and near lower, inner margin of compound
eyes. Mesosoma dark brown to black; legs
dark brown. Wing membrane lightly fuscous;
veins dark brown. Metasoma dark brown.
Labrum with scattered, short, erect, simple
setae. Face with scattered, short, suberect,
simple setae. Postgena with moderate-length,
erect, simple setae. Mesoscutum with moderate-length, erect, simple seate, a few with
minute branches. Scutellum with pubescence
as described for mesoscutum except setae
longer and with more numerous branches
along posterior border. Metanotum with
short, scattered setae each with minute
branches. Pleura with scattered, long, simple
setae, a few with branches, setae becoming
progressively longer and more dense ventrally. Scopa on metatibia and metabasitarus
composed of dense, long, plumose setae; metafemur with long, branched setae on upper
and anterior border. Terga with sparse, minute, simple setae; sterna with scattered,
long, simple setae.
MATERIAL: One specimen. Holotype. Female (SAMH) labeled: Holotype, Eomacropis glaesaria Engel.
ETYMOLOGY: The specific epithet is derived from the Latin word glaesarius (meaning of amber).
COMMENTS: The holotype of Eomacropis
is preserved with a staphyliniform beetle lar-
49
va on its dorsum (fig. 22, pl. 1f). Presumably

the two insects were accidentally preserved
together, as there is no such association
known among living bees today; this, however, does not preclude such an association
in the past. Moreover, the beetle is simply
too large to have been carried by the bee in
the same manner as bees carry triungulin
meloids.
LONG-TONGUED BEES
Family MEGACHILIDAE Latreille
DIAGNOSIS: The megachilids are most notable for the development of the scopa on the
metasomal sterna [subsequently reduced on
the hind legs in Lithurginae (tribe Lithurgini)
and Megachilinae; see below] and the structure of the subantennal suture which meets
the outer margin of the antennal socket in
Lithurginae and Megachilinae. The labrum is
broadly articulated to the head capsule and
in all except some Fideliinae and the fossils
discussed below it is longer than broad.
DESCRIPTION: Labrum broadly articulated
to clypeus, frequently longer than wide. Single subantennal suture; suture meets lower
margin of antennal socket in Fideliinae,
meets outer margin of antennal socket in Megachilinae and Lithurginae. Facial foveae absent. Lower lateral margins of clypeus not
bent posteriorly on either side of labrum. Flabellum present; glossa acute; labial palpus
with first two segments flattened, sheath-like,
and elongate; submentum strongly V-shaped
and sclerotized, submental arms articulating
with cardines slightly above cardo-stipital
articulations. Galeal comb absent; stipital
comb and concavity present; basistiptial process elongate. Preepisternal groove absent.
Mesocoxa entirely exposed. Metabasitibial
plate variable (present in Protolithurgini and
some Lithurgini along posterior margin; absent in Fideliinae and Megachilinae). Jugal
lobe short. Metasomal scopa present (in nonparasitic females). Metapostnotum setose.
Pygidial plate and fimbria of female present
Figs. 2325. Holotype female of Eomacropis glaesaria, new species. 23. Forewing. 24. Inner surface
of metatibia and metatarsus. 25. Outer surface of metatibia, apex of metafemur, and basal portion of
metabasitarsus. Scale bars 5 1 mm (a 5 fig. 23; b 5 figs. 24, 25).
50
in Fideliinae and Lithurginae, absent in Megachilinae.

COMMENTS: The family Megachilidae consists of three subfamilies: Fideliinae, Lithurginae, and Megachilinae (see table 7). The
fideliines have at times been accorded family
rank (e.g., Moure and Michener, 1955; Michener and Greenberg, 1980). Within the family the scopa on the hind legs has generally
been reduced (owing to the use of the metasomal scopa). It would appear that the scopa of the hind legs has been primitively retained in Fideliinae and an extinct tribe of
Lithurginae (Protolithurgini; see below);
however, these setae are not used for the
transport of pollen (like other megachilids,
pollen is transported in the metasomal scopa)
and are instead used in flinging sand during
nest building (Rozen, 1970, 1973). Thus,
these setae in fideliines are not a true scopa.
NO. 259
TABLE 7
Hierarchical Suprageneric Classification of

Megachilidae
Key to Subfamilies of Megachilidae

in Baltic Amber
1. Metabasitibial plate present (represented solely along posterior border in living taxa); T1
short and flat in profile; pygidial plate present (sometimes reduced to strong apical
spine) . . . . . . . . . . . Lithurginae Newman
Metabasitibial plate absent; T1 not shortened, convex in profile; pygidial plate absent . . . . . . . . . . Megachilinae Latreille
Subfamily LITHURGINAE Newman

DIAGNOSIS: The Lithurginae can be immediately differentiated by the short, flat first
metasomal tergum and the presence of both
metabasitibial and pygidial plates. From the
Fideliinae it differs by the presence of only
two submarginal cells, the reduced seventh
sternum in males, and the fusion or loss of
volsellae. Megachilines have lost the metabasitibial and pygidial plates and have a longer first metasomal tergum that is convex in
profile. Extant lithurgines can further be recognized by their distinctly spiculate tibiae
and elongate glossae.
DESCRIPTION: Proboscis elongate, often
reaching metasoma in repose (only in the extant tribe Lithurgini); third segment of labial
palpus flattened, on same axis as second segment. Labrum longer than broad (except in
Protolithurgus where the length is apparently
equal to the width). Mandibles in recent spe-
cies with three or more teeth, typically short

and robust. Subantennal sutures directed to
outer margins of antennal sockets. Outer surfaces of tibiae with hairless spicules (except
in Protolithurgus and some males of recent
species); metabasitibial plate present; metabasitarsus slender and cylindrical (except in
Protolithurgus). Forewing with two submarginal cells; jugal lobe of hind wing half or
more as long as vannal lobe. Metasomal T1
shortened, flattened in profile, posterior margin rounded. Pygidial plate and pygidial fimbria present. Females typically with a facial
prominence below antennal sockets, composed of either clypeus or supraclypeal area
(all species of Microthurge, nearly all Lithurgus, and a few Trichothurgus).
COMMENTS: Although classically given subfamilial rank (e.g., Michener, 1944, 1983;
Moure, 1949), the group has recently been
downgraded to a tribe at the base of the larger subfamily Megachilinae (Michener,
2000a). This has its advantages by emphasizing that lithurgines share more characters
in common with the megachilines and that
2001
the difference between the Fideliinae and the

remainder of Megachilidae (i.e., Lithurginae
and Megachilinae) is great. However, the lithurgines are just as distinctive in their own
right, with numerous great differences from
the Megachilinae, while differences among
the megachiline tribes are far less significant.
Thus, since the monophyly of the groups is
well established, the recognition of the lithurgines as a separate subfamily seems well
warranted and of some use to emphasize
their biological and morphological separation
from the Megachilinae. Therefore, I here return them to subfamilial rank within the Megachilidae. The subfamily Lithurginae is distributed throughout the world with its greatest diversity in arid regions of South America.
The subfamily has not previously been recorded from any amber deposit and, although
one compression fossil has been assigned to
Lithurgus, definitive lithurgines are not
known from the geological record. Lithurgus
adamiticus (Heer), a compression fossil from
the Miocene of Oeningen, was originally
proposed as a species of honey bee (Heer,
1865). Cockerell (1909a) later transferred the
species to Lithurgus on the basis of overall
body shape and wing venation. In actuality,
the fossil does not preserve characters that
would definitively assign it to this subfamily.
Thus, the amber inclusion described below is
the first definitive fossil lithurgine and serves
as both the oldest record of the subfamily
and the only one preserved in amber.
A hypothesis of generic relationships within the Lithurginae, Recent and fossil, is presented below (see Cladistic Analyses).
PROTOLITHURGINI, New Tribe
TYPE GENUS: Protolithurgus Engel, new
genus.
DIAGNOSIS: This group can be readily separated from its living sister taxon, the Lithurgini, by the large, strongly bordered metabasitibial plate, the flattened metabasitarsus, the bidentate mandibles, the absence of
metatibial spicules, and the relatively short
labrum.
DESCRIPTION: Third labial palp segment on
same axis as second. Labrum about as long
as wide. Mandible bidentate (fig. 26). Outer
51
surfaces of tibiae not spiculate; metabasitibial plate present, strongly bordered on all
sides (fig. 28); metatibia with weakly formed
scopa in addition to metasomal scopa; metabasitarsus flattened; female claw cleft (fig.
29); female arolium present (fig. 29).
COMMENTS: The Protolithurgini is significant among lithurgines for retaining the plesiomorphic condition of a relatively short
glossa, bidentate mandibles, and metatibial
scopa (as well as a metasomal scopa). It can
be confidently classified in the Lithurginae
owing to the apomorphic presence of the
shortened and flat first metasomal tergum
(unique to lithurgines among the Megachilidae), the third labial palpus on the same axis
as the second, and the retention of a metabasitibial plate (although even more well developed than in extant lithurgines).
Protolithurgus, new genus
TYPE SPECIES: Protolithurgus ditomeus Engel, new species.
DIAGNOSIS: Labrum apparently as long as
wide, with numerous, erect setae on surface.
Mandible not short and stout (as is typical
for extant lithurgines), with only two strongly developed teeth (fig. 26). Clypeus not extending below lower tangent of compound
eyes (figs. 26, 27); epistomal sulcus forming
a slightly obtuse angle; supraclypeal area
convex, gently swollen and protuberant in
lateral view, extending above level of clypeus (fig. 27). Subantennal sutures meeting
outer margin of antennal sockets (fig. 26); F1
distinctly longer than F2 (fig. 26). Compound
eyes not emarginate, inner margins parallel
(fig. 26). Preoccipital ridge rounded. Tibiae
not spiculate; metabasitibial plate present,
strongly bordered on all sides, large and
broadly rounded (fig. 28); claws cleft (fig.
29); arolium present (fig. 29). Metasomal
scopa strong and present; T1 relatively short,
flat in lateral view.
is a combination of protos (Greek, first)
and the genus name Lithurgus, type genus of
the subfamily. The name is masculine.
Protolithurgus ditomeus, new species
Figures 2629
Plate 2bd

D ESCRIPTION : Female. Preserved body
52
NO. 259
Figs. 2629. Holotype female of Protolithurgus ditomeus, new species. 26. Frontal view of head.
27. Right lateral view of head. 28. Outer surface of metatibia. 29. Inner view of claw and arolium.
Scale bars 5 1 mm (a 5 fig. 26; b 5 fig. 27; c 5 figure 28; d 5 fig. 29).
2001
length 12.86 mm (from head to approximately the basal portion of T5). Head wider
than long (length 3.10 mm, width 4.40 mm).
Clypeus not extending below lower tangent
of compound eyes; convex and somewhat
protuberant in lateral view (fig. 27). Upper
interorbital distance 2.35 mm; lower interorbital distance 2.37 mm. Interocellar distance 0.55 mm; ocellocular distance 0.60
mm; median to lateral ocellus 0.15 mm. Intertegular distance 3.38 mm. Scutellum
slightly more than three times length of metanotum; metanotum about as long as basal
area or propodeum. Basal vein basad by vein
width, straight; first submarginal cell apparently subequal in length to second submarginal cell.
Labrum, clypeus, supraclypeal area, face,
vertex, gena, and postgena imbricate. Mesosoma imbricate. Metasoma apparently
(where evident) imbricate.
Head and mesosoma brown with nonmetallic, greenish highlights; antennae, metasoma, and legs dark brown without highlights. Wing membrane fuscous; veins dark
brown to black.
Pubescence fuscous to black. Labrum with
numerous, long (22.5 OD), simple, erect setae, setae not obscuring integument. Clypeus,
supraclypeal area, and face below level of
antennal sockets with scattered, simple, erect,
long (23 OD) setae; clypeal margin with
submedial patch of setae; face above level of
antennal sockets, vertex, and gena with setae
sparser and shorter (1 OD). Postgena with
long (23.5 OD), erect, simple setae. Pronotum with minute, sparse, simple setae except dorsolateral angle with plumose (1 OD)
setae. Mesoscutum with minute, sparse, simple setae. Scutellum as described for mesoscutum except axilla with patches of erect,
plumose (1 OD) setae. Metanotum without
setae. Hypoepimeral area, metepisternum,
and lateral surface of propodeum with sparse,
suberect, simple (1 OD) setae; preepisternum
and mesepisternum with scattered, long (2.5
3 OD), erect setae each with a few short
branches. Basal area of propodeum without
pubescence. Metafemur without scopal setae;
metatibia and metabasitarsus with dense,
simple setae, a few with two or three short
branches. Metasomal terga apparently (where
evident) with sparse, minute, simple setae;
53
metasomal sterna with scopa, sterna with

dense, elongate (45.5 OD), simple, erect setae, setae with sinuous apices.
MATERIAL: One specimen. Holotype. Female, B-W 157 (AMNH) labeled: Baltic
Holotype, Protolithurgus ditomeus Engel.
The caudal end of the holotype of Protolithurgus is badly damaged. The apex of the
wings and metasoma reach to the amber surface where the amber becomes extremely
turbulent (numerous, strong, bending flow
lines). The apices of the wings are missing
as is, apparently, the apex of the metasoma.
The apical halves of the forewings appear as
though they have been cleared and even heated during preservation (while the resin was
still soft). Although the veins are no longer
pigmented the membrane of the wing is still
preserved (the minute setae of the wing
membrane can be easily seen at higher magnification) and by slowly searching the wing
membrane it is possible to locate the remnants of veins and to follow their courses.
Thus, it is possible to determine to some degree what the forewing venation of this species had been. It is also difficult to examine
T1 but its apical margin can be seen from
above and between the bases of the forewings (which are folded over the metasoma)
as well as in lateral view.
ETYMOLOGY: The specific epithet is a reference to the two teeth of the mandible by
comparison to the three teeth of Recent lithurgines (Gr. di, meaning two, and tomeus, meaning tooth).
COMMENTS: The holotype of Protolithurgus ditomeus has at least eight triungulin
meloids hanging onto seta of the mesosomal
venter as well as one attached to a seta of
the gena. Although a triungulin is recorded
by Larsson (1978) in Baltic amber, the specimen was found in isolation from any other
insects and its identity was suspect. The
triungula reported here are presently the oldest, definitive record of the host-parasite association between bees and meloid beetles as
well as the oldest, definitive fossil of a meloid triungulin.
Subfamily MEGACHILINAE Latreille
Megachiles Latreille, 1802b: 381. Type genus:
Megachile Latreille, 1802a.
54
DIAGNOSIS: Refer to Diagnosis for subfamily Lithurginae (above).

DESCRIPTION: Proboscis typically not elongate (as in Recent lithurgines); third segment
of labial palpus projecting laterally from axis
of second segment (except in some Chelostoma and other groups). Labrum longer than
broad (except in Glyptapina and Ctenoplectrellina). Mandibles with a variable number of
teeth, typically short and robust. Subantennal
sutures directed to outer margin of antennal
sockets. Outer surface of tibiae frequently not
spiculate (except in some Anthidiini); metabasitibial plate absent; metabasitarsus flattened. Forewing with two submarginal cells
(i.e., 1rs-m absent); jugal lobe of hind wing
short. Metasomal T1 not shortened, convex in
profile, posterior margin straight or convex.
Pygidial plate and pygidial fimbria absent.
Tribe OSMIINI Newman
Osmiites Newman, 1834: 401. Type genus: Osmia
Panzer, 1806.
Chelostomidae Kirby, 1837: 270. Type genus:
Chelostoma Latreille, 1809.
Glyptapinae Cockerell, 1909b: 13. Type genus:
Glyptapis Cockerell, 1909a. NEW SYNONYMY.
Trypetini Robertson, 1903: 164. Type genus: Trypetes Schenck, 1861.
Heriadini Michener, 1941: 152. Type genus: Heriades Spinola, 1808.
DIAGNOSIS: Megachiline bees with an arolium, an elongate pterostigma, 2m-cu basad

2rs-m, and frequently with simple claws (except in the Baltic amber genera and one living group).
DESCRIPTION: Arolium present; claws simple (except cleft in Glyptapina, Ctenoplectrellina, and Metallinella). Pterostigma longer than wide; 2m-cu basad 2rs-m. Integument without maculations (except in Ochreriades) and frequently metallic or with
metallic highlights.
COMMENTS: Phylogenetic affinities among
osmiine genera are unclear and difficult to
resolve. The tribe is very likely paraphyletic
with respect to at least the Megachilini.
Michener (1941) attempted to divide the
group into two tribes, the Osmiini s.s. and
the Heriadini [roughly equivalent to the Osmia- and Heriades-genus groups of Griswold
(1985), Griswold and Michener (1997), and
Michener (2000a)]. Although these groups
NO. 259
are not presently well defined, I prefer to tentatively recognize them as subtribes for purposes of comparison with the plesiomorphic
fossils discussed below. The Osmiini can
therefore be divided into four groups or subtribes as indicated in table 7; the fossils fall
into two distinct subtribes: Glyptapina Cockerell and Ctenoplectrellina (new subtribe).
The two fossil subtribes retain a short labrum, a plesiomorphic feature for Lithurginae and Megachilinae; however, the presence
of several derived characters typical of Megachilinae (e.g., complete absence of metabasitibial plate, absence of pygidial plate)
suggests that this character is a reversal. Subtribes of Osmiini can be distinguished as follows (based on females only):
Osmiina Newman: Maxillary palpi fouror five-segmented; labrum longer than wide;
mandible variously structured; parapsidal
lines punctiform (in Osmia) or linear; omaular ridge rounded; basal area of propodeum
declivitous; claws simple (except in Metalinella).
Trypetina Robertson, nomen translatum
[5 Heriadini of Michener (1941)]: Maxillary
palpi two- to four-segmented; labrum longer
than wide; mandible variously structured;
parapsidal lines linear; omaular ridge usually
rounded (carinate in some); basal area of propodeum usually horizontal; claws simple.
The family-group name based on Trypetes (a
junior synonym of Heriades) has priority
over that based on Heriades. The name Trypetini cannot be dismissed simply owing to
the synonymy of its type genus (ICZN,
1999b: Art. 40.1) and, thus, the subtribal
name should be Trypetina Robertson.
Glyptapina Cockerell, nomen translatum
(includes only Glyptapis Cockerell): Maxillary palpi four-segmented; labrum wider than
long; mandible with long, upper cutting-edge
(figs. 30, 32) (mandible in general structure
similar to some species of Anthidiini; e.g.,
some Dianthidium); parapsidal lines linear;
omaular ridge strongly carinate (as in some
Old World Anthidiini); basal area of propodeum declivitous; claws cleft (fig. 35).
Ctenoplectrellina, new subtribe (type genus: Ctenoplectrella Cockerell; subtribe also
includes Glaesosmia, n. gen.): Maxillary palpi four-segmented; labrum wider than long;
mandible variously structured; parapsidal
2001
Fig. 30.
55
Right lateral view of head of holotype female of Glyptapis mirabilis Cockerell.
lines linear; omaular ridge rounded; basal

area of propodeum gently sloping, not
strongly declivitous (fig. 46); claws cleft (fig.
42).
It may be that Osmiini, incorporating the
fossil lineages, is paraphyletic with respect
to not only the Megachilini but the Anthidiini as well. It is possible that Cockerells
Glyptapina (Cockerell, 1909b: Glyptapinae
in his system) is more closely allied to Anthidiini, while Ctenoplectrellina is sister to
living Osmiini 1 Megachilini. Once the classification of Osmiini has been clarified
through a detailed cladistic study, the fossil
groups will likely need to be elevated to tribal status (as will other generic groups of Osmiini).
Key to Genera of Osmiini
in Baltic Amber
1. Mesepisternum punctate, not areolate; omaular carina absent, omaulus rounded; propodeum imbricate or smooth, without are-
olae; mandible with three teeth, short cutting edge between uppermost tooth and
lower two teeth (fig. 40); mesoscutum and
scutellum normally punctate or impunctate;
compound eyes bare (fig. 40) . . . . . . . . 2
Mesepisternum coarsely areolate and rugose
(figs. 33, 39); omaular carina present and
strong (fig. 33); propodeum strongly areolate (fig. 31); mandible with a single, small
lower tooth and large upper cutting edge
(fig. 32); mesoscutum and scutellum coarsely punctate, punctures deep and craterlike
(figs. 31, 38); compound eyes hirsute, setae
long (fig. 32) . . . . . . Glyptapis Cockerell
2. Upper gena much narrower than compound
eye in lateral view; subantennal sutures
longer than diameter of antennal sockets
(fig. 40); distance between compound eyes
equal to or slightly less than length of compound eyes; second abscissa Rs orthogonal
to vein M (figs. 41, 43) . . . . . . . . . . . . . .
. . . . . . . . . . . . . Ctenoplectrella Cockerell
Upper gena as broad as compound eye in lateral view; subantennal sutures as long as
diameter of antennal sockets; distance be-
56
tween compound eyes greater than length

of compound eyes; second abscissa of Rs
angulate such that angle between this vein
and M opening toward wing apex is acute
(fig. 47) . . . . . . . . . . Glaesosmia, n. gen.
Genus Glyptapis Cockerell

Glyptapis Cockerell, 1909a: 314. Type species:
Glyptapis mirabilis Cockerell, 1909a, monobasic [also designated by Cockerell, 1909b].
Cockerell, 1909b: 13.
DIAGNOSIS: This genus can be instantly

recognized by the strong areolate sculpturing
of the pleura and propodeum as well as the
strong and large punctures of the mesoscutum and scutellum. Furthermore, the long
cutting edge of the mandible with a single,
sharp, lower tooth and the hirsute compound
eyes serve to distinguish Glyptapis species
from not only Baltic amber megachilids but
from all other Baltic amber bees. Glyptapis
can be separated from other genera, living
and extinct, placed in the paraphyletic Osmiini by the characters presented above for
the subtribe Glyptapina.
D ESCRIPTION : Mandible with a single,
small, sharp tooth on lower apical margin
and large upper cutting edge (the mandible
is similar in general structure to that of some
anthidiines) (figs. 30, 32, 33). Malar space
short, much shorter than basal mandibular
width. Clypeus flat; extending just below
lower tangent of compound eyes (fig. 32).
Hypostomal ridge carinate, anterior angle
rounded. Compound eyes hirsute, setae long
(much longer than thrice ommatidial diameter) (fig. 32); compound eye broader than
gena in lateral view; inner margins straight
and slightly converging below. F1 longer
than F2; length of F2 approximately equal to
that of F3. Posterior margin of vertex gently
concave. Preoccipital area sharply angled
and weakly carinate. Mesoscutum and scutellum coarsely and deeply punctate (figs. 31,
38), mesoscutal anterior border broadly
rounded; median line moderately impressed;
parapsidal lines faintly impressed and linear
(not punctiform); tegula oval; scutellum low,
weakly convex, not overhanging metanotum,
anterior margin with distinct, small, median
V-shaped notch, smaller lateral notches present along anterior margin where scutellum
laterally bends toward mesoscutum; metan-
NO. 259
otum slightly inclined but generally not far

off from horizontal; mesepisternum strongly
areolate, with precoxal triangle along anterior border of mesocoxa; scrobal suture absent; anterior and posterior borders of metepisternum converging ventrally and meeting
one another at point of mesocoxal base; propodeum strongly areolate. Claws with strong
inner tooth, inner tooth shorter than outer;
arolium present (fig. 35); outer apex of mesotibia with a very short, broad, apical spine,
similar spine on protibial apex, although
much shorter than that of mesotibia; two metatibial spurs, spurs minutely ciliate; metatiba
with scattered, elongate, simple setae (fig.
36). Basal vein strongly arcuate (fig. 37). No
apparent maculations on integument (coloration is, however, not well preserved in any
of the known specimens).
COMMENTS: I had earlier not considered
Glyptapis to be a megachilid (Engel, 1999d);
however, having now seen type material of
this genus in the Institut und Museum fur
Geologie und Palaontologie, I recognize that
this genus is, in fact, quite clearly an enigmatic lineage of megachilines (as is the genus Ctenoplectrella; see below). The presence of a single subantennal suture that
meets the antennal sockets at their outer margin is known only in Megachilidae and the
absence of basitibial and pygidial plates as
well as possession of only two submarginal
cells is indicative of Megachilinae. Although
the coarse sculpturing on the pleura is somewhat suggestive of some Old World Anthidiini, as is the presence of a strong omaular
carina, the apparent absence of integumental
markings, the basal position of 1m-cu relative to the second abscissa of Rs, and the
relatively long pterostigma excludes Glyptapis from that tribe (see also Comments
above under Osmiini). The subtribe Glyptapina (defined above) may eventually prove to
represent a lineage more closely allied to the
anthidiines (or from which the anthidiines
arose: i.e., glyptapines may be stem-group
anthidiines), in which case tribal status for
this fossil group would be warranted owing
to its numerous, plesiomorphic osmiine features, not too mention its own peculiar apomorphies (i.e., sculpturing of mesosoma,
elongate eye setae).
2001
Key to Species of GLYPTAPIS

1. Mesoscutum with punctures approximately
one-third ocellar diameter or less and well
spaced (not nearly contiguous) (fig. 38);
precoxal triangle bordered anteriorly by
narrow, glabrous margin many times less
than width of areolae . . . . . . . . . . . . . . . 2
Mesoscutum with punctures large, approximately one-half ocellar diameter, and
nearly contiguous (fig. 31); precoxal triangle bordered anteriorly by wide glabrous margin at least 1.5 times width of
areolae . . . . . . . . . G. mirabilis Cockerell
2. Precoxal triangle containing distinct transverse ridges or striae to form a single, dorsoventral row of areolae . . . . . . . . . . . . 3
Precoxal triangle without areolae (fig. 33) . .
. . . . . . . . . . . . . . . . . G. disareolata, n. sp.
3. Deep, crater-like punctures of scutellum not
restricted to posterior two-thirds; metanotum with minute punctures separated by 2
3 times puncture width. . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . G. fuscula Cockerell
Deep, crater-like punctures of scutellum restricted to posterior two-thirds; metanotum
with small punctures densely packed, separated by puncture width or less . . . . . .
. . . . . . . . . . . . . . G. densopunctata, n. sp.
Glyptapis mirabilis Cockerell

Figures 30, 31
Frontispiece and Plate 2e
Glyptapis mirabilis Cockerell, 1909a: 314.
DIAGNOSIS: This species can be separated

from all other Glyptapis by the large and
nearly contiguous punctures of the mesoscutum. Additionally, the precoxal triangle is
well separated from the areolate sculpturing
of the remainder of the mesepisternum; the
width of the glabrous margin demarcating
the anterior border of the precoxal triangle is
more than the diameter of the areolae (approximately 1.5 times areolae diameter in
width).
width 1.56 mm). Upper interorbital distance
ocellus 0.12 mm. Intertegular distance 1.48
mm. Basal vein strongly arcuate, confluent
with cu-a; first submarginal cell longer than
57
second. 1m-cu enters second submarginal

cell about four times vein width distad of
second abscissa of Rs; 2rs-m distad 2m-cu
by two times vein width; second submarginal
cell gently narrowed anteriorly; 2rs-m gently
curved; cu-a long; inner margin of pterostigma in first submarginal cell longer than width
of pterostigma; inner margin of pterostigma
in marginal cell gently convex; marginal cell
apex acutely rounded, offset from wing margin by width of vein, not appendiculate. Six
distal hamuli arranged in a single, evenly
spaced series.
Integument generally shining. Mandible
smooth and shining, without outer grooves,
with faint, small punctures separated by less
than a puncture width on basal half. Clypeus
with small, faint punctures separated by 12
times puncture width, such punctures particularly faint on apical third, integument between punctures smooth. Supraclypeal area
sculptured as on clypeus. Face with minute
punctures, more well-defined than those of
clypeus and supraclypeal area, punctures
separated by 12 times puncture width, integument between punctures smooth; punctures becoming faint on vertex. Gena with
minute, faint punctures separated by 23
times puncture width, integument between
punctures smooth. Postgena smooth and impunctate. Pronotum with very sparse, minute
punctures, also with minute, transverse striae
running along posterior edge of lateral ridge,
integument otherwise smooth. Mesoscutum
strongly punctured; punctures large (ca. 0.5
OD) and deep, separated by less than onehalf a puncture diameter (i.e., nearly contiguous), integument between such punctures
smooth, with minute punctures separated
(where apparent) by three or more times their
own diameter (fig. 31). Tegula with minute
punctures separated by two times puncture
width, integument between smooth. Scutellum sculptured as on mesoscutum. Metanotum with large lateral pits, medially with minute punctures separated by 23 times puncture width, integument otherwise smooth.
Preepisternal area (here used for anterior-facing surface of mesepisternum anterior to
omaular carina: preepisternal suture is absent, as in other long-tongued bees) smooth
and impunctate. Mesepisternum coarsely reticulate-areolate, integument inside areolae
58
Fig. 31.
NO. 259
Dorsal view of mesosoma of holotype female of Glyptapis mirabilis Cockerell.
2001
smooth; scrobal suture absent (perhaps utterly obscured by coarse areolate sculpturing?);
precoxal triangle with eight dorsoventral areolae (the number of areolae, however, are
variable; in one other specimen there are
only four areolae); precoxal triangle anteriorly separated from remainder of mesepisternum by broad glabrous margin that is approximately 1.5 times areola diameter in
width. Metepisternum transversely striate, integument between striae smooth. Basal area
of propodeum strongly areolate, areolae easily 1 OD in diameter, basal area two areolae
in length and eight areolae wide, lateral areolae more rectangular than square areolae
near midline; metapostnotum smooth and
impunctate, bordered by areolae (0.5 OD in
diameter) that converge ventrally just below
propodeal pit; outside of areolate borders of
metapostnotum on posterior propodeal surface a small glabrous zone of about 1 OD in
width; lateral and ventral borders of posterior
surface strongly areolate; lateral surface of
propodeum dorsoventrally striate, integument between striae smooth. Metasomal terga and sterna faintly imbricate and impunctate.
Color not well preserved, apparently entirely dark brown without maculations. Wing
membrane hyaline; veins strong and dark
brown.
Mandible with minute, simple, subappressed
setae, longer on ventral border. Galea with
long setae at apex and on inner border near
apex. Clypeus with minute, appressed, simple setae not obscuring integument, such setae also on face below level of antennal
sockets. Widely scattered, longer, erect setae
on face, clypeus, supraclypeal area; appressed, minute setae disappearing by supraclypeal area, replaced on upper face by
aforementioned longer, erect setae, although
now more numerous, such setae long on
vertex and each bearing a few minute
branches. Gena with similar setae as those
of vertex, although shorter, subappressed
(not obscuring integument), and simple.
Postgena with long, erect, widely scattered
setae. Pronotum with scattered, minute, simple setae, appressed but not obscuring the
integument except on pronotal lobe where
setae appear dense and short and along dorsal carina, with setae short, erect, and each
59
with a few minute branches. Mesoscutum

with short, plumose setae arising out of
large, deep punctures, also with minute,
sparse setae in interspaces between punctures. Tegula with minute, appressed, simple
setae not obscuring integument. Scutellum
with pubescence as on mesoscutum except
setae slightly longer. Metanotum with minute, erect, simple setae widely scattered.
Preepisternal area without pubescence; setae
of mesepisternum (posterior to omaular carina) as on mesoscutum; metepisternum
with scattered, short, simple setae. Basal
area and posterior surface of propodeum
without pubescence; lateral surface of propodeum with pubescence as described for
metepisternum. Pubescence of legs generally simple and short except those setae of
tarsi long; inner surface of metafemur without pubescence, outer, anterior, and posterior surfaces with short, branched setae; metatibia with numerous, somewhat long, setae
on all surfaces, most of those on inner surface simple but a few with minute branches,
those of outer surface with several short
branches. Central discs of metasomal terga
with sparse, minute, simple, appressed setae, setae longer and erect to suberect on
lateral borders; sterna as for terga except
along apical thirds setae longer, erect, simple, and more numerous (to form metasomal
scopa).
MATERIAL: Three specimens. Holotype.
Female, K72 (IMGP) labeled: K72, Glyptapis mirabilis n. sp., Geologisch-Palaontologisches Institut und Museum, Gottingen
// Holotype, Glyptapis mirabilis Cockerell.
Non-type. Female (CMSE) labeled:
Glyptapis mirabilis? Cockerell, det. M. S.
Engel.
Non-type. Female, Nr. 3300524 (CGHG)
labeled: Nr. 3300524 // Glyptapis mirabilis Cockerell, det. M. S. Engel.
Glyptapis disareolata, new species
Figures 3235
Plate 2f
DIAGNOSIS: This species is most similar to

G. fuscula but differs most notably by the
absence of areolae in the precoxal triangle.
60
Fig. 32. Frontal view of head of holotype female of Glyptapis disareolata, new species. Scale
bar 5 1 mm.

mm. Basal vein confluent with cu-a; first
submarginal cell longer than second; 1m-cu
enters second submarginal cell about four
times vein width distad of second abscissa of
Rs; 2rs-m distad 2m-cu by two times vein
width; second submarginal cell gently narrowed anteriorly; 2rs-m gently curved; cu-a
long; inner margin of pterostigma in first
submarginal cell longer than width of pterostigma; inner margin of pterostigma in marginal cell gently convex; marginal cell apex
acutely rounded, offset from wing margin by
width of vein, not appendiculate; six distal
hamuli, arranged in a single evenly spaced
series (fig. 34).
with minute punctures separated by 12
times a puncture width, integument between
smooth. Clypeus with minute punctures separated by 12 times puncture width, integument between smooth. Supraclypeal area
sculptured as on clypeus. Face and vertex
NO. 259
with minute punctures, more well-defined

than those of clypeus and supraclypeal area,
punctures separated by a puncture width, integument between smooth. Gena and postgena with minute punctures separated by a
puncture width, integument between smooth.
Pronotum with minute punctures separated
by a puncture width, also with minute, transverse striae running along posterior edge of
lateral ridge, integument between smooth.
Mesoscutum strongly punctured; punctures
moderately large (ca. 1/3 OD) and deep, separated by less 0.52 times puncture width,
integument between such punctures smooth
times their own diameter; anterior border of
mesoscutum with deeper puncture closely
packed. Tegula with minute punctures separated by a puncture width, integument between smooth. Scutellum sculptured as on
mesoscutum except deep punctures restricted
to posterior two-thirds. Metanotum with minute punctures separated by a puncture
width, integument between smooth. Preepisternal area (here used for anterior-facing surface of mesepisternum anterior to omaular
carina: preepisternal suture is absent, as in
other long-tongued bees) with minute punctures separated by a puncture width, integument between smooth. Mesepisternum
coarsely reticulate-areolate, integument inside areolae smooth; precoxal triangle anteriorly separated from remainder of mesepisternum by narrow glabrous margin that is
less than 0.5 times areola diameter in width,
precoxal triangle without areolae (fig. 33).
Metepisternum transversely striate, integument between striae smooth. Basal area of
propodeum strongly areolate, areolae easily
1 OD in diameter, basal area two areolae in
length and eight areolae wide; metapostnotum smooth and impunctate, bordered by
areolae (0.5 OD in diameter) that converge
ventrally just below propodeal pit; outside of
areolate borders of metapostnotum on posterior propodeal surface a small glabrous
zone of about 1 OD in width; lateral and ventral borders of posterior surface strongly areolate; lateral surface of propodeum dorsoventrally striate, integument between striae
smooth. Metasomal terga and sterna faintly
imbricate.
Coloration not well preserved; however,
2001
61
Fig. 33. Right ventrolateral oblique view of head and mesosoma of holotype female of Glyptapis
disareolata, new species.
where evident, dark brown and without maculations. Wing membrane hyaline; veins
strong and dark brown.
setae, longer on ventral border. Clypeus with
Fig. 34.
minute, appressed, simple setae not obscuring integument, such setae also on face below level of antennal sockets. Widely scattered, longer, erect setae on face, clypeus,
and supraclypeal area; appressed, minute se-
Hind wing of holotype female of Glyptapis disareolata, new species. Scale bar 5 1 mm.
62
Figs. 3536. Leg structures of Glyptapis species. 35. Claw and arolium of Glyptapis disareolata, new species. 36. Metafemur, metatibia, and
metatarsus of female of Glyptapis fuscula Cockerell. Scale bars 5 0.5 mm (a), 0.25 mm (b); (a
5 fig. 36; b 5 fig. 35).
tae disappearing by supraclypeal area; face

with longer erect setae intermixed with short,
suberect setae; vertex with long, erect setae,
each bearing a few minute branches. Gena
with similar setae as those of vertex, although shorter, subappressed (not obscuring
integument), and simple. Postgena with long,
erect, widely scattered setae. Pronotum with
scattered, minute, simple setae, appressed but
not obscuring the integument except on pronotal lobe setae dense and short and along
dorsal carina setae short, erect, and each with
a few minute branches. Mesoscutum with
short, plumose setae arising out of large,
deep punctures, also with minute, sparse setae in interspaces between punctures. Tegula
with minute, appressed, simple setae not obscuring integument. Scutellum with pubescence as on mesoscutum except setae slightly
longer. Metanotum with minute, erect, simple
setae widely scattered. Preepisternal area
without pubescence; setae of mesepisternum
NO. 259
(posterior to omaular carina) as on mesoscutum; metepisternum with scattered, short,

simple setae. Basal area and posterior surface
of propodeum without pubescence; lateral
surface of propodeum with pubescence as
described for metepisternum. Pubescence of
legs generally simple and short except those
setae of tarsi long; inner surface of metafemur without pubescence, outer, anterior, and
posterior surfaces with short, branched setae;
metatibia with numerous, somewhat long, setae on all surfaces, most of those on inner
surface simple but a few with minute branches, those of outer surface with several, short
with sparse, minute, simple, appressed setae,
setae longer and erect to suberect on lateral
borders; sternal scopa composed of dense,
long, erect, simple setae.
MATERIAL: Two specimens. Holotype. Female, B-JH 104 (AMNH) labeled: Baltic
Holotype, Glyptapis disareolata Engel.
Paratype. Female, B-W 156 (AMNH) labeled: Baltic amber: Eocene, Kaliningrad,
Yantarny // Paratype, Glyptapis disareolata Engel.
ETYMOLOGY: The specific epithet is a combination of the Latin words dis (meaning
without) and areolatus (meaning small
spaces) and is a reference to the absence of
areolae in the precoxal triangle.
Glyptapis densopunctata, new species
Figure 37
Plate 3b

G. disareolata; both species have the deep,
crater-like punctures of the scutellum restricted to the posterior two-thirds and dense
punctures on the metanotum. These characters serve to separate both G. densopunctata
and G. disareolata from G. fuscula. Glyptapis densopunctata differs from G. disareolata
by the presence of areolae in the precoxal
triangle.
mm. Interocellar distance 0.38 mm; ocello-
2001
Fig. 37. Forewing of holotype female of

Glyptapis densopunctata, new species. Scale bar
5 1 mm.
cular distance 0.31 mm; median to lateral

enters second submarginal cell about four
times vein width distad of second abscissa of
Rs; 2rs-m distad 2m-cu by two times vein
width; second submarginal cell gently narrowed anteriorly; 2rs-m gently curved; cu-a
long; inner margin of pterostigma in first
submarginal cell longer than width of pterostigma; inner margin of pterostigma in marginal cell gently convex; marginal cell apex
acutely rounded, offset from wing margin by
width of vein, not appendiculate (fig. 37);
eight distal hamuli, arranged in a single,
evenly spaced series.
moderately large (ca. 1/3 OD) and deep, separated by less 0.52 times puncture width,
integument between such punctures smooth,
63

mesoscutum with deeper punctures closely
packed. Tegula with minute punctures separated by a puncture width, integument between smooth. Scutellum sculptured as on
mesoscutum except deep punctures restricted
to posterior two-thirds. Metanotum with minute punctures separated by a puncture width
or less, integument between (where evident)
smooth. Preepisternal area (here used for anterior-facing surface of mesepisternum anterior to omaular carina: preepisternal suture is
absent, as in other long-tongued bees) with
minute punctures separated by a puncture
width, integument between smooth. Mesepisternum coarsely reticulate-areolate, integument inside areolae smooth; precoxal triangle
anteriorly separated from remainder of mesepisternum by narrow glabrous margin that
is less than 0.5 times areola diameter in
width, precoxal triangle with weak transverse
striae forming dorsoventral row of areolae.
Metepisternum transversely striate, integument between striae smooth. Basal area of
propodeum strongly areolate, areolae 1 OD
or more in diameter, basal area two areolae
in length and eight areolae wide; lateral surface of propodeum dorsoventrally striate, integument between striae smooth. Metasomal
terga and sterna faintly imbricate.
Color not entirely preserved; however,
where evident, dark brown and without maculations. Wing membrane hyaline; veins
minute, appressed, simple setae not obscuring integument, such setae also on face below level of antennal sockets. Widely scattered, longer, erect setae on face, clypeus, supraclypeal area; appressed, minute setae disappearing by supraclypeal area; face with
longer erect setae intermixed with short, suberect setae; vertex with long, erect setae,
each bearing a few minute branches. Gena
with similar setae as those of vertex, although shorter, subappressed (not obscuring
integument), and simple. Postgena with long,
erect, widely scattered setae. Pronotum with
scattered, minute, simple setae, appressed but
not obscuring the integument except on pronotal lobe setae dense and short and along
dorsal carina setae short, erect, and each with
64
a few minute branches. Mesoscutum with

short, plumose setae arising out of large,
deep punctures, also with minute, sparse setae in interspaces between punctures. Tegula
with minute, appressed, simple setae not obscuring integument. Scutellum with pubescence as on mesoscutum except setae slightly
longer. Metanotum with minute, erect, simple
setae widely scattered. Preepisternal area
without pubescence; setae of mesepisternum
(posterior to omaular carina) as on mesoscutum; metepisternum with scattered, short,
simple setae. Basal area and posterior surface
of propodeum without pubescence; lateral
surface of propodeum with pubescence as
described for metepisternum. Pubescence of
legs generally simple and short except those
setae on tarsi long; inner surface of metafemur without pubescence, outer, anterior, and
posterior surfaces with short, branched setae;
metatibia with numerous, somewhat long setae on all surfaces, most of those on inner
surface simple but a few with minute branches, those of outer surface with several short
with sparse, minute, simple, appressed setae,
setae longer and erect to suberect on lateral
borders; sternal scopa composed of dense,
long, erect, simple setae.
MATERIAL: Four specimens. Holotype. Female, B-W 161 (AMNH) labeled: Baltic
Holotype, Glyptapis densopunctata Engel.
Paratype. Female, B-JH 89 (AMNH) labeled: Baltic amber: Eocene, Kaliningrad,
Yantarny // Paratype, Glyptapis densopunctata Engel.
Paratype. Female, Nr. 2623 (CCGG) labeled: Nr. 2623 // Paratype, Glyptapis
densopunctata Engel.
Paratype. Female, Nr. 857 (CHFG) labeled: Nr. 857, Balt. Bernstein // Paratype, Glyptapis densopunctata Engel.
ETYMOLOGY: The specific epithet is a combination of the Latin words densus (meaning
dense) and punctatum (meaning punctures) and is a reference to the dense punctures of the metanotum.
Glyptapis fuscula Cockerell
Figures 36, 38, 39
Plate 3a
Glyptapis fuscula Cockerell, 1909b: 17.

Glyptapis reticulata Cockerell, 1909b: 16.
SYNONYMY.
NEW
NO. 259
Glyptapis reducta Cockerell, 1909b: 18. NEW SYNONYMY.

Glyptapis neglecta Salt, 1931: 136. NEW SYNONYMY.
Dasypoda sp. Gerlach, 1989: 253. [misidentification; 5 G. reducta Cockerell]
DIAGNOSIS: This species is similar to G.

densopunctata and G. disareolata but has the
deep, crater-like punctures not restricted to
the posterior two-thirds of the scutellum and
the punctures of the metanotum more widely
spaced.
enters second submarginal cell about eight
times vein width distad second abscissa of
Rs; 2rs-m distad 2m-cu by vein width; second submarginal cell gently narrowed anteriorly; 2rs-m gently curved; inner margin of
pterostigma in first submarginal cell longer
than width of pterostigma; inner margin of
pterostigma in marginal cell gently convex;
marginal cell apex acutely rounded, offset
from wing margin by width of vein, not appendiculate; six distal hamuli, arranged in a
single, evenly spaced series.
moderately large (ca. 1/3 OD) and deep, sep-
2001
Fig. 38.
65
Dorsolateral view of mesosoma of holotype female of Glyptapis fuscula Cockerell.
arated by less 0.52 times puncture width,

integument between such punctures smooth,
mesoscutum with deeper puncture closely
packed (fig. 38). Tegula with minute punctures separated by a puncture width, integument between smooth. Scutellum sculptured
as on mesoscutum, deep, crater-like punctures on all parts of scutellum (not restricted
to posterior two-thirds). Metanotum with minute punctures separated by 23 times puncture width, integument between smooth.
Preepisternal area (here used for anterior-facing surface of mesepisternum anterior to
omaular carina: preepisternal suture is absent, as in other long-tongued bees) with minute punctures separated by a puncture
width, integument between smooth. Mesepisternum coarsely reticulate-areolate, integument inside areolae smooth; precoxal triangle
anteriorly separated from remainder of mesepisternum by narrow glabrous margin that
is less than one-half times areola diameter in

width, precoxal triangle without strong areolae, instead with weak longitudinal striae
that extend only one-half of precoxal triangle. Metepisternum transversely striate, integument between striae smooth. Basal area
of propodeum strongly areolate, areolae easily 1 OD in diameter, basal area two areolae
in length and eight areolae wide; metapostnotum smooth and impunctate, bordered by
areolae (0.5 OD in diameter) that converge
ventrally just below propodeal pit; small glabrous zone of about 1 OD in width appearing
outside of areolate borders of metapostnotum
on posterior propodeal surface; lateral and
ventral borders of posterior surface strongly
areolate; lateral surface of propodeum dorsoventrally striate, integument between striae
smooth. Metasomal terga and sterna faintly
imbricate.
Color dark brown without maculations.
Wing membrane hyaline; veins dark brown.
66
NO. 259
Fig. 39. Lateral view (posterior portion of head in lower right-hand corner) of mesosoma of holotype
female of Glyptapis fuscula Cockerell. Mesepisternum sculpturing in focus; precoxal triangle cannot be
seen in this image.

minute, appressed, simple setae not obscuring integument, such setae also on face below level of antennal sockets. Widely scattered, longer, erect setae on face, clypeus, supraclypeal area; appressed, minute setae disappearing by supraclypeal area, replaced on
upper face by aforementioned longer, erect
setae although now more numerous, such setae long on vertex and each bearing a few
minute branches. Gena with similar setae as
those of vertex although shorter, subappressed
(not obscuring integument), and simple.
Postgena with long, erect, widely scattered
setae. Pronotum with scattered, minute, simple setae, appressed but not obscuring the integument except on pronotal lobe setae dense
and short and along dorsal carina setae short,
erect, and each with a few minute branches.
Mesoscutum with short, plumose setae arising out of large, deep punctures, also with
minute, sparse setae in interspaces between

punctures. Tegula with minute, appressed,
simple setae not obscuring integument. Scutellum with pubescence as on mesoscutum
except setae slightly longer. Metanotum with
minute, erect, simple setae widely scattered
and a medial patch of long, erect setae, each
with a few minute branches. Preepisternal
area without pubescence; setae of mesepisternum (posterior to omaular carina) as on
mesoscutum; metepisternum with scattered,
short, simple setae. Basal area and posterior
surface of propodeum without pubescence;
lateral surface of propodeum with pubescence as described for metepisternum. Pubescence of legs generally simple and short
except those setae on tarsi long; inner surface
of metafemur without pubescence, outer, anterior, and posterior surfaces with short,
branched setae; metatibia with numerous,
somewhat long, setae on all surfaces, most
2001
of those on inner surface simple but a few

with minute branches, those of outer surface
with several, short branches. Central discs of
metasomal terga with sparse, minute, simple,
appressed setae, setae longer and erect to
suberect on lateral borders; sternal scopa
composed of dense, long, erect, simple setae.
M ATERIAL : Nine specimens. Holotype
(fuscula). Female, K66 (IMGP) labeled:
Holotype, Glyptapis fuscula Cockerell //
Holotype, K66, Glyptapis fuscula n. sp.
Cockerell, Geologisch-Palaontologisches Institut und Museum, Gottingen // Glyptapis
fuscula // Glyptapis fuscula Cock.
[5Cockerell], K66.
Neotype (reticulata; here designated).
Female, B-JH 78 (AMNH) labeled: Baltic
Neotype, Glyptapis reticulata Cockerell //
Glyptapis fuscula Cockerell, det. M. S. Engel.
Holotype (reducta). Female, G4668
(IMGP) labeled: Holotype, Glyptapis reducta Cockerell // Holotype, G4668, Glyptapis reducta n. sp. Cockerell // Reservierte
Registiernummer, Dasypoda: G4668 [Gerlach handwriting] // Dasypoda sp. indet.,
Gerlach, J., 1987 // Glytapis fuscula
Cockerell, det. M. S. Engel. The eye setae
are difficult to see but present. The coarse
sculpturing, although also difficult to view
through the fine layer of Schimmel on the
specimen, can be seen.
Neotype (neglecta; here designated). Female B-JH 78 (AMNH) labeled: Neotype,
Glyptapis neglecta Salt, desig. M. S. Engel
// Glyptapis fuscula Cockerell, det. M. S.
Engel (see Comments below). This is the
same specimen designated as the neotype of
G. reticulata and thus all other labels are
identical with those listed above.
Non-type. Female (ZMUC) labeled:
Apoidea, A. Henningsen, 99, 1974 //
Glyptapis fuscula Cockerell, det. M. S. Engel. This is the specimen figured by Poinar
(1994: 75).
Non-type. Female, B-W 155 (AMNH) labeled: Baltic amber: Eocene, Kaliningrad,
Yantarny // Glyptapis fuscula? Cockerell,
det. M. S. Engel.
Non-type. Female, MB.I.1936 (ZMHB)
labeled: Museum fur Naturkunde Berlin,
Palaontologisches Museum, Inv. Nr.
67
MB.I.1936 (No. 18) // 18 // Glyptapis

fuscula Cockerell, det. M. S. Engel, 1999.
MB.I.1937, Slg. Berendt // Glyptapis
fuscula Cockerell, det. M. S. Engel.
Non-type. Female, Nr. 1015 (CHMG) labeled: Nr. 1015 // Glyptapis fuscula
Cockerell, det. M. S. Engel.
Non-type. Female (CFEG) labeled:
Glyptapis fuscula Cockerell, det. M. S. Engel.
COMMENTS: Salts description of G. neglecta agrees with G. fuscula in every respect,
particularly the sculpturing of the mesoscutum. Salt considered the distinguishing feature between his specimen and G. fuscula to
be the hyaline wing membrane in his species;
however, the surviving holotype of G. fuscula does not actually have fuscous wing
membranes. I therefore consider G. neglecta
and G. fuscula to be conspecific. The holotype of G. neglecta, however, has since become lost (it was originally in the ill-fated
Konigsberg collection). I, therefore, here
designate a neotype of a specimen from the
AMNH collection which agrees with Salts
description. Acting as first reviser (ICZN,
1999b: Art. 24.2) I have chosen G. fuscula
as the valid name for this taxon and consider
G. reducta and G. reticulata, both published
in the same article, to be junior synonyms.
Gerlach (1989) misidentified the holotype
of G. reducta as a species of the melittid genus Dasypoda (Melittidae: Dasypodainae).
There is a thin layer of mold over most of
the specimen but the punctures of the mesosoma can be easily seen through and within gaps of the mold. There is absolutely no
resemblance between G. reducta (nor any
Glyptapis for that matter) and species of Dasypoda.
Glyptapis sp. indet.
MATERIAL: One specimen. Non-type. Female, MB.I.1944 (ZMHB) labeled: Museum fur Naturkunde Berlin, Palaontologisches
Museum, Inv. Nr. MB.I.1944 // Glyptapis
sp. indet., det. M. S. Engel.
68
NO. 259
Glyptapis sp. indet.

MATERIAL: One specimen. Non-type. Female, Nr. 577/1 (CHFG) labeled: Nr. 577/
1, Bittf. [Bitterfeld] Bernstein // Glyptapis
sp. indet., det. M. S. Engel.
Genus Ctenoplectrella Cockerell
Ctenoplectrella Cockerell, 1909a: 314. Type species: Ctenoplectrella viridiceps Cockerell,
1909a, monobasic. Cockerell, 1909b: 19. Zeuner and Manning, 1976: 172.
DIAGNOSIS: Among Baltic amber Megachlinae Ctenoplectrella is most similar to Glaesosmia but differs by the longer supraclypeal
area, narrow gena, and more orthogonal second abscissa of Rs.
DESCRIPTION: Mandible with three teeth
along apical margin, upper tooth separated
from lower two teeth by a short, gently concave cutting-edge (fig. 40). Malar space
short, much shorter than basal mandibular
width. Clypeus flat; not extending below
lower tangent of compound eyes (fig. 40).
Hypostomal ridge carinate, anterior angle
rounded. Subantennal sutures longer than antennal socket diameter. Compound eyes bare
(fig. 40); compound eye broader than gena
in lateral view; inner margins straight and
slightly converging below. F1 slightly longer
than F2; length of F2 approximately equal to
that of F3. Posterior margin of vertex gently
concave. Preoccipital area rounded. Mesoscutum and scutellum not coarsely sculptured;
mesoscutal anterior border broadly rounded
(fig. 45); median line moderately impressed;
parapsidal lines faintly impressed and linear;
tegula oval; scutellum low, weakly convex,
not overhanging metanotum, anterior margin
without notches; metanotum slightly inclined
but generally not far off from horizontal (fig.
46); anterior and posterior borders of metepisternum converging ventrally and meeting
one another at point of mesocoxal base; propodeum not strongly sculptured. Claws with
inner tooth, inner tooth shorter than outer
(fig. 42); arolium present (fig. 42); outer apex
of mesotibia without spine; two metatibial
spurs, spurs serrate. Basal vein strongly arcuate; second abscissa Rs approximately orthogonal to M (figs. 41, 43). No apparent
maculations on integument.
COMMENTS: This genus has long been as-
Fig. 40. Frontal view of head of neotype female of Ctenoplectrella viridiceps Cockerell.
Scale bar 5 1 mm.
sociated with the apine genera Ctenoplectra

Kirby (In Kirby and Spence, 1826) and Ctenoplectrina Cockerell (1930b), together forming the tribe Ctenoplectrini. Oddly, the unity
of Ctenoplectrella with Ctenoplectrini seems
to be simply based upon two entirely unfounded criteria, 1. the name chosen by
Cockerell (1909a), which suggests some affinity between the groups (Ctenoplectrella is
a diminutive form of Ctenoplectra); and 2.
an anecdotal comment by Cockerell (1909b)
who said (p. 13), Their [Glyptapis and
Ctenoplectrella] nearest relative in the modern fauna appears to be Ctenoplectra..., but
provided no characters to support such a relationship. Indeed, the wing venation of
Ctenoplectrella shares a superficial similarity
to that of some Ctenoplectrini. Unfortunately, the similarity between the two groups
ends at this point (detailed above and further
below). The Ctenoplectrini was originally
proposed as a family by Cockerell (1930a),
later moved into the Melittidae as a subfamily (Michener, 1944), resurrected as a family
intermediate between short- and longtongued bees by Michener and Greenberg
(1980), only to be later recognized as a de-
2001
rived group of Apinae where it is currently

placed (Roig-Alsina and Michener, 1993;
Michener, 2000a). The placement of Ctenoplectrella has generally followed that of
Ctenoplectra (e.g., Zeuner and Manning,
1976; Michener and Greenberg, 1980), once
again, presumably owing to the common derivation of their names and Cockerells comment rather than any shared characteristics.
Zeuner and Manning (1976) followed Cockerells assertion of affinity with Ctenoplectra
and placed Ctenoplectrella and Glyptapis in
Ctenoplectrinae, at that time in the Melittidae
(sensu Michener, 1944). It is, unfortunately,
apparent that neither of these authors had actually seen specimens of the latter two genera. For instance, Zeuner and Manning (op.
cit.) characterize these groups as having the
inner metatibial spur modified as in Ctenoplectra; namely, broadened basally with numerous, elongate teeth, as well as having a
pygidial plate in females. Although the holotype for Ctenoplectrella is lost (a neotype
is designated below), the types of Glyptapis
have survived in the Institut und Museum fur
Geologie und Palaontologie, Gottingen and
none of these characters are present in Glyptapis (nor in the neotype of Ctenoplectrella).
Moreover, these distinctive characters are
never once mentioned by Cockerell (1909b)
who was quite adept and would undoubtedly
have noticed these peculiar features. Zeuner
and Manning (1976) do not provide descriptions for these taxa, instead referring to the
original works, once again suggesting that
Zeuner and Manning did not have the opportunity to see specimens of these taxa before their untimely deaths. The type of Ctenoplectrella viridiceps was certainly not available to Zeuner and Manning since it was lost
during World War II. The specimen here considered as the neotype for Ctenoplectrella
agrees in every respect with Cockerells description and, of particular importance, with
his figures (especially in the shape of the
pterostigma, which is quite distinctive). I am
therefore confident that the Ctenoplectrella
of Cockerell is the same as my own.
Ctenoplectrella differs from Ctenoplectrini
(Apidae) by the flat clypeus (not bent posteriorly at the apicolateral margins), the
coarsely serrate inner metatibial spur, the absence of a pygidial plate, the broadened man-
69
dible that is reminiscent of the Megachilinae,

the presence of a true metasomal scopa, the
strongly arcuate basal vein, the absence of
hairs on the metapostnotum, and the absence
of a metabasitibial plate, among numerous
other characters.
The absence of a preepisternal groove, the
shortened jugal lobe, and the fully exposed
mesocoxa are all characteristic of Melittidae
1 long-tongued bees, while most Apidae
have the lateral portions of the clypeus
strongly bent posteriorly, a feature absent in
Megachilidae and Melittidae (as well as
Ctenoplectrella and Glyptapis; unobservable
for Glaesosmia). Among the Megachilinae,
however, Ctenoplectrella is remarkable for
the slanting metanotum and the broad labrum
(the latter plesiomorphic trait shared with
Glyptapis).
Key to Species of CTENOPLECTRELLA
1. Crossvein 2rs-m strongly and doubly arcuate,
thus second submarginal cell more strongly
produced toward wing apex along its posterior margin (fig. 43); medioapical margin
of clypeus straight . . . . . . . . . . . . . . . . . 2
Crossvein 2rs-m relatively straight and therefore second submarginal cell not more
strongly produced toward wing apex along
its posterior margin (fig. 41); medioapical
margin of clypeus gently convex (fig. 40)
. . . . . . . . . . . . . . C. viridiceps Cockerell
2. Mesoscutum and scutellum punctate . . . .
. . . . . . . . . . . . . . . . . . C. cockerelli, n. sp.
Mesoscutum and scutellum impunctate . .
. . . . . . . . . . . . . . . . . . . C. grimaldii, n. sp.
Ctenoplectrella viridiceps Cockerell

Figures 4042
Plate 3ce
Ctenoplectrella viridiceps Cockerell, 1909a: 314.

Ctenoplectrella dentata Salt, 1931: 139. NEW SYNONYMY.
Ctenoplectrella splendens Kelner-Pillault, 1970a:
13. NEW SYNONYMY.
DIAGNOSIS: This species is distinguished

by the gently concave mediapical margin of
the clypeus (fig. 40) and weakly arcuate 2rsm (fig. 41).
70
NO. 259
Fig. 41. Forewing of neotype female of Ctenoplectrella viridiceps Cockerell. Scale bar 5 1
mm.
Fig. 42. Claw and arolium of neotype female

of Ctenoplectrella viridiceps Cockerell. Scale bar
5 0.25 mm.

mm. Interocellar distance 0.44 mm; ocellocular distance 0.33 mm; median ocellus to
lateral ocellus 0.16 mm. Median and parapsidal lines moderately impressed, parapsidal
lines elongate (not punctiform). Intertegular
distance 1.31 mm. Basal vein confluent with
cu-a; second abscissa Rs basad 1m-cu by 6.5
times vein width; 2rs-m distad 2m-cu by two
times vein width, 2rs-m relatively straight,
arching only toward its apex before meeting
vein M; second submarginal cell slightly longer than first submarginal cell; six distal
hamuli arranged in a single, evenly spaced
series.
Outer surface of mandible with minute
less, integument between smooth. Clypeus
with faint, coarse punctures separated by a
puncture width or less, integument between
smooth. Supraclypeal area and face with
punctures more well-defined than those of
clypeus and separated by 12 times puncture
width, integument between smooth. Vertex
and gena with sculpturing as on face although punctures slightly more closely
spaced, separated by about a puncture width,
integument between smooth. Mesoscutum
width or slightly less, integument between
smooth. Tegula with minute punctures separated by 12 times a puncture width, integument between smooth. Scutellum sculptured
as on mesoscutum. Metanotum impunctate
and smooth. Preepisternal area (i.e., anteriorfacing surface anterior to omaulus) essentially impunctate and smooth except for a few,
exceedingly faint, widely spaced, coarse
punctures. Mesepisternum with coarse,
slightly faint punctures separated by 12

smooth, punctures becoming somewhat
fainter ventrally and posteriorly. Metepisternum impunctate and smooth. Propodeum
(basal, lateral, and posterior surfaces) finely
imbricate. Terga with minute, faint punctures
separated by 13 times a puncture width, integument between finely imbricate; sterna
imbricate.
Coloration dark brown without maculations. Wing membrane hyaline except between C and Sc1R slightly fuscous in some
specimens; veins strong and dark brown.
Mandible with minute, simple, appressed
setae. Clypeus with minute, appressed, simple setae not obscuring integument; similar
setae also on face below level of antennal
sockets, intermixed with scattered, longer,
erect setae on face, clypeus, supraclypeal
area, gena, and postgena. Mesoscutum and
scutellum with scattered, short, simple setae
except those of scutellum slightly longer.
Pleural pubescence as described for mesoscutum. Basal area of propodeum without pubescence; lateral and posterior surfaces with
pubescence as described for mesoscutum.
Pubescence of legs generally simple and
short; inner surface of metafemur without
pubescence, outer, anterior, and posterior surfaces with short, branched setae; metatibia
with scattered, simple, moderate-length setae, a few with minute branches. Central
discs of metasomal terga with sparse, minute,
simple, appressed setae, setae longer and
erect to suberect on lateral borders; sternal
scopa composed of dense, long, erect, simple
setae.
Male. As described for the female except
2001
as indicated: Total body length 6.93 mm;

forewing length 4.00 mm. Head slightly wider than long (length 1.70 mm, width 1.86
mm). Upper interorbital distance 1.10 mm;
lower interorbital distance 1.00 mm. Interocellar distance 0.40 mm; ocellocular distance
0.17 mm. Intertegular distance 1.45 mm. Flagellum elongate, reaching to posterior margin of mesoscutum; F1 longer than F2; F2
equal in length to F3. T6 deeply bifid at apex
and slightly protuberant. Metasomal scopa
absent.
MATERIAL: Ten specimens. Neotype (viridiceps; here designated). Female, B-JH 95
(AMNH) labeled: Baltic amber: Eocene,
Kaliningrad, Yantarny // Neotype, Ctenoplectrella viridiceps Cockerell, desig. M. S.
Engel.
Neotype (dentata; here designated).
Male, NB.I.1949 (ZMHB) labeled: Neotype, Ctenoplectrella dentata Salt, desig. M.
S. Engel. This is the same as specimen as
the holotype for C. splendens and therefore
the remaining labels of the specimen are given below under C. splendens.
Holotype (splendens). Male, NB.I.1949
(ZMHB) labeled: 49 // Ctenoplectrella
splendens, Holotype, S. Kelner-Pillault det.
// Holotyp [sic], Palaontologisches Museum
Berlin, Ctenoplectrella splendens Kelner-Pillault, 1970, Baltischer Bernstein // Holotype, Ctenoplectrella splendens Kelner-Pillault.
Non-type. Female, B-JH 93 (AMNH) labeled: Baltic amber: Eocene, Kaliningrad,
Yantarny // Ctenoplectrella viridiceps
Non-type. Female, B-JH 83 (AMNH) labeled: Baltic amber: Eocene, Kaliningrad,
Yantarny // Ctenoplectrella viridiceps
Yantarny // Ctenoplectrella viridiceps?
MB.I.1940 (No. 29) // 29 // Ctenoplectrella viridiceps Cockerell, det. M. S.
Engel.
71

MB.I.1941 (No. 17) // 17 // Ctenoplectrella viridiceps Cockerell, det. M. S. Engel.
Non-type. Female, Nr. 24 (CFKG) labeled: 1987, Nr. 24 // Ctenoplectrella
viridiceps Cockerell, det. M. S. Engel.
Non-type. Female (CJVG) labeled:
Ctenoplectrella viridiceps Cockerell, det.
M. S. Engel.
Non-type. Female, Nr. 1615 (CCGG) labeled: Nr. 1615 // Ctenoplectrella viridiceps Cockerell, det. M. S. Engel.
COMMENTS: The species was proposed as
new again by Cockerell (1909b) but the
name was already made available by Cockerell (1909a).
The specimen here designated as the neotype agrees with Cockerells original description and figures, particularly in the shape of
the pterostigma. Zeuner and Manning (1976)
indicate the holotype to have been moved to
the Museum fur Naturkunde, HumboldtUniversitat, Berlin, but upon inspection of
their collection I failed to find any specimens
from the Konigsberg collection. Most material from the former Konigsberg collection
eventually ended up in the Institut und Museum fur Geologie und Palaontologie, Gottingen, where some of Cockerells other
types were discovered (e.g., those of Glyptapis), but Ctenoplectrella was not among
them. Ctenoplectrella was cataloged in the
Gottingen collection as specimen K65 but
it was not to be found. I therefore consider
the holotype to have been lost and here designate a neotype.
The species proposed by Salt (1931), a
male of Ctenoplectrella, is undoubtedly the
male for C. viridiceps. The neotype of C.
splendens in Berlin is the only existing male
specimen (of any genus!) preserved in Baltic
amber.
Ctenoplectrella cockerelli, new species
Figures 43, 44
DIAGNOSIS: This is most similar to C. grimaldii (see below) in the strongly produced
second submarginal cell (fig. 43), straight
clypeal apex, and narrow gena but differs by
72
NO. 259
Figs. 4344. Wings of holotype female of Ctenoplectrella cockerelli, new species. 43. Forewing.
44. Hind wing. Scale bar 5 1 mm.
the presence of punctures on the mesoscutum

and scutellum (absent in C. grimaldii).
lateral ocellus 0.13 mm. Intertegular distance
1.28 mm. Basal vein confluent with cu-a;
second abscissa of Rs basad 1m-cu by seven
times vein width; 2rs-m distad 2m-cu by vein
width, 2rs-m doubly arcuate; second submarginal cell slightly longer than first submarginal cell; eight distal hamuli, arranged
in a single, evenly spaced series (fig. 44).
Outer surface of mandible with minute
less, integument between smooth. Clypeus

Supraclypeal area sculptured as on clypeus.
Face with punctures separated by 11.5 times
smooth. Vertex and gena with small punctures separated by about a puncture width,
width or slightly less, integument between
smooth. Tegula with minute punctures separated by a puncture width, integument between smooth. Scutellum sculptured as on
mesoscutum. Metanotum impunctate and
smooth. Preepisternal area (i.e., anterior-facing surface anterior to omaulus) essentially
impunctate and smooth except for a few, exceedingly faint, widely spaced, coarse punc-
2001
tures. Mesepisternum with coarse, slightly

faint punctures separated by a puncture
width, integument between smooth, punctures becoming somewhat fainter ventrally
and posteriorly. Metepisternum impunctate
and smooth. Propodeum (basal, lateral, and
posterior surfaces) impunctate and smooth.
Terga with small punctures separated by 1
2 times a puncture width, integument between smooth; sterna imbricate.
Color brown without maculations. Wing
brown.
scopa composed of bands of dense, long,
erect, simple setae.
Female, B-W 160 (AMNH) labeled: Baltic
Holotype, Ctenoplectrella cockerelli Engel.
Paratype. Female, MB.I.1942 (ZMHB)
MB.I.1942 (No. 31) // 31 // Paratype,
Ctenoplectrella cockerelli Engel.
Paratype. Female, Nr. 1011 (CCGG) labeled: Nr. 1011 // Paratype, Ctenoplectrella cockerelli Engel.
ETYMOLOGY: The specific epithet is a patronymic honoring the late Prof. Theodore D.
A. Cockerell (18661948). Cockerell was the
73
first to critically examine the Baltic amber

bees and to attempt to place them into a phylogenetic context. Weber (1965, 2000) has
presented a bibliography of Cockerells papers as well as a sampling of his published
writings on topics ranging from entomology
to philosophy.
Ctenoplectrella grimaldii, new species
Figures 45, 46
Plate 3f
DIAGNOSIS: Refer to diagnosis presented

for C. cockerelli (above).
1.13 mm. Basal vein confluent with cu-a;
second abscissa of Rs basad 1m-cu by seven
times vein width; 2rs-m distad 2m-cu by vein
width, 2rs-m doubly arcuate; second submarginal cell slightly longer than first submarginal cell; eight distal hamuli, arranged
in a single, evenly spaced series.
Mandibular sculpturing not visible. Clypeus with small punctures separated by a
smooth. Supraclypeal area sculptured as on
clypeus. Face with punctures separated by 1
1.5 times a puncture width, integument between smooth. Vertex and gena with small
punctures separated by about a puncture
width, integument between smooth. Mesoscutum, scutellum, and metanotum impunctate and finely imbricate. Tegula with minute,
faint punctures separated by 12 times a
puncture width, integument between faintly
imbricate. Pleura impunctate and finely imbricate. Propodeum impunctate and imbricate. Terga and sterna imbricate.
Color dark brown or black without maculations. Wing membrane hyaline; veins
74
NO. 259
Figs. 4546. Holotype female of Ctenoplectrella grimaldii, new species. 45. Dorsal view of holotype. 46. Left lateral view of mesosomal dorsum. Scale bar 5 1 mm.

scopa composed of dense, long, erect, simple

setae.
MATERIAL: One specimen. Holotype. Female, B-JH 88 (AMNH) labeled: Baltic amber: Eocene, Kaliningrad, Yantarny // Holotype, Ctenoplectrella grimaldii Engel.
ETYMOLOGY: The specific epithet is a patronymic honoring Dr. David A. Grimaldi for
his friendship, advice, and assistance with
my paleontological studies.
Ctenoplectrella sp. indet.
MATERIAL: One specimen. Non-type. Female, Nr. 1616 (CCGG) labeled: Nr. 1616
// Ctenoplectrella sp. indet., det. M. S. Engel.
COMMENTS: It is possible that this speci-
2001
75
men represents another individual of C. viridiceps.

Ctenoplectrella sp. indet.
MATERIAL: One specimen. Non-type. Female, Nr. 3000 (GPUH) labeled: Nr. 3000
// Ctenoplectrella sp. indet., det. M. S. Engel.
Ctenoplectrella? sp. indet.
Megachile sp. Gerlach, 1989: 255. [misidentification]
MATERIAL: One specimen. Non-type. Female, Nr. G4669 (IMGP) labeled Reservierte Registriernummer, Megachile: G4669
[Gerlach handwriting] // Megachile sp. indet., Gerlach, J., 1987 // G4669, Ctenoplectrella sp. indet. (sensu Ckll.), det. M. S.
Engel, 1999, Geologisch-Palaontologisches
Institut und Museum, Gottingen.
COMMENTS: This specimen is poorly preserved and is mostly covered by Schimmel
except for portions of the compound eyes, an
exposed wing, apices of the legs; other structures dimly visible through thinner regions of
Schimmel. Those few visible features generally suggest a species of Ctenoplectrella
(i.e., an osmiine without eye setae, relatively
orthogonal second abscissa Rs to M, and
with an apparently narrow gena). The strongly arcuate 2rs-m suggests C. cockerelli or C.
grimaldii but further identification as to species is impossible. The somewhat larger body
size (ca. 8 mm) suggests that the specimen
may represent an otherwise unknown species
(if indeed it is a true Ctenoplectrella at all!).
The body size is closer to that of Glaesosmia.
Glaesosmia, new genus
TYPE SPECIES: Glaesosmia genalis Engel,
new species.
DIAGNOSIS: This genus is most similar to
Ctenoplectrella but differs by the short supraclypeal area, broad gena, angulate second
abscissa of Rs (fig. 47), and broad head (refer
to key to genera, above).
DESCRIPTION: Mandible with three lower
teeth along apical margin, upper tooth separated from lower two teeth by a short, gently
concave cutting-edge. Malar space short,
much shorter than basal mandibular width.
Clypeus flat; not extending below lower tan-

Glaesosmia genalis, new species. Scale bar 5 1
mm.
gent of compound eyes. Subantennal sutures

as long as antennal socket diameter. Compound eyes bare; compound eye as broad as
gena in lateral view; inner margins straight
and slightly converging below. F1 slightly
longer than F2; length of F2 approximately
equal to that of F3. Posterior margin of vertex gently concave. Preoccipital area rounded. Mesoscutum and scutellum not coarsely
sculptured; mesoscutal anterior border broadly rounded; median line moderately impressed; parapsidal lines moderately impressed and linear; tegula oval; scutellum
low, weakly convex, not overhanging metanotum; propodeum not strongly sculptured.
Claws with inner tooth, inner tooth shorter
than outer; arolium present; outer apex of
mesotibia without spine; two metatibial
spurs, spurs serrate. Basal vein strongly arcuate; second abscissa Rs forming an acute
angle with M (fig. 47). No apparent maculations on integument.
is a combination of glaesum (Latin, meaning
amber) and the generic name Osmia. The
name is feminine.
Glaesosmia genalis, new species
Figure 47

much wider than long (length 2.05 mm,
1.85 mm. Basal vein basad cu-a by vein
76
width; second abscissa Rs basad 1m-cu by

10 times vein width; 2rs-m distad 2m-cu by
vein width, 2rs-m doubly arcuate; second
submarginal cell slightly longer than first
submarginal cell, lower apical margin of second submarginal cell extending apically; 10
distal hamuli, arranged in a single, evenly
spaced series.
Clypeus with small punctures separated by
a puncture width or less, integument between
smooth. Supraclypeal area and face with
small punctures separated by a puncture
Vertex and gena with sculpturing as on face,
although punctures slightly more widely
spaced, separated by about 0.51.5 times a
Mesoscutum with small punctures separated
by less than a puncture width, integument between smooth. Tegula with minute punctures
separated by a puncture width or less, integument between smooth. Scutellum sculptured
as on mesoscutum. Metanotum rugulose.
Preepisternal area (i.e., anterior-facing surface anterior to omaulus) essentially impunctate and smooth except for a few, exceedingly faint, widely spaced, coarse punctures.
Mesepisternum with small punctures separated a puncture width, integument between
smooth, punctures becoming faint on posterior third. Metepisternum impunctate and
smooth. Basal area and posterior surface of
propodeum finely imbricate and impunctate,
lateral surface finely imbricate with a few,
sparse, faint, small punctures. T1T2 impunctate and smooth; remaining terga with
coarse, faint punctures separated by a puncture width, integument between smooth; sternal integument obscured.
Coloration brown, without maculations.
Wing membrane hyaline except between
C1Sc and R slightly fuscous in some specimens; veins dark brown.
Clypeus with minute, appressed, simple
setae not obscuring integument. Face, clypeus, supraclypeal area, gena, and postgena
with scattered, short, erect or suberect, simple setae. Mesoscutum and scutellum with
scattered, short, simple setae. Pleural pubescence as described for mesoscutum. Basal
area of propodeum without pubescence; lateral and posterior surfaces with pubescence
as described for mesoscutum. Pubescence of
NO. 259
legs generally simple and short; inner surface

of metafemur without pubescence, outer, anterior, and posterior surfaces with short,
branched setae; metatibia with scattered, simple, moderate-length setae, a few with minute branches. Central discs of metasomal
terga with sparse, minute, simple, appressed
setae, setae longer and erect to suberect on
lateral borders; sternal scopa composed of
dense, long, erect, simple setae.
MATERIAL: One specimen. Holotype. Female, B-JH 80 (AMNH) labeled: Baltic amber: Eocene, Kaliningrad, Yantarny // Holotype, Glaesosmia genalis Engel.
ETYMOLOGY: The specific epithet is a reference to the enlarged gena (L. gena, meaning cheek).
Family APIDAE Latreille
DIAGNOSIS: Long-tongued bees lacking a
strong metasomal scopa in nonparasitic
forms and with the clypeus bent posteriorly
lateral to the narrowed labral articulation.
DESCRIPTION: Single subantennal suture;
suture meeting lower margin of antennal
socket. Facial foveae absent. Lower lateral
margins of clypeus bent posteriorly on either
side of labrum. Flabellum present; glossa
acute; labial palpus with first two segments
flattened, sheath-like, and elongate; submentum strongly V-shaped and sclerotized, submental arms articulating with cardines
slightly above cardo-stipital articulations.
Galeal comb absent; stipital comb and concavity present; basistiptial process elongate.
Preepisternal groove absent. Mesocoxa entirely exposed. Metabasitibial frequently present. Jugal lobe short. Strong metasomal scopa absent; scopa variously formed on metafemur and metatibia, metatibia sometimes
developed into a corbicula (in nonparasitic
females). Metapostnotum setose. Pygidial
plate and fimbria of female typically present.
Key to Subfamilies of Apidae
in Baltic Amber
1. Metatibia modified into corbicula; procoxa
slightly wider than long or as long as wide;
rastellum present (corbiculate Apinae) . . .
. . . . . . . . . . . . . . . . . . . . . Apinae Latreille
Metatibia not modified into corbicula, typical
scopa present but reduced; procoxa much
wider than long; rastellum absent . . . . .
. . . . . . . . . . . . . . . . . Xylocopinae Latreille
2001
Subfamily XYLOCOPINAE Latreille

Xylocopae Latreille 1802b: 379. Type genus: Xylocopa Latreille, 1802b.
D IAGNOSIS : Apid bees with distinctly

broadened procoxae and a reduced metatibial
scopa.
DESCRIPTION: Flabellum set at apex of bare
shank (Michener and Brooks, 1984). Procoxae wider than long; metatibial scopa reduced
(in nonparasitic females), not developed into
corbicula.
COMMENTS: This morphologically diverse
subfamily of bees contains the familiar carpenter bees of the genera Xylocopa and Ceratina. The subfamily is partitioned into four
tribes; the Allodapini of the Old World tropics and south temperate regions, the Manueliini of southern South America, and the
cosmopolitan tribes Ceratinini (small carpenter bees of the genera Ceratina and Megaceratina; Pithitis is considered a subgenus of
Ceratina by Michener, 2000a) and Xylocopini (the large carpenter bees of the genus
Xylocopa; including Proxylocopa and Lestis
according to Minckley, 1998). A key to these
four tribes was presented by Sakagami and
Michener (1987) and Michener (2000a). Individuals of Ceratinini and Xylocopini have
been discovered in various fossil deposits
[Heer, 1849, 1865 (as Bombus); Cockerell,
1906; Statz, 1936; Zeuner, 1938; Zhang,
1990; Zhang et al., 1994; Engel, in press-a];
however, no Xylocopinae have previously
been recorded as amber inclusions. These
fossils are also the oldest record of the subfamily. A cladistic study of the subfamily is
presented below based on exemplar genera
(see Cladistic Analyses), while phylogenetic
studies on the relationships within Xylocopini have been undertaken by Minckley
(1998) and for Allodapini by Reyes (1998).
Species of Manueliini were reviewed by
Daly et al. (1987) but relationships were not
explored cladistically. No cladogram has yet
been produced for Ceratinini.
BOREALLODAPINI, New Tribe
TYPE GENUS: Boreallodape Engel, new genus.
DIAGNOSIS: Among xylocopine bees the
boreallodapines are most closely related to
77
the tribes Ceratinini and Allodapini; all three

have the abruptly narrowed mandible (fig.
54), have the anterior tentorial pit situation
near the middle of the clypeus (fig. 49), and
have flattened labral surfaces (Xylocopini
and Manueliini have tapering mandibles, the
anterior tentorial pits situated well above the
middle of the clypeus, and basal labral elevations). From the Allodapini the new tribe
differs by having an elongate supraclypeal
area (longer than the diameter of the antennal
sockets) (fig. 49), unflattened distal metasomal terga (figs. 48, 53, 55), the lateral margins of the clypeus concave (not biconvex)
(fig. 49), and in the hind wing the first abscissa of M is more than three-quarters the
length of the second abscissa of M1Cu (fig.
51). From the Ceratinini the boreallodapines
can be distinguished by having only two submarginal cells (fig. 50) and lacking a metabasitibial plate.
DESCRIPTION: Mandible abruptly narrowed
near midpoint (fig. 54). Anterior tentorial pits
near (frequently slightly above) midpoint of
clypeus; lateral margins of clypeus strongly
concave (fig. 49). Subantennal sutures longer
than antennal socket diameter (i.e., supraclypeal area more elongate than that in Allodapini in which the subantennal sutures are
shorter than the diameter of the antennal socket) (fig. 49). Scape short, not reaching to median ocellus; F1 less than combined lengths
of F21F3. Prosternum with short basisternum
and anterior process, lateral processes slightly
elongate, without constriction between basisternum and furcasternum (figs. 5658); furcasternum elongate with strong medial groove
from apophyseal pit that extends slightly less
than three-quarters of total furcasternal length.
Claw with strong inner tooth; arolium large
(fig. 52). Forewing membrane pubescent, not
papillate; marginal cell apex rounded on wing
margin (figs. 50, 55); pterostigma of moderate
size (not obsolete as in Xylocopini); prestigma short; two submarginal cells (figs. 50, 55);
hind wing with first abscissa of M more than
three-quarters length of second abscissa of
M1Cu (fig. 51); jugal lobe one-half length of
vannal lobe. Metabasitibial plate absent. Distal metasomal terga distinctly not flattened
dorsally (in Allodapini the distal three metasomal terga are flattened dorsally) (fig. 55);
pygidial fimbria absent; pygidial plate absent.
78
Fig. 48.
NO. 259
Right lateral view of holotype female of Boreallodape baltica, new species.
Boreallodape, new genus

TYPE SPECIES: Boreallodape baltica Engel,
new species.
DESCRIPTION: As for the tribe with the following additions: Clypeus not protuberant in
lateral view, gently and slightly curved,
slightly extending below lower tangent of
compound eyes, with scattered, faint punctures; lateroclypeal carina absent; upper clypeal margin relatively straight. Antennal sockets slightly above level of middle of compound eyes. Frons not protuberant. Vertex
Fig. 49. Frontal view of head of holotype female of Boreallodape baltica, new species. Scale
bar 5 0.5 mm.
behind ocelli strongly concave, vertex relatively short. Preoccipital area rounded. Gena
narrower than compound eye in lateral view.
Malar space short, much less than basal mandibular width. Mandible bidentate (fig. 54).
Mesoscutum broadly rounded anteriorly; median and parapsidal lines moderately impressed. Tibial combs absent; coxae without
spines or carinae. Wing membrane hyaline;
veins dark brown; basal vein strongly curved
(figs. 50, 55); pterostigma well developed,
margin inside marginal cell slightly convex;
apex of marginal cell acutely rounded, set off
from wing margin by width of vein at most;
second submarginal cell shorter than first;
1rs-m basad of 1m-cu; five distal hamuli arranged in a single, evenly spaced series.
ETYMOLOGY: The new genus-group name is
derived from boreas (Greek, meaning
north) and Allodape, type genus of the xylocopine tribe Allodapini. The name is a combined reference to the resemblance of species
of this tribe to those of their sister tribe Allodapini and to the fact that these fossils are
at the highest latitude recorded for any allodapine-like bee. The name is feminine.
COMMENTS: Species of Boreallodape su-
2001
79
Figs. 5051. Wings of holotype female of Boreallodape baltica, new species. 50. Forewing. 51.
Hind wing. Scale bar 5 0.5 mm.
perficially resemble the living allodapine genus Braunsapis but can be immediately separated from the latter (as well as all other
Allodapini) by the tribal characteristics.
Key to Species of BOREALLODAPE
1. Pleura strongly punctate, punctures well-defined, small and separated by a puncture
width or less; tergal punctation closely
spaced, separated by a puncture width at
most; basal area of propodeum variable;
larger species (body length more than 4
mm) . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2
Pleura impunctate; tergal punctation minute
and sparse, puncture separated by 13
times a puncture width; basal area of propodeum glabrous; smaller species (body
length less than 4 mm) . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . B. mollyae, n. sp.
2. Lower half of hypoepimeral area impunctate;
punctures of mesepisternum not nearly

contiguous, separated by 11.5 times a
puncture width of smooth integument . . .
. . . . . . . . . . . . . . . . . . . . B. baltica, n. sp.
Hypoepimeral area uniformly punctured;
punctures of mesepisternum nearly contiguous, when separated, then separated by
less than a puncture width . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . B. striebichi, n. sp.
Boreallodape baltica, new species

Figures 4852
Plates 4a,b, 5b

B. striebichi but can be distinguished by the
glabrous basal area of the propodeum, the
more well-spaced pleural punctures (separated by smooth integument), and the presence
80
Fig. 52. Claw and arolium of holotype female

of Boreallodape baltica, new species. Scale bar
5 0.25 mm.
of a distinct impunctate region on the hypoepimeral area.

0.08 mm. Intertegular distance 0.80 mm.
Basal vein distad cu-a by two times vein
four times vein width; 2rs-m relatively
straight, distad 2m-cu by two times vein
width; first submarginal cell longer than second submarginal cell.
24 times a puncture width, integument between smooth. Clypeus with exceedingly
faint, small punctures sparsely scattered. Supraclypeal area sculptured as on clypeus. Remainder of face with sparse, small punctures,
integument between smooth, punctures become more closely packed (separated by two
times a puncture width) toward upper part of
face and vertex, integument between smooth;
small impunctate zone lateral to and bordering
lateral ocellus, impunctate zone less than 1OD
in width and length. Gena with small punctures separated by 12 times a puncture width,
integument between smooth; postgena sculptured as on gena. Pronotum with small punctures separated by 12 times a puncture width,
NO. 259
width. Tegula with small, faint punctures on

inner two-thirds, punctures separated by 12
times a puncture width. Scutellum with small
width, integument between smooth. Metanotum with minute punctures separated by a
smooth. Pleura strongly punctured, punctures
approximately twice the diameter of those on
mesoscutum, punctures separated by 11.5
times a puncture width except lower half of
hypoepimeral area impunctate, integument
between smooth. Metepisternum sculptured
like mesepisternum except punctures separated by a puncture width or less and slightly
smaller. Basal area of propodeum, metapostnotum, and posterior surface of propodeum
impunctate and smooth; lateral surface of propodeum with small punctures separated by a
First metasomal tergum impunctate; T2 with
coarse, faint punctures separated by a puncture width or less except central disc impunctate and punctures bordering this area more
widely separated and more faint; remaining
terga with coarse, faint punctures separated by
a puncture width or less, without impunctate
zone on central discs.
Integumental coloration not preserved.
Wing membrane hyaline; veins strong and
dark brown.
Setae simple, minute or short, and sparse
except those of sterna erect, short and more
dense.
MATERIAL: Eight specimens. Holotype. Female, B-JH 152 (AMNH) labeled Baltic amber: Eocene, Kaliningrad, Yantarny // Holotype, Boreallodape baltica Engel. Paratype. Female, Nr. 15: 1967 (ZMUC) labeled:
Apidae, C. V. Henningsen, 15, 1967 //
Paratype, Boreallodape striebichi Engel.
This specimen is missing the very apex of the
head (right mandible, lateral two-thirds of labrum, apex of right compound eye, and lateral
third of clypeus), which is right at the surface
of the amber. There are also some thin layers
of Schimmel on the bee.
Paratype. Female, Nr. 810 (CDTG) labeled: Nr. 810 // Paratype, Boreallodape
baltica Engel.
Paratype. Female, Nr. 873 (CHFG) labeled: Nr. 873, Balt. Bernstein // Paratype, Boreallodape baltica Engel.
2001
Fig. 53.
81
Dorsal view of holotype female of Boreallodape mollyae, new species.
Paratype. Female, Nr. 70 (CJDL) labeled:
Nr. 70 // Paratype, Boreallodape baltica
Engel.
Yantarny // Boreallodape baltica? Engel,
det. M. S. Engel.
ETYMOLOGY: The specific epithet is a reference to the Baltic region where the amber
originates.
Boreallodape mollyae, new species
Figures 53, 54
Plates 4e, 5a
DIAGNOSIS: This is the smallest species of

the genus. Boreallodape mollyae can be recognized by the combination of impunctate
pleural integument (unique for the genus)
and minute and relatively sparse metasomal
punctation.

13 times a puncture width, integument between smooth. Clypeus with small punctures
separated by 23 times a puncture width. Supraclypeal area sculptured as on clypeus. Remainder of face with sparse, small punctures,
integument between smooth, punctures becoming gradually more closely spaced again
(separated by 24 times a puncture width) on
upper part of face and vertex, integument between smooth. Gena with minute punctures
separated by 34 times a puncture width, integument between smooth; postgena with
Pronotum impunctate, faintly imbricate. Mesoscutum with small punctures separated by
82
NO. 259
MATERIAL: Six specimens. Holotype. Female, B-JH 81 (AMNH) labeled: Baltic amber: Eocene, Kaliningrad, Yantarny // Holotype, Boreallodape mollyae Engel.
Paratypes. Five females, Nr. 19948
(ZMPA) labeled: Muzeum Ziemi w Warszawie, 19948, Baltic amber with bees, Apidae, from Baltic beach in Stupsk, 1983 //
Baltic Amber: Eocene, Nr. 19948, Muzeum
Ziemi Pan, M. S. Engel study // Paratype,
Boreallodape mollyae Engel.
ETYMOLOGY: The specific epithet is a patronymic in loving honor of Ms. Molly G.
Rightmyer, brilliant hymenopterist and gifted
artist, who executed the numerous magnificent illustrations presented herein.
Boreallodape striebichi, new species
Figures 5558
Plate 4c,d,f
Fig. 54. Frontoventral oblique view of head

of holotype female of Boreallodape mollyae, new
species. Scale bar 5 0.5 mm.

smooth. Tegula with minute punctures, punctures separated by 12 times a puncture
width. Scutellum sculptured as on mesoscutum. Metanotum with minute punctures separated by 23 times a puncture width, integument between smooth. Mesepisternum impunctate and smooth except for a few exceedingly faint punctures sparsely scattered,
hypoepimeral area impunctate. Metepisternum impunctate. Propodeum impunctate and
smooth. T1T2 impunctate; T3T5 with minute punctures separated by 23 times a
puncture width, integument between smooth;
T6 with minute punctures separated by 12
smooth; sterna imbricate.
Where preserved (mesoscutum, scutellum,
metasoma) integument dark brown and shining with reddish-brown highlights. Wing
membrane hyaline; veins dark brown.
except those of sterna erect, short and more
dense.

the type species for the genus, B. baltica, but
differs in the punctation of the pleura and
sculpturing of the propodeum (refer to Diagnosis for B. baltica).
24 times a puncture width, integument between smooth. Clypeus with faint, small
punctures sparsely separated by 13 times a
puncture width. Supraclypeal area and face
sculptured as on clypeus; punctures becoming
more closely packed (separated by less than a
puncture width) toward upper part of face and
vertex, integument between smooth; small impunctate zone lateral to and bordering lateral
ocellus, impunctate zone less than 1OD in
width and length. Gena with small punctures
separated by less than a puncture width, integument between finely imbricate; postgena
2001
Fig. 55.
Right dorsolateral habitus of holotype female of Boreallodape striebichi, new species.
83
84
NO. 259
Fig. 56. Scanning electron micrograph of paratype female of Boreallodape striebichi, new species;
partial specimen preserved at amber surface of B-BS 153 (AMNH).
sculptured as on gena. Pronotum with small

less, integument between finely imbricate.
Mesoscutum with small punctures separated
by a puncture width, integument between
smooth. Tegula with minute, faint punctures
on inner two-thirds, punctures separated by
two times a puncture width. Scutellum sculptured as on mesoscutum. Metanotum with minute punctures separated by a puncture width
or less, integument between smooth. Pleura
strongly punctured, punctures separated by
less than a puncture width or more often contiguous, hypoepimeral area without impunctate zone, integument between (where evident) smooth. Metepisternum sculptured as on
mesepisternum. Basal area of propodeum,
metapostnotum, and posterior surface of propodeum imbricate; lateral surface of propodeum with small punctures separated by less
than a puncture width, integument between
faintly imbricate. First metasomal tergum impunctate; T2 with coarse, faint punctures separated by a puncture width or less except
punctures of central disc more widely separated and faint, without impunctate area; remaining terga with coarse, faint punctures
separated by a puncture width or less.
Integumental coloration dark brown where
preserved. On some individuals yellow markings are easily apparent along the inner margin of the compound eyes below the level of
the anterior tentorial pits as well as small areas on the lateral apical borders of the clypeus. Wing membrane hyaline; veins dark
brown.
except those of sterna erect, short, and more
dense.
MATERIAL: Fourteen specimens. Holotype.
Female (pl. 4c, 4d), B-BS 153 (AMNH) labeled: Baltic amber: Eocene, Kaliningrad,
2001
85
Fig. 57. Scanning electron micrograph of paratype female of Boreallodape striebichi, new species.
Magnified view of prosternum from figure 56.
Yantarny // Holotype, Boreallodape striebichi Engel. There are several specimens in

this single piece (designated as paratypes below). The holotype is the single individual
depicted in plate 4c, 4d. There is some pyrrhite on the specimen (e.g., on supraclypeal
area) that makes the sculpturing difficult to
interpret in some areas.
Paratypes. Eight whole females and five
partial females, B-BS 153 (AMNH): in the
same block of amber as the holotype.
ETYMOLOGY: The specific epithet is a patronymic honoring Herr Bernhard Striebich,
Buxtehude (Germany), who generously donated the holotype and its associated paratypes to the AMNH.
COMMENTS: The shape of the clypeus in
this species appears to be slightly different
from that of other Boreallodape. The lateral
margins of the clypeus are not as strongly
concave and the portion of the clypeus above
the bend in the lateral margins and above the

anterior tentorial pits is shorter (thereby the
supraclypeal area is correspondingly longer
than in the other species). Thus, the clypeus
is more plesiomorphic (more closely approximating that shape seen in Manueliini) than
other Boreallodape species. I presently interpret this species as the basalmost species of
Boreallodape.
Boreallodape sp. indet.
MATERIAL: One specimen. Non-type. Female, B-W 163 (AMNH) labeled: Baltic
Boreallodape sp. indet., det. M. S. Engel.
COMMENTS: This specimen is mostly covered by debris, and although the wing venation, metasoma, and legs can be examined
easily, the remainder of the body cannot be
seen clearly.
86
NO. 259
Fig. 58. Scanning electron micrograph of paratype female of Boreallodape striebichi, new species.
Magnified and slightly oblique view of prosternum.
Subfamily APINAE Latreille

Corbiculate Apinae
Apiariae Latreille, 1802a: 373. Type genus: Apis
Linnaeus, 1758.
DIAGNOSIS: The corbiculate apines, as the

name implies, can be immediately recognized by the modification of the metatibia
into a corbicula in nonparasitic females and
workers. The presence of a rastellum, an expanded mandibular pollex, as well as the absence of the metabasitibial plate and pygidial
plate also serve to distinguish this clade from
other apines as well as the Xylocopinae (the
only other subfamily of Apidae presently
known in Baltic amber). At present, only the
corbiculate tribes of Apinae are known from
Baltic amber.
DESCRIPTION: Mandibular pollex expanded.
Metatibia modified into corbicula; metabasitibial plate absent; inner apical margin of me-
tatibia with rastellum; auricle frequently present (except in parasitic forms and Meliponini). Wings uniformly setose. Prepygidial
fimbria and pygidial plate absent.
COMMENTS: The name corbiculate bees,
or Corbiculata, was first coined by Shuckard
(1866: 165) for this specialized group of apine bees. This distinctive clade was once
considered as the family Apidae (e.g., Michener, 1965, 1990) with the other apids relegated to the paraphyletic family Anthophoridae. The recent study of long-tongued bee
phylogeny by Roig-Alsina and Michener
(1993) resurrected the expanded concept of
Apidae as it was employed by Michener
(1944) and the corbiculate bees were incorporated into an expanded subfamily Apinae
without a separate, formal rank. Should a formal rank be useful for recognizing the corbiculate apines, the supertribal name Apiti
could be applied. Presently, an apine clade
2001
containing the non-corbiculate tribes Anthophorini, Centridini, Melectini, and Ericrocidini6 is believed to be the closest relative of
the corbiculate Apinae (refer to fig. 123).
This is the most common group of bees in
Baltic amber. Three of the living tribes are
eusocial to varying degrees and the phylogenetic position of the extinct corbiculate
groups suggests that they too were eusocial
(refer to sections on Cladistic Analyses, below). This social lifestyle perhaps can account for their abundance in amber in comparison to other groups. Living corbiculate
bees typically collect resins and it is possible
that the fossil species described below did as
well, thus subjecting them more frequently
to entrapment and fossilization.
The corbiculate clade consists of four extant tribes, briefly summarized as follows:
The Apini (honey bees) are originally an Old
World group but have been spread throughout the world by humans for agricultural purposes. Presently, fossils of Apini are only
known from as far back as the early Oligocene (Engel, 1998c, 1999c). The Euglossini
(orchid bees) are strictly neotropical with
two species in Miocene amber from the Dominican Republic (Engel, 1999b) and one
extant species occurring in South American
copal (Engel, personal obs.; Ross, 1998). The
bumble bees, tribe Bombini, are distributed
throughout the western hemisphere, the Palearctic, and the Oriental regions. The report
of a bumble bee from tropical Africa (Tkalcu, 1966) is actually based upon on an introduced species from South America (Sakagami, 1976; Michener, 1979, 1990; Williams,
1998). Numerous fossil bumble bees are
known [Heer, 1867; Unger, 1867; Novak,
1877; Cockerell, 1906 (as Calyptapis), 1931;
Piton, 1940 (as Probombus); Zhang, 1990;
Zhang et al., 1994; Rasnitsyn and Michener,
1991; Riou, 1999] but no true bombine has
yet been discovered in amber. The report of
a bombine in Paleocene amber from France
(In Grimaldi, 1999) has yet to be confirmed;
it is possible that this fossil is an electrobom6 Despite that shown in figure 123, relationships
among these four non-corbiculate tribes are somewhat
uncertain and nodes are not strongly supported. It is very
likely that, once resolved, Anthophorini and Melectini
will be sister groups and together sister to a monophyletic Centridini 1 Ericrocidini clade.
87
bine (if a small jugal lobe is present in the

hind wing). Lastly, the stingless bees, tribe
Meliponini, are diverse pantropically. Species are known in Late Cretaceous New Jersey amber (Michener and Grimaldi, 1988a,
1988b; Engel, 2000b), Miocene Dominican
amber (Wille and Chandler, 1964; Michener,
1982; Camargo et al., 2000), Oligocene-Miocene Mexican amber (Wille, 1959), Colombian copal (Engel, personal obs.), Burmese
copal [Cockerell, 1921 (not Miocene amber!)], Sicilian amber [Tosi, 1896 (perhaps
copal?)], and African copal [Engel, personal
obs.; Stuckenberg, 1975 (not Baltic amber!);
Zeuner and Manning, 1976; Wille, 1977]. Of
all of these Recent tribes, only the meliponines are presently known in Baltic amber.
Key to Tribes of Corbiculate Apinae
in Baltic Amber
1. Distal wing venation complete (e.g., figs. 62,
68, 75, 77); marginal cell closed at apex;
metatibial spurs present (fig. 71); auricle
present (e.g., figs. 81, 90, 106); supra-alar
carina present; claws with inner tooth (e.g.,
figs. 61, 108) . . . . . . . . . . . . . . . . . . . . . 2
Distal wing venation incomplete (figs. 111,
113); marginal cell open at apex; metatibial
spurs absent; auricle absent (fig. 114; supra-alar carina absent; claws simple (fig.
115) . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
. . Meliponini Lepeletier de Saint Fargeau
2. Outer grooves of mandible present; hamuli
not reduced, 10 or more (fig. 63); 1m-cu
not strongly angulate (e.g., figs. 62, 66);
one or two metatibial spurs present; robust,
Bombus- or Apis-like bees . . . . . . . . . . . 3
Outer grooves of mandible absent; hamuli reduced in number, less than 10 (figs. 92,
104, 110); 1m-cu short and strongly angulate (figs. 91, 103, 109); single metatibial
spur present (figs. 90, 102, 107); frequently
small, Trigona-like bees . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . Melikertini, n. tribe
3. Alar papillae absent (figs. 6668, 7377, 85);
outer mandibular grooves reduced but present; one metatibial spur present (figs. 71,
83); marginal cell either weakly truncate or
appendiculate . . . . . . . . Electrapini Engel
Alar papillae present (figs. 62, 63); outer
mandibular grooves present and strong;
two metatibial spurs present; marginal cell
narrowly rounded at apex, not truncate or
appendiculate . . . Electrobombini, n. tribe
88
ELECTROBOMBINI, New Tribe

TYPE GENUS: Electrobombus Engel, new
genus.
DIAGNOSIS: Robust Bombus-like bees that
resemble in almost every respect the tribe
Bombini; however, the presence of a distinct
jugal lobe at the base of the hind wing, a
well-developed pterostigma that is much
larger than the prestigma, and a strongly developed arolium serve to distinguish the
electrobombines from bombines. Electrobombini can be separated from primitive
Electrapini by the presence of strongly developed outer mandibular grooves and two
elongate metatibial spurs.
DESCRIPTION: Large (ca. 18 mm long), robust, densely pubescent bees. Mandible with
strong outer mandibular grooves present. Labral width two times length (fig. 59). Clypeus
slightly convex and weakly protuberant in
lateral view. Compound eyes bare. Supraalar
carina present; scutellum broadly rounded
posteriorly and projecting over metanotum
and (perhaps?) propodeum. Claws of female
with inner tooth; arolium present and strong
(fig. 61); two metatibial spurs present; malus
of strigilis with short anterior velum in addition to primary ventral velum; metabasitarsus with distinct auricle at base (fig. 60); metatibia without penicillum (fig. 60). Distal venation of forewing strong and present; marginal cell large and narrowly rounded at
apex, slightly offset from wing margin, not
truncate or appendiculate, cell longer than
distance from its apex to wing apex (fig. 62);
pterostigma present and moderately sized,
much longer than prestigma, r-rs arising
slightly prior to its midpoint, margin within
marginal cell straight; 1m-cu relatively
straight; hind wing with small but distinct jugal lobe, lobe broadly incised (fig. 63); hamuli numerous (15 in a single series for Electrobombus samlandensis); wing membrane
papillate in outer margins (figs. 62, 63). Sting
not reduced, not barbed.
Electrobombus, new genus
TYPE SPECIES: Electrobombus samlandensis Engel, new species.
DESCRIPTION: Mandible with a few weak
apical teeth. Malar space short, shorter than
NO. 259
basal mandibular width. Epistomal sulcus

forming obtuse angle (fig. 59). Inner margins
of compound eyes slightly converging below, not emarginate. F1 longer than F23
combined; F3 longer than F2. Preoccipital
ridge rounded. Mesoscutal anterior border
broadly rounded; scutellum projecting over
metanotum and basal area of propodeum.
Basal area of propodeum apparently strongly
declivitous. Keirotrichiate field not elevated;
metatibial spurs serrate; auricular basket
sparse, setae short; rastellum strong and
formed of stiff setae; metabasitarsus twice as
long as wide. Margin of pterostigma within
marginal cell straight; marginal cell apex offset from wing margin by slightly less than
pterostigma width, apex rounded, not appendiculate.
is a combination of elektron (Greek, meaning
amber) and the generic name Bombus
(originally taken from the Greek bombos,
meaning buzzing, which itself is related to
bombylios, meaning bumble bee).
Electrobombus samlandensis, new species
Figures 5963
Plate 5c, e

longer than wide (length 3.40 mm, width
3.24 mm). Upper interorbital distance 1.50
mm; lower interorbital distance 1.45 mm. Interocellar distance indeterminate (owing to
bubble that obscures the position of one lateral ocellus); ocellocular distance 0.35 mm;
median to lateral ocellus 0.20 mm. Intertegular distance 3.75 mm. Basal vein basad cua by three times vein width; 1rs-m distad by
1m-cu by seven times vein width; 2rs-m distad 2m-cu by vein width; first submarginal
cell shorter than second and third combined;
length of anterior border of second submarginal cell one-half that of posterior border;
length of anterior border of third submarginal
cell one-half of that of posterior border, 1.5
times length of anterior border of second
submarginal cell; 15 hamuli, arranged in a
Integument, where visible between dense
pubescence of head, mesosoma, and meta-
2001
89
Figs. 5961. Holotype female of Electrobombus samlandensis, new species. 59. Oblique frontal
view of head. 60. Outer surface of metatibia and basal portion of metabasitarsus. 61. Claw and arolium.
Scale bars 5 1 mm (a 5 fig. 60; b 5 fig. 59; c 5 fig. 61).
soma, smooth and impunctate except face,

clypeus, and supraclypeal area with small,
faint punctures separated by a puncture width
or less, tegula with microscopic punctures
separated by 23 times a puncture width, and
terga and sterna imbricate.
Coloration of head and mesosoma black;
metasoma, antennae, and legs dark brown.
dark brown.
Pubescence generally yellowish-white.
Mandible with dense, minute setae in apical
halves of outer grooves. Labrum with widely
scattered, short, simple, erect setae. Clypeus
with scattered, erect setae, such setae slightly
more numerous along lateral borders. Face
with dense, short, appressed, plumose setae
mostly obscuring integument intermixed
with longer, erect to suberect, plumose setae.
Vertex with long, plumose, erect setae. Gena
with pubescence as described for vertex except setae with only a few, minute branches.
Mesosomal (dorsal, lateral, and ventral) pubescence long, feathery plumose, and exceedingly dense, mostly obscuring integument. Tegula with rather sparse, short, appressed setae not obscuring integument.
Long corbicular setae with numerous, minute

branches intermixed with long, simple setae.
Terga and sterna with minutely branched,
dense, moderate-length, subappressed setae
that partly obscure the integument.
MATERIAL: Two specimens. Holotype. Female (caste indeterminate), B-JH 94
Kaliningrad, Yantarny // Holotype, Electrobombus samlandensis Engel.
Non-type. Female (caste indeterminate),
Nr. 500 (CJDL) labeled: Nr. 500 // Electrobombus samlandensis?, det. M. S. Engel. This individual is entirely covered in
Schimmel. Those few characters that can be
determined suggest that the specimen is a
second individual of E. samlandensis but this
will perhaps never be known with certainty.
The specimen is even larger than the holotype and is the largest bee specimen known
from Baltic amber (total body length 22
mm).
ETYMOLOGY: The specific epithet is a reference to the Samland Peninsula where most
Baltic amber has been uncovered.
COMMENTS: This is presently the largest
90
NO. 259
Figs. 6263. Wings of holotype female of Electrobombus samlandensis, new species. 62. Forewing.
63. Hind wing. Scale bar 5 1 mm.
bee species preserved in amber (of any deposit!).

TRIBE ELECTRAPINI ENGEL,
NOMEN TRANSLATUM
Electrapina Engel 1998a: 99. Type genus: Electrapis Cockerell, 1908b.
DIAGNOSIS: The electrapine bees are a heterogeneous group of three genera resembling
either bumble bees (Bombini) or, to a much
lesser degree, honey bees (Apini). From the
former tribe electrapines differ by the presence of a jugal lobe, absence of alar papillae,
truncated or appendiculate marginal cell
apex, reduction of the outer mandibular
grooves, and presence of only one, reduced
metatibial spur. From the Apini, the electrapines differ by the deep incision demarking
the jugal lobe, the presence of outer mandibular grooves, the single metatibial spur (entirely absent in Apini), the absence of long
eye setae (some microscopic setae are present in many specimens as is typical for
many bees), and the shape of the marginal
and submarginal cells in the forewing,
among other characters.
DESCRIPTION: Moderately sized (ca. 5.516
mm long), robust, densely pubescent bees.
Mandible with weakened outer mandibular
grooves. Labral width 23 times length.
Clypeus slightly convex and weakly protuberant in lateral view or flat (in Thaumastobombus). Compound eyes bare or with
sparse, microscopic setae. Supraalar carina
present; scutellum broadly rounded posteriorly and variously produced (ranging from
projecting over metanotum and propodeum
2001
to over metanotum only). Claws of female

with inner tooth (e.g., figs. 72, 83); arolium
strong and present; single, reduced metatibial
spur present; malus of strigilis with short,
thickened anterior prong in addition to primary ventral velum (e.g., fig. 82); metabasitarsus with distinct auricle at base; metatibia
without penicillum. Distal venation of forewing strong and present; marginal cell large
and narrowly rounded at apex, slightly offset
from wing margin, truncate or feebly appendiculate, cell longer than distance from its
apex to wing apex; pterostigma present and
short to moderately sized, much longer than
prestigma, r-rs arising near to just after midpoint, margin within marginal cell variously
produced; 1m-cu relatively straight; hind
wing with distinct jugal lobe, lobe broadly
and deeply incised; hamuli numerous or reduced (numerous in Electrapis and Protobombus, reduced in Thaumastobombus);
wing membrane without alar papillae. Sting
not reduced; with or without microscopic
barbs.
COMMENTS: There has been much confusion over the identity of Electrapis and its
tribal position among the corbiculate bees.
Recently I erected the Electrapina as a subtribe of Apini to accommodate the nominate
genus as it had been conceived by more recent authors (e.g., Zeuner and Manning,
1976; Kelner-Pillault, 1970a; Engel, 1998a)
but my recent revelations over the true identity and nature of the type species (see below
in Comments under Electrapis and E. meliponoides) have resulted in a radical reinterpretation of the group. Those bees believed
to be Electrapis s.l. in recent decades, and
thereby somewhat intermediate in character
between Apini and Meliponini are, in fact,
members of the Melikertini (treated below).
The electrapines are, in fact, intermediate in
some respects between bombines and the
Apini 1 Meliponini clade (see Cladistic
Analyses, below). Cockerells (1908b,
1909b, 1909c) notes on the genus, as well as
the genus Protobombus, clearly indicate
overall Bombus-like bees (although some
have a definite Apis-like habitus) and provide
the necessary characters to associate his species with the specimens described below as
Electrapis and Protobombus. Other taxa (i.e.,
those thought to resemble meliponines) at
91
TABLE 8
Hierarchical Classification of Tribe

Electrapini
one time placed in or near Electrapis by Kelner-Pillault (1970a), Manning (1960), Zeuner
and Manning (1976), as well as myself (Engel, 1998a), are reassigned to the Melikertini
or in a few instances to other electrapine genera following the results of the cladistic analysis presented below.
The tribe Electrapini, even after the removal of Melikertes and Roussyana, may
still be paraphyletic with respect to an Apini
1 Meliponini 1 Melikertini clade. Among
the three genera of electrapines known, Electrapis is the most plesiomorphic and resembles not only Electrobombus (from which it
differs most significantly by the tribal characters mentioned above) but living bombines
as well. Protobombus appears somewhat intermediate between Electrapis and Thaumastobombus, the latter appearing most similar
to the higher corbiculates (i.e., Apini, Meliponini, and Melikertini). However, there is
presently not enough cladistic information to
resolve the issue, and this scheme is presented only as a working hypothesis upon
which future studies can build. It is appropriate to pose the possibility that electrapines
may represent a stem-group lineage from
which the higher corbiculates originated. Table 8 summarizes the present classification of
Electrapini.
92
Fig. 64.
NO. 259
Frontal view of head of female Electrapis martialis (Cockerell). Scale bar 5 1 mm.
Key to Genera of Electrapini

1. Metabasitarsus elongate, 1.52 times as long
as maximum width (fig. 65) . . . . . . . . .
. . . . . . . . . . . . . . . . . . Electrapis Cockerell
Metabasitarsus quadrangular, as long as maximal width (figs. 70, 78, 81) . . . . . . . . . 2
2. Lateral margins of clypeus strongly concave,
epistomal sulcus forming an obtuse angle
opening toward compound eye (fig. 69) . .
. . . . . . . . . . . . . . . Protobombus Cockerell
Lateral margins of clypeus linear, epistomal
sulcus straight (figs. 79, 80) . . . . . . . . .
. . . . . . . . . . . . Thaumastobombus, n. gen.
Genus Electrapis Cockerell, revised status

Electrapis Cockerell, 1908b: 326. Type species:
Apis meliponoides Buttel-Reepen, 1906, by designation of Cockerell, 1909b. Cockerell, 1909b:
7. Zeuner and Manning, 1976: 227.
Elektrapis Bischoff, 1927: 454. Lapsus calami.
Eckfeldapis Lutz, 1993: 180. Type species: Eckfeldapis electrapoides Lutz, 1993, monobasic
and original designation.
DIAGNOSIS: This genus can be separated

from other genera in the tribe by the angled
epistomal sulcus (fig. 64) and elongate metabasitarsus (fig. 65).
DESCRIPTION: Mandible with a few, weak
apical teeth. Malar space short, shorter than
basal mandibular width (fig. 64). Epistomal
sulcus forming orthogonal or obtuse angle

(fig. 64). Inner margins of compound eyes
slightly converging below, weakly emarginate. F1 longer than F2, not longer than combined lengths of F2 and F3; F3 longer than
F2. Preoccipital ridge rounded. Prosternum
without constriction between basisternum
and furcasternum; similar in construction to
Bombus; apophyseal pit present. Mesoscutal
anterior border broadly rounded; scutellum
projecting over metanotum and basal area of
propodeum. Basal area of propodeum strongly declivitous. Keirotrichiate field not elevated; metatibial spur serrate; auricular basket
sparse, setae short; rastellum strong and
formed of stiff setae; metabasitarsus 1.52
times as long as wide (fig. 65). Margin of
pterostigma within marginal cell concave to
convex; marginal cell apex offset from wing
margin by pterostigma width or more, apex
either rounded and appendiculate or truncate.
Sting usually not barbed (except in some
specimens of Protobombus and one of Thaumastobombus microscopic barbs are apparently present).
COMMENTS: One species of Electrapis has
been discovered as a compression fossil in
the middle Eocene oil-shale sediments of
Eckfelder Maar Formation [E. electrapoides
2001
(Lutz) (Lutz, 1993; Engel, 1998a: 101)].

With the splitting of Electrapis into multiple
genera, Eckfeldapis remains a junior synonym of Electrapis s.s. (e.g., elongate metabasitarsus, identical wing venation). Electrapis electrapoides has not been included in
the following key to species.
An individual of a species of Electrapis
(GPUH Nr. 3001; see below) is preserved
with its dorsum open at the amber surface
thereby allowing examination of the prosternal structure. It is from this individual that
the above definition of the Electrapis prosternal construction is based.
Key to Species of ELECTRAPIS in
Baltic Amber
1. Pterostigma size normal, longer than wide
(fig. 66) . . . . . . . . . . . . . . . . . . . . . . . . . 2
Pterostigma exceptionally short, length equal
to width (fig. 68) . . . . . . . . . . . . . . . . . . .
. . . . . . . . E. meliponoides (Buttel-Reepen)
2. Labral apex without distinctive patches of
elongate setae . . . . . . . . . . . . . . . . . . . . 3
Labral apex with patches of elongate setae
bordering midpoint . . . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . E. tornquisti Cockerell
3. Apical margins of T25 with bands of dense,
short, plumose setae; hind wing with first abscissa M 3.54.5 times length of rs-m; forewing 2rs-m distad 2m-cu by at least twice
vein width . . . . . . . E. martialis (Cockerell)
Apical margins of T25 without bands of
dense, short, plumose setae; hind wing with
first abscissa M 1.52 times length of rs-m
(fig. 67); forewing 2rs-m confluent with
2m-cu . . . . . . . . . . . E. krishnorum, n. sp.
Electrapis martialis (Cockerell),

new combination
Figure 64
Chalcobombus martialis Cockerell, 1908b: 326.

Cockerell, 1909c: 22. Zeuner and Manning,
1976: 208.
DIAGNOSIS: Electrapis martialis is most

similar to E. krishnorum but differs by the
presence of dense, plumose bands on the apical margins of the metasomal terga, the more
distad position of 2rs-m in the forewing, and
the more elongate first abscissa of M in the
hind wing.
wider than long (length 2.55, width 3.10).
93
Upper interorbital distance 1.65 mm; lower

interorbital distance 1.30 mm. Interocellar
distance 0.50 mm; ocellocular distance 0.45
mm; median to lateral ocellus 0.20 mm. Intertegular distance 2.55 mm. Basal vein basad cu-a by two times vein width, straight;
second abscissa Rs relatively straight; 1rs-m
distad 1m-cu by six times vein width; 2rs-m
confluent with 2m-cu; first submarginal cell
shorter than combined lengths of second and
third submarginal cells; posterior border of
second submarginal cell 1.5 times length of
anterior border; posterior border of third submarginal cell twice as long as anterior border,
anterior border approximately equal to anterior border of second submarginal cell; pterostigma longer than wide, margin inside marginal cell straight; marginal cell bending
away from wing apex by pterostigmal width,
feebly appendiculate; hind wing with 13 distal hamuli arranged in a single, evenly
spaced series.
Labrum smooth. Integument of face obscured by fine layer of Schimmel (setae visible extending out of this Schimmel and described below). Mesosoma finely imbricate
and impunctate except ventrally on mesepisternum with small punctures separated by a
puncture width and mesoscutum and scutellum with sparse, faint punctures separated by
a puncture width or less. Metasomal terga
and sterna finely imbricate and impunctate.
Color apparently dark brown without maculations. Wing membrane hyaline; veins
brown.
Pubescence white. Labrum with scattered,
erect, short, simple setae. Clypeus and supraclypeal area with scattered, moderate-length,
simple setae, such setae more numerous on
face and plumose, those setae of frons appearing to be dense enough to have obscured
integument if it were not already obscured
by fine layer of Schimmel. Setae on vertex
quite long, fuscous, and with minute branches by comparison to those setae of frons. Mesoscutum with dense, long, erect, plumose
setae partially obscuring integument. Tegula
with short, appressed, simple setae, setae
with a few minute branches. Scutellar setae
very long (1.52 times length of those on
mesoscutum, most setae as long as or longer
than scutellum) and plumose; setae entirely
obscuring view of metanotum and propo-
94
Fig. 65.
1 mm.
NO. 259
Left lateral view of holotype female of Electrapis krishnorum, new species. Scale bar 5
deum. Pleural pubescence long, dense, and

plumose. Setae of legs long, numerous, and
stiff on tibiae and tarsi, and mostly plumose;
dense tuft of very long, sinuous, simple setae
on ventral surface of mesotrochanter, setae of
tuft directed posteriorly; corbicula with long,
simple setae along lateral margins; sparse setae on outer metabasitarsus. Metasomal terga
with sparse, minute, appressed, simple setae
except on apical margins of T25 with bands
of dense, short, plumose, suberect setae; sterna with scattered, short, erect or suberect,
simple setae.
MATERIAL: Two total specimens. Neotype
(here designated). Female, worker caste, B-
JH 91 (AMNH) labeled Neotype, Chalcobombus martialis Cockerell, desig. M. S. Engel // Electrapis martialis (Cockerell), det.
M. S. Engel.
Non-type. Female, worker caste, Nr. 502
(CJDL) labeled: Nr. 502 // Electrapis
martialis (Cockerell), det. M. S. Engel.
Electrapis krishnorum, new species
Figures 6567
Apis sp. Gerlach, 1989: 257. [misidentification]

E. martialis in that both are relatively small
by comparison to E. meliponoides and E.
2001
tornquisti and both have pterostigmas that

are longer than wide while lacking distinctive setal patches on their labra. From E.
martialis, however, the new species differs
by the absence of dense bands of plumose
setae on the apical margins of T25, by the
shortened first abscissa of M in the hind
wing, and by the confluence of 2rs-m and
2m-cu in the forewing.
apparently wider than long (direct, frontal
view of head not possible). Mandibular
grooves weakened, those present appear to
be (from upper to lower margin) acetabular
groove, outer upper groove, and outer lower
groove. Intertegular distance 2.15 mm. Basal
vein basad cu-a by vein width, straight; second abscissa Rs relatively straight; 1rs-m distad 1m-cu by six times vein width; 2rs-m distad 2m-cu by vein width; first submarginal
cell shorter than combined lengths of second
and third submarginal cells; posterior border
of second submarginal cell 1.25 times length
of anterior border; posterior border of third
submarginal cell nearly twice as long as anterior border, anterior border approximately
equal to anterior border of second submarginal cell; pterostigma longer than wide,
margin inside marginal cell slightly convex;
marginal cell bending away from wing apex
by slightly less than pterostigma width, feebly appendiculate; hind wing with 13 distal
series; first abscissa M slightly less than
twice as long as rs-m.
Mandibular integument smooth except appearing minutely granular within weakened
mandibular grooves. Labrum minutely granular. Clypeus and supraclypeal area with
coarse, faint punctures separated by two
times a puncture width or more, integument
between smooth. Face, vertex, gena, and
postgena smooth and impunctate. Mesosoma
smooth and impunctate except ventrally on
mesepisternum; posteriorly and laterally on
mesoscutum and scutellum faint punctures
separated by a puncture width or less. Metasomal terga and sterna finely imbricate and
impunctate.
Color apparently dark brown, without
maculations. Wing membrane hyaline; veins
dark brown.
95
Pubescence silvery except those setae of

vertex, tarsi, and apical terga slightly fuscous. Mandible with a few, very short, widely scattered, simple setae except lower margin with long, simple, scattered setae. Labrum with scattered, erect, short, simple setae. Clypeus and supraclypeal area with
scattered, moderate-length, simple setae,
such setae more numerous on face and some
with a few, minute branches, those setae of
face partially obscuring integument. Setae on
vertex quite long, slightly fuscous, and with
more minute branches than those setae of
face. Scape with minute, simple, scattered,
appressed setae. Postgena with scattered,
long, erect, simple setae. Mesoscutum with
dense, long, erect, plumose setae partially
obscuring integument. Tegula with short, appressed, simple setae scattered on surface except anteriorly setae longer and with a few
minute branches. Scutellar setae very long
(at least 1.5 times length of those on mesoscutum, most setae as long as or longer than
scutellum itself) and plumose; setae entirely
obscure view of metanotum and propodeum.
Pleural pubescence long, dense, and either
simple or minutely plumose. Setae of legs
long, numerous and stiff on tibiae and tarsi;
dense tuft of very long, sinuous, simple setae
on ventral surface of mesotrochanter, setae of
tuft directed posteriorly; corbicula with long
setae along lateral margins; stiff scattered setae on outer metabasitarsus and stiff comb
rows on inner surface. Metasomal terga with
sparse, minute, simple, appressed setae, terga
without apical setal bands; sterna with scattered, short, erect, simple setae.
MATERIAL: Four specimens. Holotype. Female, worker caste, II1926 (BMNH) labeled:
II1926(1) // Bombus?, Baltic amber//
Electrapis // Holotype, Electrapis krishnorum Engel.
Paratype. Female, worker caste, 5589
(ZMPA) labeled: Muzeum Ziemi w Warszawie, 5589 (in two pieces), Apidae in Baltic Amber, Gerlach 1989: Apis L., Electrapis:
det. Engel, Golarisk-Stogi, coll. T. Gierewicz, 1972 // Baltic Amber, Eocene, 5589,
Muzeum Ziemi Pan, M. S. Engel study //
Paratype, Electrapis krishnorum Engel.
Paratype. Female, worker caste, Nr. 25
(CSUL) labeled: Paratype, Electrapis krishnorum Engel.
96
NO. 259
Figs. 6667. Wings of holotype female of Electrapis krishnorum, new species. 66. Forewing. 67.
Hind wing. Scale bar 5 1 mm.

(CJDL) labeled: Nr. 71 // Paratype, Electrapis krishnorum Engel.
ETYMOLOGY: The specific epithet is a patronymic honoring my dear friends and colleagues Drs. Kumar and Valerie Krishna,
worlds authorities on termites and medieval
literature, respectively.
Electrapis tornquisti Cockerell
Electrapis tornquisti Cockerell, 1908b: 326.
1976: 231.
Electrapis hermenaui Bischoff In Kelner-Pillault,
1974: 624. Nomen nudum [5 Electrapis bombusoides Kelner-Pillault, 1974].
Electrapis cockerelli Bischoff In Kelner-Pillault,

1974: 624. Nomen nudum [5 Electrapis bombusoides Kelner-Pillault, 1974].
Electrapis cokerelli Kelner-Pillault, 1974: 624.
Lapsus calami.
Electrapis bombusoides Kelner-Pillault, 1974:
625. NEW SYNONYMY.
DIAGNOSIS: Electrapis tornquisti, like E.

meliponoides, is a larger species than E. martialis and E. krishnorum. The species can be
distinguished from E. meliponoides by the
longer pterostigma and the presence of medioapical patches of elongate setae on the labrum.
2001
longer than wide (length 4.41 mm, width

mm; lower interorbital distance 2.09 mm.
Basal vein basad cu-a by three times vein
width, straight; second abscissa Rs relatively
straight; 1rs-m distad 1m-cu by seven times
vein width; 2rs-m distad 2m-cu by two times
vein width; first submarginal cell shorter than
combined lengths of second and third submarginal cells; posterior border of second
submarginal cell 1.5 times length of anterior
border; posterior border of third submarginal
cell twice as long as anterior border, anterior
border slightly less than length of anterior
border of second submarginal cell; pterostigma longer than wide, margin inside marginal
cell slightly concave; marginal cell bending
away from wing apex by pterostigmal width,
feebly appendiculate; hind wing with 17 distal hamuli arranged in a single, evenly
spaced series; first abscissa M four times longer than rs-m.
Mandibular integument smooth except appearing minutely granular within weakened
mandibular grooves. Labrum minutely granular. Clypeus and supraclypeal area with
coarse, faint punctures separated by two
times a puncture width or more, integument
between smooth. Face, vertex, gena, and
postgena finely imbricate and impunctate.
Mesosoma finely imbricate and impunctate
except some sparse, faint punctures on mesoscutum and scutellum. Metasomal terga
and sterna finely imbricate; terga impunctate;
S12 apparently impunctate, S35 minutely
punctured on apical thirds, S6 minutely
punctured on apical half, punctures on all
sterna separated by 13 times a puncture
width.
Color, where evident, dark brown without
black.
Pubescence white except those setae of
protibia and tarsi black. Mandible with a few,
very short, widely scattered, simple setae except lower margin with long, simple, scattered setae. Labrum with scattered, erect,
short, simple setae and apical margin with
mediolateral patches of elongate, simple setae. Clypeus and supraclypeal area with scattered, moderate-length, simple setae, setae
more numerous on face and minutely plumose. Setae on vertex quite long, white (not
97
fuscous as in E. krishnorum), and minutely

plumose. Mesoscutum with dense, long,
erect, plumose setae partially obscuring integument. Scutellar setae very long, dense,
and plumose. Pleural pubescence long,
dense, and minutely plumose. Setae of legs
long, numerous, and stiff on tibiae and tarsi;
inner surface of metafemur covered by keirotrichiae except medially this field narrows
slightly; corbicula with long setae along lateral margins, inner surface of metatibia entirely covered by keirotrichiae; stiff scattered
setae laterally on metabasitarsus, outer surface with sparse, simple setae, inner surface
with comb rows. Metasomal terga with scattered, simple or minutely plumose setae, apical margins of terga with dense bands of
short, plumose setae; sterna with scattered,
erect, simple setae.
M ATERIAL : Three specimens. Neotype
(tornquisti; here designated). Female,
worker caste, MB.I.1947 (ZMHB) labeled:
Neotype, Electrapis tornquisti Cockerell,
desig. M. S. Engel [red label]. This is also
the holotype specimen of E. bombusoides
and bears additional labels reading: Electrapis bombusoides n. sp., Holotype, S. Kelner-Pillault // Palaont. Mus. Berlin, coll.
Kuhl [actually from the Berendt collection]
// Electrapis hermanaui, Bisch., Paratypus,
1930 // Elektrapis [sic] hermanaui, 20 /
/ Holotyp [sic], Palaontologisches Museum
Berlin, Electrapis bombusoides Kelner-Pillault, 1974, slg. Berendt.
Holotype (bombusoides). Same specimen
as neotype of E. tornquisti (above).
Paratype (bombusoides). Female, worker
caste, MB.I.1948 (ZMHB) labeled: 27 //
Paratype, Electrapis bombusoides //
MB.I.1948 // Paratyp [sic], Palaontologisches Museum Berlin, Electrapis bombusoides Kelner-Pillault, 1974.
Non-type. Female, worker caste (IMGP)
labeled: Electrapis bombusoides n. sp., det.
S. Kelner-Pillault [in Kelner-Pillaults handwriting] // Electrapis cockerelli Bisch.,
Paratype, 1930 // Electrapis cockerelli
Bisch., Paratypus, 1930 // 27 Electrapis
Cockerelli [sic] Bisch., Chalcobombus? humilis Cockerell, Electrapis bombusoides S.
Kelner P. // 6021, Electrapis bombuscoides [sic] n. sp., Kelner-Pillault, GeologischPalaontologisches Institut Gottingen //
98
Chalcobombus humilis Cock. [stricken

through], Electrapis Cockerelli [sic] Bisch..
This specimen is very poorly preserved and
is only tentatively placed in E. tornquisti.
COMMENTS: At first notice the specimen
from Gottingen labeled as C. humilis and
then as E. cockerelli might be considered as
the missing type of C. humilis and simply
misinterpreted by Bischoff and Kelner-Pillault as Electrapis. However, Chalcobombus
was defined as having a quadrangular metabasitarsus (among other features) and this is
not the case with the fossil (the metabasitarsus being twice as long as broad; typical of
Electrapis). I therefore concur with Bischoff
and Kelner-Pillault that this cannot be Cockerells missing type (a neotype for C. humilis
is designated below). Moreover, the labels
reading Chalcobombus are all in Bischoffs
handwriting; none are in Cockerells distinctive script.
Bischoff had originally intended to produce a paper on this material and to summarize the then known bees in Baltic amber
under the title Apiden des Bernsteins (to
have appeared in the 1939 volume of Bernstein Forschungen); however, this paper was
never published. Kelner-Pillault subsequently
borrowed this material from Berlin and prepared a description of Bischoffs specimens.
At that time she mentioned the unpublished
species names with which Bischoff had labeled the specimens (i.e., E. hermenaui, E.
cockerelli). Under Recommendation 50C
(ICZN, 1999b) authorship of these unavailable names should be attributed to Bischoff
in Kelner-Pillault.
In Kelner-Pillaults (1974) illustration of
the hind wing of the holotype female of E.
bombusoides she omitted the jugal lobe and
the wing drawing therefore strongly resembles that of a true bombine. The holotype
specimen clearly has a jugal lobe present at
the base of the hind wing; this structure is
even more apparent in the specimen she
chose as a paratype.
Electrapis meliponoides (Buttel-Reepen)
Figure 68
Plate 6c
Apis meliponoides Buttel-Reepen, 1906: 158.

Electrapis meliponoides (Buttel-Reepen); Cock-
NO. 259
Fig. 68. Forewing of neotype female of Electrapis meliponoides (Buttel-Reepen). Scale bar 5
1 mm.
erell, 1908b: 326. Cockerell, 1909b: 9. Zeuner

and Manning, 1976: 229.
DIAGNOSIS: Electrapis meliponoides is distinctive among all known Electrapis species

for the shortened pterostigma. The species is
large like E. tornquisti but lacks the medioapical patches of setae on the labrum.
longer than wide (length 2.85 mm, upper
width 2.40 mm: lower width not preserved
owing to damage to lower right of head). Intertegular distance 3.40 mm. Basal vein basad cu-a by vein width, straight; second abscissa Rs arched; 1rs-m distad 1m-cu by five
times vein width; 2rs-m distad 2m-cu by four
times vein width; first submarginal cell shorter than combined lengths of second and third
submarginal cells; posterior border of second
submarginal cell 1.25 times length of anterior
cell twice as long as anterior border, length
of anterior border slightly less than anterior
border of second submarginal cell; pterostigma short, as long as wide, margin inside marginal cell slightly concave; marginal cell
bending away from wing apex by nearly
twice pterostigma width, feebly appendiculate; hind wing with 13 distal hamuli arranged in a single, evenly spaced series; first
abscissa M 4.5 times as long as rs-m.
Preserved portion of facial integument obscured by fine layer of Schimmel; where evident, appearing imbricate. Mesosoma and
metasoma, where evident, finely imbricate.
Color dark brown without maculations.
Wing membrane hyaline; veins dark brown.
Pubescence white. Mandible with a few,
very short, widely scattered, simple setae except lower margin with long, simple, scat-
2001
tered setae. Labrum with scattered, erect,

long, simple setae. Clypeus and supraclypeal
area with scattered, moderate-length, simple
setae, such setae more numerous on face and
some with a few minute branches; those setae of face fairly dense and plumose. Setae
on vertex quite long and simple. Genal setae
as described for vertex except much shorter.
Mesoscutum with dense, long, erect, plumose setae partially obscuring integument.
Tegula with dense, long, appressed, plumose
setae directed laterally. Scutellar setae as described for mesoscutum except along posterior third exceedingly long (longer than scutellum) and plumose. Pleural pubescence
long, dense, and plumose. Setae of legs long,
numerous and stiff on tibiae and tarsi; corbicula with long setae along lateral margins,
setae of anterior margin simple, posterior
margin minutely plumose; inner surface of
metatibia entirely covered by keirotrichiae;
stiff, elongate, scattered setae on lateral margins of metabasitarsus, outer surface with
sparse, shorter, simple setae, inner surface
with dense field of setae not arranged into
rows. Metasomal terga with scattered, short,
plumose setae, apparently with dense, apical
bands on T25; sterna with scattered, long,
erect or suberect, minutely plumose setae.
MATERIAL: One specimen. Neotype (here
designated). Female, worker caste, B-JH 97
Kaliningrad, Yantarny // Neotype, Apis
meliponoides Buttel-Reepen, desig. M. S.
Engel // Electrapis meliponoides (ButtelReepen), det. M. S. Engel.
Electrapis cf. krishnorum
MATERIAL: One specimen. Non-type. Female, worker caste, Nr. 501 (CJDL) labeled:
Nr. 501 // Electrapis sp., det. M. S. Engel.
COMMENTS: This specimen represents a
species near to E. krishnorum but has distinctive differences in both the wing venation
as well as the mouthparts.
Electrapis sp. indet.
MATERIAL: One specimen. Non-type. Female, worker caste, Nr. 577/2 (CHFG) labeled: Nr. 577/2, Bittf. [Bitterfeld] Bern-
99
stein // Electrapis sp. indet., det. M. S.

Engel.
MATERIAL: One specimen. Non-type. Female?, Nr. 461 (GPUH) labeled: Nr. 461
(Scheele), Apis meliponoides Buttel-Reepen
5 Electrapis meliponoides (Buttel-Reepen)
// Electrapis sp. indet., det. M. S. Engel.
COMMENTS: This specimen was identified
by Zeuner and Manning (1976) as belonging
to E. meliponoides but the individual is entirely covered by fractures, Schimmel, and
pyrite and such an indentification cannot actually be made. The size of the specimen is
about that of E. meliponoides.
MATERIAL: One specimen. Non-type. Female, worker caste, Nr. 1900937 (CGHG)
labeled: Nr. 1900937 // Electrapis sp.
indet., det. M. S. Engel.
MATERIAL: Three specimens. Non-type.
Females worker caste? (CMGG) labeled:
Electrapis sp. indet., det. M. S. Engel.
MATERIAL: One specimen. Non-type. Female, worker caste, 3001 (GPUH) labeled:
Electrapis sp. indet., det. M. S. Engel,
1999.
COMMENTS: This individual is preserved
with its dorsum at the amber surface and its
mesosoma and metasoma opened. The internal sclerites are preserved and can be examined with some ease. The generic identification of the specimen is still possible owing to the relatively good preservation of the
ventral half of the bee and it can be assigned
to Electrapis with confidence. Specific identification, however, is not possible. It is from
this individual that the above definition of the
prosternal shape in Electrapis was made.
Genus Protobombus Cockerell
Protobombus Cockerell, 1908b: 326. Type species: Protobombus indecisus Cockerell, 1908b,
monobasic [also designated by Cockerell,
100
NO. 259
1909b]. Cockerell, 1909b: 9. Manning, 1960:

306. Zeuner and Manning, 1976: 231.
Chalcobombus Cockerell, 1908b: 326. Type species: Chalcobombus humilis Cockerell, 1908b
[5 Protobombus indecisus Cockerell, 1908b],
by designation of Cockerell, 1909b; isotypic
with Protobombus Cockerell, 1908b by synonymy. Cockerell, 1909b: 11. Zeuner and Manning, 1976: 206. NEW SYNONYMY.
Sophrobombus Cockerell, 1908b: 326. Type species: Sophrobombus fatalis Cockerell, 1908b,
monobasic [also designated by Cockerell,
1909c]. Cockerell, 1909c: 21. Zeuner and Manning, 1976: 213. NEW SYNONYMY.
DIAGNOSIS: Electrapine bees with a short,

quadrangular metabasitarsus; similar in this
respect to Thaumastobombus (described below). Protobombus differs from Thaumastobombus most notably, however, in the obtuse
angle formed by the epistomal sulcus. Some
Protobombus, like Thaumastobombus, have
a superficial Apis-like habitus.
apical teeth. Malar space shorter than basal
mandibular width. Epistomal sulcus forming
obtuse angle (fig. 69). Inner margins of compound eyes parallel, not emarginate (fig. 69).
F1 longer than F2, not longer than combined
lengths of F2 and F3; F3 longer than F2. Preoccipital ridge rounded. Mesoscutal anterior
border broadly rounded; scutellum projecting
over metanotum and basal area of propodeum. Basal area of propodeum strongly declivitous. Keirotrichiate field not elevated;
metatibial spur serrate or minutely ciliate; auricular basket sparse, setae short; rastellum
strong and formed of stiff setae; metabasitarsus as long as wide. Margin of pterostigma
within marginal cell convex; marginal cell
apex offset from wing margin by pterostigma
width or slightly less, apex rounded and feebly appendiculate. Sting variable, usually not
barbed (some specimens appear to have microscopic barbs).
COMMENTS: The genus Sophrobombus was
regarded as a genus of Meliponini by Zeuner
and Manning (1976) because the forewing
venation was apparently reduced. The
type species, Sophrobombus fatalis Cockerell, is notable for the presence of only two
submarginal cells owing to the apparent absence of 1m-cu (i.e., the second transversocubital vein in Cockerells system) (Cocker-
Fig. 69. Frontal view of head of neotype female of Protobombus indecisus Cockerell. Scale
bar 5 1 mm.
ell, 1908b, 1909c). The type specimen of S.

fatalis has been lost but Zeuner and Manning
(op. cit.) located a specimen in the Scheele
Collection (presently in the Geologisch-Palaontologisches Institut und Museum, Universitat Hamburg) that matches Cockerells
description in all respects. This specimen is
here designated as the neotype for S. fatalis.
The presence of only two submarginal cells
is not the same forewing venation reduction as is diagnostic for Meliponini. In meliponines the distal wing veins (i.e., 2rs-m
and 2m-cu) are lost, the marginal cell is typically open, and frequently other veins are
also missing or greatly reduced. None of
these are true for S. fatalis. The presence of
a single inner metatibial spur, of an auricle,
a supra-alar carina, an inner tooth on the
claw, as well as numerous other characters
separate this species from the Meliponini.
Sophrobombus is actually an electrapine bee
and similar in all respects to species of Protobombus, differing only in the number of
2001
submarginal cells. Cockerell (1908b), who

did not ally this species with the stingless
bees (see fig. 120), separated Sophrobombus
from Protobombus by the number of submarginal cells and by the basal vein being
basad cu-a in the former. Below I describe a
new species (P. basilaris) that blends the defining characters of the genera (i.e., basal
vein basad cu-a but with three submarginal
cells). I have therefore placed Sophrobombus
as a junior synonym of Protobombus.
Chalcobombus was differentiated from
Protobombus by the shape the pterostigma as
well as the third submarginal cell. The shape
of the third submarginal cell varies among
the species of Electrapini and, although species-specific characters can be identified,
these differences in shape are not correlated
with other structural characters. Similarly,
the shape of the pterostigma is quite variable
and was incongruent with all other characters
(including the shape of the third submarginal
cell!). Chalcobombus appears to be synonymous with Protobombus.
Key to Species of PROTOBOMBUS
1. Forewing with three submarginal cells (i.e.,
1rs-m present) (figs. 75, 77) . . . . . . . . . 2
Forewing with two submarginal cells (i.e.,
1rs-m absent) (fig. 74) . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . P. fatalis (Cockerell)
2. Basal vein confluent with cu-a (fig. 77); hind
wing with first abcissa of M approximately
equal in length to second abcissa of M1Cu
(fig. 73) . . . . . . . . . . . . . . . . . . . . . . . . . 3
Basal vein basad cu-a by twice vein width
(fig. 75); hind wing with first abcissa of M
distinctly shorter than second abcissa of
M1Cu (fig. 76) . . . . . P. basilaris, n. sp.
3. Apical third of clypeus extending below lower tangent of compound eyes; keirotrichiate
field of metatibial inner surface separated
from metatibial apex by glabrous zone of
variable length . . . . . . . . . . . . . . . . . . . . 4
Clypeus not extending below lower tangent
of compound eyes; keirotrichiate field of
metatibial inner surface separated from metatibial apex by glabrous zone that is half
width of keirotrichiate field in length . . . .
. . . . . . . . . . . . . . . . P. tristellus Cockerell
4. Keirotrichiate field of metatibial inner surface
separated from metatibial apex by glabrous
zone that is half width of keirotrichiate field
in length; posterior margin of third sub-
101
maringal cell twice length of anterior border (fig. 77) . . . . . P. hirsutus (Cockerell)
Keirotrichiate field of metatibial inner surface
separated from metatibial apex by glabrous
zone that is equal to width of keirotrichiate
field in length (fig. 71); posterior margin of
third submarginal cell thrice length of anterior border . . . . . P. indecisus Cockerell
Protobombus indecisus Cockerell

Figures 6973
Plate 6a,b
Protobombus indecisus Cockerell, 1908b: 326.

Cockerell, 1909b: 10.
Chalcobombus humilis Cockerell, 1908b: 326.
NEW SYNONYMY.
Electrapis (Electrapis) apoides Manning, 1960:
307. Zeuner and Manning, 1976: 227. NEW SYNONYMY.
Electrapis (Protobombus) indecisus (Cockerell);
Zeuner and Manning, 1976: 232.
DIAGNOSIS: This species can be recognized

by the combination of three submarginal
cells, apical third of clypeus below lower
tangent of compund eyes, and the keirotrichiate field on the inner surface of the metatibia separated from metatibial apex by distance equal to its own apical width.
0.38 mm; median to lateral ocellus 0.09 mm.
Lower half of clypeus below lower tangent
of compound eyes. Intertegular distance 2.15
mm (measured from second specimen in amber piece). Basal vein confluent with cu-a,
straight; second abscissa Rs relatively
straight; 1m-cu basad 1rs-m by seven times
vein width; 2rs-m distad 2m-cu by three
times vein width, strongly arched; anterior
border of second submarginal cell approximately one-third length of posterior border;
posterior border of third submarginal cell
slightly more than twice length of anterior
border, anterior border twice as long as anterior border of second submarginal cell;
hind wing with 10 distal hamuli arranged in
a single, evenly spaced series; hind wing
with first abscissa M approximately equal in
length to second abscissa M1Cu.
102
NO. 259
Figs. 7072. Leg structures of neotype female of Protobombus indecisus Cockerell. 70. Outer surface of metatibia and metabasitarsus. 71. Inner surface of metatibia. 72. Claw and arolium. Scale bars
5 0.25 mm (a), 1 mm (b); (a 5 fig. 72; b 5 figs. 70, 71).
Labrum imbricate and impunctate. Clypeus with sparse, small, faint punctures, integument between faintly imbricate. Supraclypeal area, face, vertex, gena, and postgena
with sparse, small, faint punctures, integument between smooth. Pronotum finely imbricate. Mesoscutum and scutellum with
sparse, faint punctures, integument between
faintly imbricate. Metanotum imbricate and
impunctate. Pleura with sparse, small punc-
Fig. 73.
tures, integument between finely imbricate.

Propodeum faintly imbricate and impunctate.
Corbicula smooth and impunctate. Metasomal terga and sterna finely imbricate.
Where evident integument dark brown and
shining. Wing membrane very lightly fuscous; veins dark brown.
Pubescence golden. Labrum with short,
scattered, erect, simple setae. Clypeus with
scattered, short, simple, suberect setae inter-
Hind wing of neotype female of Protobombus indecisus Cockerell. Scale bar 5 1 mm.
2001
mixed with scattered, minute, appressed,

simple setae. Face below level of antennal
sockets with appressed, short, minutely plumose setae, setae numerous but not obscuring integument; supraclypeal area, vertex,
gena, and remainder of face with scattered,
simple, suberect or erect setae a few on vertex and gena with minute branches. Postgena
with scattered, short, erect, simple setae.
Pronotum with sparse, short, suberect simple
setae except along posterior border and covering pronotal lobe with moderately long,
erect, dense, plumose setae. Mesoscutum
with scattered, short to moderate length,
erect, minutely plumose setae. Tegula with
short, scattered, minutely plumose setae anteriorly, posteriorly with minute, simple setae. Scutellar setae as described for mesoscutum except setae dense on posterior border
and on axilla. Metanotum with sparse, minute, appressed, simple setae. Pleura with
short, scattered, erect, simple setae except a
few with minute branches. Lateral and posterior surfaces of propodeum as described for
pleura except setae minute; basal area of propodeum without pubescence. Corbicular setae mostly simple but a few minutely plumose also with a few minute setae on corbicular surface itself (fig. 70), inner surface
of metatibia mostly covered by keirotrichiate
field, field separated from metatibial apex by
length approximately equal to its width; metabasitarsus with scattered, short, simple setae, posterior edge with longer setae, inner
surface with 56 comb rows. Metasomal terga with sparse, minute, appressed, simple setae; sterna with suberect to erect, moderately
long, scattered setae, each with a few minute
branches, setae entirely postgradular.
MATERIAL: Four specimens. Neotype (indecisus; here designated). Female, worker
caste, B-JH 98 (AMNH) labeled: Baltic
Neotype, Protobombus indecisus Cockerell,
desig. M. S. Engel. There are two individuals in the same block of amber; the neotype
is the one figured in plate 6b and the righthand bee in plate 6a.
Neotype (humilis; here designated).
Same specimen as the neotype of P. indecisus (above); bears one additional label reading: Neotype, Chalcobombus humilis Cockerell, desig. M. S. Engel.
103
Holotype (apoides). Female, worker caste

(MNHN). Not seen. I have based my identification on photographs of the holotype
specimen as well as illustrations of the wing
venation and descriptive notes given by both
Manning (1960) and Kelner-Pillault (1970a).
From this information it is quite clear that E.
apoides does not belong in Electrapis (see
comments below) and is, in fact, conspecific
with the specimen designated above as the
neotype for P. indecisus.
Non-type. Female, worker caste, B-JH 98
(AMNH) preserved in same piece with neotype (see above).
(CCGG) labeled: Nr. 1521 // Protobombus indecisus Cockerell, det. M. S. Engel.
COMMENTS: Although I was not able to examine the holotype of E. apoides, the placement of this species in Protobombus is simple owing to the quadrate metabasitarsus not
only described in the original paper by Manning (1960) and by Zeuner and Manning
(1976), but also nicely depicted in the published photograph of the holotype by KelnerPillault (1970a). Additionally, the confluence
of the basal vein among other characters
clearly shown in the photograph all suggest
that this species is conspecific with Cockerells P. indecisus. The line illustrations of the
wing venation presented by Kelner-Pillault
(1970b) are quite poor and I have not trusted
them, instead referring to the descriptions
and other illustrations. In fact, the holotype
of E. apoides agrees quite well with not only
Cockerells descriptive notes about P. indecisus but also with the specimens designated
here as neotypes. I therefore feel confident in
the above synonymy.
Protobombus fatalis (Cockerell),
new combination
Figure 74
Sophrobombus fatalis Cockerell, 1908b: 326.

1976: 214.
DIAGNOSIS: This species is immediately

recognizable for the complete absence of 1rsm in the forewing thus forming only two
submarginal cells (fig. 74). The basal vein is
basad cu-a and therefore resembles to some
104
Fig. 74. Forewing of neotype female of Protobombus fatalis (Cockerell). Scale bar 5 1 mm.
degree P. basilaris. The mandible of P. fatalis bears a strong, broad subapical tooth
(see Material and Comments, below).
width 1.94 mm). Mandible with strong,
broad subapical tooth; tooth separated from
remainder of the apical mandibular margin
by deep incision. Interocellar distance 0.38
mm; ocellocular distance 0.34 mm; median
to lateral ocellus 0.16 mm. Intertegular distance 1.59 mm. Basal vein basad cu-a by
three times vein width, straight; second abscissa Rs arched; 1m-cu distad second abscissa Rs by seven times vein width; 2rs-m
distad 2m-cu by three times vein width,
strongly arched; second submarginal cell extremely elongate owing to absence of 1rs-m.
Face above antennal sockets, vertex and
gena finely imbricate and impunctate. Pronotum finely imbricate and impunctate. Mesoscutum and scutellum faintly imbricate and
impunctate. Mesepisternum sculptured as on
mesoscutum. Corbicula smooth and impunctate. Metasomal terga and sterna finely imbricate.
Coloration not preserved. Wing membrane
hyaline; veins brown.
Pubescence whitish. Face above level of
antennal sockets and vertex with scattered,
simple or minutely branched, erect setae.
Gena with sparse, minute, appressed, simple
setae. Pronotum with sparse, short, suberect,
simple setae except along posterior border
and covering pronotal lobe with moderately
long, erect, dense, plumose setae. Mesoscutum with scattered, short to moderate length,
erect, minutely plumose setae. Tegula with
short, scattered, minutely plumose setae anteriorly, posteriorly with minute, simple se-
NO. 259
tae. Scutellar setae as described for mesoscutum. Mesepisternum with short, scattered,
erect, simple setae except a few with minute
branches. Corbicular setae mostly simple and
sinuous, also with three elongate, simple, sinuous setae on corbicular surface, corbicula
occupying glabrous region on lower fourfifths of metatibia, inner surface of metatibia
covered by keirotrichiate field, field separated from metatibial apex by length approximately equal to its width; metabasitarsus
with scattered, short, simple setae, posterior
edge with longer setae, inner surface with
seven comb rows. Metasomal terga with
sparse, minute, appressed, simple setae; sterna with suberect, moderately long, scattered,
simple setae, setae entirely postgradular.
MATERIAL: Two specimens. Neotype (here
designated). Female, Nr. 1225 (GPUH) labeled: Neotype, Sophrobombus fatalis
Cockerell, desig. M. S. Engel // Geol.-Palaont. Inst. Univ. Hamburg, Nr. 1225 (Scheele) Sophrobombus fatalis Cockerell //
Protobombus fatalis (Cockerell), det. M. S.
Engel, 1999. This specimen is poorly preserved with the ventral portion of the head
lost into a series of fractures that eventually
lead into a cavity on the lower surface of the
amber piece. This cavity appears to open into
what would have been the lowermost portion
of the head and the anteroventral area of the
mesosoma. In addition, numerous small fracture planes and fine layers of Schimmel obscure various structures of the bee. It can
still, however, be placed confidently into the
species as defined by Cockerell owing to the
preservation of the metatibia, metabasitarsus,
and forewing venation, all of which indicate
this specimen to be conspecific with Cockerells original material.
Non-type. Female, collection of Michael
Baatjer (Hamburg, Germany) labeled: Protobombus fatalis (Cockerell), det. M. S. Engel, 2000. Although this specimen is in better condition than the neotype, it is part of a
private collection and is presently available
for purchase; thus, the final depository of this
specimen remains unknown. In the interest
of stability I have therefore chosen the less
perfect individual to serve as the neotype
since the whereabouts of this specimen will
certainly change in the coming months and
years rendering it difficult, if not impossible,
2001
to locate again. This specimen was received

at the time when the remainder of the manuscript was going to press. It was therefore
not possible to prepare illustrations. In all
comparable features this specimen is conspecific with the neotype designated above. The
abdomen is unfortunately distended and portions of the legs and mesosoma cleared in
preservation but is otherwise a nice specimen. It is from this specimen that information on the mandibular structure of P. fatalis
was extracted. Similarly, information on the
structure of other characters (e.g., the epistomal sulcus) can all be seen in this individual and confirms the placement of P. fatalis
in the Electrapini and Protobombus. Furthermore, the information provided by this new
material helps to reinforce the identity of the
neotype as well as this specimen for being
conspecific with Cockerells (1908b, 1909c)
missing original material.
COMMENTS: The specimen I studied was
the same one examined by Zeuner and Manning (1976); see comments above for synonymy of the genus Sophrobombus with
Protobombus. Cockerell (1909c) noted that
his monotypic Sophrobombus was very
much like Protobombus and Chalcobombus
(above considered as synonyms) but differed
most notably by the presence of only two
submarginal cells. The specimen described
by Cockerell (1908b, 1909c) agrees in every
respect with the one selected here as the neotype, particularly in the unique trait of having
only two submarginal cells. Cockerells original specimen lacked mandibular dentition
(differing in this respect from the non-type
specimen examined here); it is possible that
the mandibles were simply worn or set close
the the face where the large subapical tooth
would be completely obscured (for instance,
in the available specimen the labrum and
clypeus totally cover the subapical tooth of
one mandible while the other, more fully
opened, displays this character nicely).
Protobombus basilaris, new species
Figures 75, 76
DIAGNOSIS: This species is notable for the

position of the basal vein that is basad cu-a.
Protobombus fatalis is similar in this respect
but has only two submarginal cells.
105

mm). Interocellar distance 0.34 mm; ocellocular distance 0.34 mm; median to lateral
ocellus 0.13 mm. Lower half of clypeus below lower tangent of compound eyes. Intertergular distance 1.72 mm. Basal vein basad
cu-a by twice vein width, straight; second abscissa Rs arched; 1m-cu basad 1rs-m by seven times vein width; 2rs-m distad 2m-cu by
vein width, strongly arched; posterior border
of second submarginal cell approximately
three times length of anterior border; posterior border of third submarginal cell 1.75
times length of anterior border, anterior border twice as long as anterior border of second
submarginal cell; hind wing with 10 distal
series.
Labrum finely imbricate and impunctate;
remainder of head and entirety of mesosoma
and metasoma smooth and impunctate.
Integument dark brown and shining. Wing
membrane hyaline; veins black.
Pubescence whitish. Labrum with short,
scattered, erect, simple setae. Clypeus with
scattered, short, simple, suberect setae. Face,
supraclypeal area, vertex, and gena with scattered, simple, suberect or erect setae, those
on vertex and gena with minute branches.
Pronotum with sparse, minute, suberect, simple setae except along posterior border and
covering pronotal lobe with short, erect, plumose setae and a few elongate, plumose setae
on dorsolateral angle that are slightly fuscous. Mesoscutum with scattered, short to
moderate length, erect, minutely plumose setae. Tegula with sparse, appressed, minute
setae. Scutellar setae as described for mesoscutum except setae dense and elongate on
posterior border. Pleura with short, scattered,
erect, simple setae. Lateral and posterior surfaces of propodeum as described for pleura
except setae minute; basal area of propodeum without pubescence. Corbicular setae
mostly simple but a few minutely plumose
also with a few minute setae on corbicular
surface itself, inner surface of metatibia
mostly covered by keirotrichiate field, field
separated from metatibial apex by length approximately equal to its width; metabasitarsus with scattered, short, simple setae, pos-
106
NO. 259
Figs. 7577. Wings of Protobombus species. 75. Forewing of Protobombus basilaris, new species.
76. Hind wing of P. basilaris, new species. 77. Forewing of P. hirsutus (Cockerell). Scale bars 5 1
mm (a 5 fig. 76; b 5 figs. 75, 77).
2001
terior edge with longer setae. Metasomal terga with sparse, minute, appressed, simple setae; sterna with suberect to erect, short,
scattered setae, each with a few minute
branches, setae entirely postgradular.
MATERIAL: One specimen. Holotype. Female, worker caste, MB.I.1939 (ZMHB) labeled: Museum fur Naturkunde Berlin, Palaontologisches Museum, Inv. Nr. MB.I.1939
(No. 36) // 36 [handwritten label] //
Holotype, Protobombus basilaris Engel.
ETYMOLOGY: The specific epithet is the
Greek word basilaris, meaning at the
base, and is a reference to the basal position
of the basal vein relative to cu-a.
Protobombus tristellus Cockerell
Protobombus tristellus Cockerell, 1909c: 24.
Electrapis (Protobombus) tristellus (Cockerell);
Zeuner and Manning, 1976: 233.
DIAGNOSIS: This species is similar to P. indecisus and P. hirsutus but can be distinguished from both by the apical margin of
the clypeus not extending below the lower
tangent of compound eyes.
Lower third of clypeus below lower tangent
of compound eyes. Intertegular distance 1.72
mm. Basal vein confluent with cu-a, straight;
second abscissa Rs arched; 1m-cu basad 1rsm by six times vein width; 2rs-m confluent
2m-cu, strongly arched; posterior border of
second submarginal cell 2.5 times length of
anterior border; posterior border of third submarginal cell twice length of anterior border,
anterior border 1.5 times as long as anterior
border of second submarginal cell.
Labrum imbricate and impunctate. Clypeus with small, faint punctures separated by
12 times a puncture width, integument between smooth. Supraclypeal area, face, vertex, gena, and postgena with sparse, small,
faint punctures, integument between smooth.
Pronotum smooth and impunctate. Mesoscutum and scutellum with sparse, faint punctures, integument between smooth. Pleura
107
with sparse, small punctures, integument between smooth. Propodeum smooth and impunctate. Corbicula smooth and impunctate.
Metasomal terga smooth and impunctate;
sterna finely imbricate and impunctate except
apical margins more strongly imbricate.
Where evident, integument dark brown
and shining. Wing membrane hyaline; veins
dark brown.
Pubescence whitish. Labrum with short,
scattered, erect setae. Clypeus with scattered,
short, simple, suberect setae intermixed with
scattered, minute, appressed, simple setae.
Face below level of antennal sockets with appressed, short, minutely plumose setae, setae
numerous but not obscuring integument; supraclypeal area, vertex, gena, and remainder
of face with scattered, suberect or erect, simple setae, setae of vertex and gena with minute branches. Postgena with scattered, short,
erect, simple setae. Pronotum with sparse,
short, suberect, simple setae except those of
dorsolateral angle and pronotal lobe dense,
moderately long, erect, and plumose setae.
Mesoscutum with scattered, short, erect, minutely plumose setae. Tegula with sparse, appressed, scattered, minutely plumose setae.
Scutellar setae as described for mesoscutum
except setae dense and elongate on posterior
border. Mesepisternum with moderately long,
scattered, erect, simple setae and intermixed
with scattered, minute, appressed, simple setae; hypoepimeral area without longer setae,
only with minute, appressed setae; metepisternum with setae as on mesepisternum except simple and distinctly more dense on dorsal half than on ventral half. Setae of lateral
and posterior surfaces of propodeum as described for ventral half of metepisternum except setae minute; basal area of propodeum
without pubescence. Corbicular setae minutely plumose, inner surface of metatibia
mostly covered by keirotrichiate field, field
separated from metatibial apex by length approximately equal to one-half its width; metabasitarsus with scattered, short, simple setae, posterior edge with longer setae, inner
surface with seven comb rows. Metasomal
terga with sparse, minute, appressed, simple
setae, slightly longer on more apical terga;
sterna with suberect to erect, short to moderately long, scattered setae, each with a few
minute branches, setae entirely postgradular.
108
MATERIAL: One specimen. Neotype (here

designated). Female, worker caste, MB.I.1943
(ZMHB) labeled: Museum fur Naturkunde
Berlin, Palaontologisches Museum, Inv. Nr.
MB.I.1943 // 50b [handwritten label] //
Neotype, Protobombus tristellus Cockerell,
desig. M. S. Engel.
Protobombus hirsutus (Cockerell),
new combination
Figure 77
Plate 5d
Chalcobombus hirsutus Cockerell, 1908b: 326.

Cockerell, 1909b: 12. Zeuner and Manning,
1976: 207.

P. indecisus but differs by the separation of
the keirotrichiate field from the metatibial
apex being equal to one-half of its apical
width and the posterior border of the third
submarginal cell only twice as long as the
anterior border.
wider than length (length 2.19 mm, width
mm; lower interorbital distance 1.19 mm. Interocellar distance 0.44 mm; ocellocular distance 0.34 mm; median to lateral ocellus 0.13
mm. Lower half of clypeus below lower tangent of compound eyes. Intertegular distance
1.88 mm. Basal vein confluent with cu-a,
straight; second abscissa Rs arched; 1m-cu
basad 1rs-m by six times vein width; 2rs-m
distad 2m-cu by vein width, strongly arched;
posterior border of second submarginal cell
approximately five times length of anterior
cell 1.75 times length of anterior border, anterior border slightly more than twice as long
as anterior border of second submarginal
cell; hind wing with eight distal hamuli arranged in a single, evenly spaced series; hind
wing with first abscissa M approximately
equal in length to second abscissa M1Cu.
Labrum imbricate and impunctate. Clypeus with sparse, small, faint punctures, integument between faintly imbricate. Supraclypeal area, face, vertex, gena, and postgena
with sparse, small, faint punctures, integument between smooth. Pronotum finely imbricate. Dorsum of mesosoma obscured by
NO. 259
Schimmel and sundry small items of debris.

Pleura impunctate finely imbricate. Corbicula
with remains presumably of pollen covering
surface. Metasomal terga and sterna finely
imbricate.
Where evident, integument dark brown
and shining. Wing membrane very lightly
fuscous; veins dark brown.
Pubescence whitish or slightly fuscous.
Labrum with short, scattered, erect, simple
setae. Clypeus with scattered, short, simple,
suberect setae intermixed with scattered, minute, appressed, simple setae. Face below
level of antennal sockets with appressed,
short, minutely plumose setae, setae numerous but not obscuring integument; supraclypeal area, vertex, gena, and remainder of face
with scattered, simple, suberect or erect setae, a few on vertex and gena with minute
branches. Postgena with scattered, short,
erect, simple setae. Pronotum with sparse,
short, suberect, simple setae except along
posterior border and covering pronotal lobe
with moderately long, erect, dense, plumose
setae. Pubescence of mesosomal dorsal surface difficult to interpret, those mesoscutal
setae appearing through layer of Schimmel
and debris appearing to be scattered, moderate-length, erect, and minutely plumose;
scutellar setae as described for mesoscutum
except setae dense on posterior border and
more elongate. Pleura with scattered, long,
erect, simple setae. Lateral and posterior surfaces of propodeum as described for pleura
except setae minute; basal area of propodeum without pubescence. Corbicular setae
simple, inner surface of metatibia covered by
keirotrichiate field, field separated from metatibial apex by length approximately equal
to one-half its width; metabasitarsus with
scattered, short, simple setae, posterior edge
with longer setae. Metasomal terga with
sparse, minute, appressed, simple setae; sterna with scattered, long, erect, fuscous, simple
setae along apical margins.
M ATERIAL : Four specimens. Neotype
(here designated). Female, worker caste, BJH 92 (AMNH) labeled: Baltic amber: Eocene, Kaliningrad, Yantarny // Neotype,
Chalcobombus hirsutus Cockerell, desig. M.
S. Engel // Protobombus hirsutus (Cockerell), det. M. S. Engel.
Non-type. Female, worker caste (CJVG)
2001
labeled: Protobombus hirsutus (Cockerell),

det. M. S. Engel.
(CHFG) labeled: Nr. 1254, Balt. Bernstein
// Protobombus hirsutus (Cockerell), det.
M. S. Engel.
(CHFG) labeled: Nr. 1255, Balt. Bernstein
// Protobombus hirsutus (Cockerell), det.
M. S. Engel.
Protobombus sp. indet.
MATERIAL: One specimen. Non-type. Female, Nr. 2608 (CCGG) labeled: Nr. 2608
// Protobombus sp. indet., det. M. S. Engel.
Protobombus sp. indet.
MATERIAL: One specimen. Non-type. Female, Nr. 1962.I.262 (BSPG): labeled Bayer. St. Slg. Palaont. Hist. Geol. (BSP), Nr.
1962.I.262, Apidae, Honigbiene, Unteroligozan, Ostpreussen // Protobombus sp. indet., det. M. S. Engel, 1999.
COMMENTS: This specimen is in exceedingly poor shape. The bee is almost completely pyritized. The general habitus, presence of a corbicula, and quadrangular metabasitarsus suggests a species of Protobombus. The identity of this specimen will
probably never be known with any reasonable degree of certainty.
Thaumastobombus, new genus
TYPE SPECIES: Thaumastobombus andreniformis Engel, new species.
DIAGNOSIS: This group superficially resembles in its general habitus a short-tongued
bee (either an andrenine, rophitine, or melittine) as well as the dwarf honey bees [Apini, Apis (Micrapis)] (fig. 78; pl. 5f). It, however, readily falls away from the former by
being a long-tongued bee and a corbiculate
apine at that. From the latter it differs in the
tribal characters [Apini lacks metatibial spurs
and outer mandibular grooves, has elongate
eye hairs, an elongate malar space, among
other features: refer to Engel (1999c) for a
diagnosis of the honey bees and their constituent groups; however, note that in the subgeneric diagnoses for Apis presented in En-
109
gel (1999c), references to the shape of the

first submarginal cell should refer to the third
submarginal cell]. Among electrapines,
Thaumastobombus is most similar to Protobombus, both sharing the quadrangular metabasitarsus. The former differs, however, by
the linear epistomal sulcus and weakly emarginate compound eyes that converge below.
apical teeth along upper half. Malar space
short, shorter than basal mandibular width.
Clypeus flat, not protuberant in lateral view.
Epistomal sulcus linear (fig. 80). Inner margins of compound eyes slightly converging
below, weakly emarginate (figs. 79, 80). F1
longer than F2, not longer than F23 combined; F3 longer than F2. Mesoscutal anterior border broadly rounded; median and parapsidal lines moderately impressed; scutellum projecting over metanotum and basal
area of propodeum. Basal area of propodeum
strongly declivitous (fig. 78). Keirotrichiate
field not elevated; metatibial spur pectinate
(figs. 83, 84); auricular basket dense, setae
long (fig. 81); rastellum strong and formed
of stiff setae (fig. 83); metabasitarsus about
as long as wide (fig. 81). Margin of pterostigma within marginal cell convex; marginal
cell apex offset from wing margin by slightly
less than pterostigma width, apex appendiculate (fig 85). Worker gonostyli minutely
setose at apices (fig. 78); sting not barbed
(except see Thaumastobombus cf. andreniformis).
is a combination of the Greek words thaumastos (meaning wonderful) and bombos
(meaning buzzing; a derivative of bombylios meaning bumble bee). The name is
masculine.
Thaumastobombus andreniformis,
new species
Figures 7885
Plate 5f

mm. Interocellar distance 0.45 mm; ocello-
110
NO. 259
Fig. 78. Left dorsolateral habitus of holotype female of Thaumastobombus andreniformis, new species. Scale bar 5 1 mm.
cular distance 0.25 mm; median to lateral

ocellus 0.14 mm. Distal third of clypeus set
below lower tangent of compound eyes. Intertegular distance 1.20 mm. Basal vein basad cu-a by vein width; 1rs-m distad 1m-cu
by seven times vein width; 2rs-m distad 2mcu by 2.5 times vein width; marginal cell
apex separated from wing margin by onehalf pterostigma width, feebly appendiculate;
first submarginal cell shorter than second and
third combined; second submarginal cell narrowed, anterior border just less than onethird length of posterior border, anterior border just less than one-half length of anterior
border of third submarginal cell; anterior
border of third submarginal cell just less than
one-third length of posterior border; seven
distal hamuli, arranged in a single, evenly
spaced series.
Integument of head and mesosoma smooth

and impunctate. Terga and sterna imbricate.
Coloration not well preserved, apparently
black and shining except antennae, metasoma, and legs dark brown; without maculations. Wing membrane hyaline; veins strong
and dark brown.
Pubescence generally pale. Labrum with
sparse, simple, erect setae. Setae of face numerous, simple, appressed, and short, each
with a few, minute branches, setae partially
obscuring integument, such setae intermixed
with sparse, erect, minutely branched setae;
setae most dense on face below level of antennal sockets and lateral to clypeus; appressed setae disappearing by vertex and
erect setae becoming gradually more numerous. Gena with appressed setae like those of
face; postgena with sparse, erect, moderate-
2001
Fig. 79.
111
Frontal view of head of holotype female of Thaumastobombus andreniformis, new species.
112
NO. 259
Sterna with dense, apical bands of moderate

to long, simple setae.
MATERIAL: One specimen. Holotype. Female, worker caste, B-JH 164 (AMNH) labeled: Baltic amber: Eocene, Kaliningrad,
Yantarny // Holotype, Thaumastobombus
andreniformis Engel.
ETYMOLOGY: The specific epithet is a reference to the superficial habitus of the species that resembles a short-tongued bee. The
species also resembles to some degree the
dwarf honey bees (A. florea and A. andreniformis), which also resemble short-tongued
bees.
Thaumastobombus cf. andreniformis
Fig. 80. Frontal view of head of holotype female of Thaumastobombus andreniformis, new
species. Scale bar 5 0.5 mm.
length, simple setae. Pubescence of mesoscutum as described for vertex. Tegula with a
few, minute, appressed, simple setae. Scutellar pubescence as on mesoscutum except setae along posterior border elongate and plumose. Pleura with dense, long, plumose setae, partially obscuring integument. Posterior
surface of metafemur with keirotrichiate
zone on apical two-thirds; inner surface of
metatibia with keirotrichiate zone not elevated but with narrow glabrous posterior margin
that is one-fifth width of keirotrichiate zone,
keirotrichiate zone separated from rastellum
and metatibial apex by glabrous zone that is
as long as width of keirotrichiate zone. Terga
with sparse setal bands along apical margin,
setae short and with a few, minute branches.
M ATERIAL : Two specimens. Non-type.

Two females, worker caste, Nr. 9 (CSUL) labeled: Thaumastobombus cf. andreniformis, det. M. S. Engel.
COMMENTS: These two, well-preserved individuals differ slightly in the shape of the
head and particularly in the structure of metatibial spur whereby the thickening of the
spur toward its apex present in the holotype
of T. andreniformis (fig. 84) is apparently absent in both of these individuals. This specimen also appears to have microscopic barbs
on the sting (perhaps also present in true T.
andreniformis?). These differences are likely
not great enough to deserve specific status
separated from T. andreniformis; however, I
hesitate to place them immediately into T.
andreniformis until more material of the genus is discovered. If these differences continue to be upheld or are found to be congruent
with additional, yet undiscovered, characters,
then specific status for this morph would be
warranted.
MELIKERTINI, New Tribe
TYPE GENUS: Melikertes Engel, 1998a.
DIAGNOSIS: This group most closely resembles species of the stingless bee tribe Me
Figs. 8184. Leg structures of holotype female of Thaumastobombus andreniformis, new species.
81. Outer surface of metatibia and metatarsus. 82. Strigil at protibia-probasitarsus junction. 83. Ventral,
slightly oblique view of metatarsus and metatibia (thus, outer surface of metatibia is visible while inner
surface of metabasitarsus is visible). 84. Magnified view of metatibial spur. Scale bars 5 0.5 mm (a 5
fig. 81; b 5 fig. 83; c 5 fig. 82).
2001
113
114
NO. 259
TABLE 9
Hierarchical Classification of Tribe

Melikertini

Thaumastobombus andreniformis, new species.
Scale bar 5 1 mm.
liponini. The general habitus of all species is

Trigona-like. The melikertines differ from
the Meliponini by the complete wing venation, presence of a single metatibial spur,
presence of a supraalar carina, toothed claws,
presence of an auricle, and presence of a
sting.
DESCRIPTION: Small to moderate size (ca.
38.5 mm long), slender, sparsely pubescent
bees. Mandible without outer mandibular
grooves. Labral width 34 times length.
Clypeus gently convex and weakly protuberant in profile. Compound eyes bare. Supraalar carina present; scutellum broadly rounded
posteriorly and variously produced (ranging
from projecting over metanotum and propodeum to not projecting over metanotum).
Claws of female with minute, inner tooth
(figs. 89, 90, 108); arolium strong and present (e.g., fig. 108); single, reduced metatibial spur present (figs. 90, 102, 107); malus
of strigilis with short anterior prong in addition to primary ventral velum; metabasitarsus with distinct auricle at base (figs. 89, 90,
106, 107); metatibia without penicillum. Distal venation of forewing strong and present
(figs. 91, 103, 109); marginal cell large and
narrowly rounded at apex, slightly offset
from wing margin, feebly appendiculate, cell
longer than distance from its apex to wing
apex; pterostigma present and moderate size,
much longer than prestigma, r-rs arising near
or just beyond midpoint, margin within marginal cell convex; 1m-cu angled; hind wing
with distinct jugal lobe, lobe broadly and
deeply incised (figs. 92, 104, 110); hamuli
reduced; wing membrane without alar papillae. Sting not reduced; apparently not barbed.
COMMENTS: This is a particularly diverse
group of Baltic amber bees and specimens

seem to be relatively commonly encountered.
Table 9 summarizes the classification of the
tribe as I have outlined it herein.
Key to Genera of Melikertini
1. Clypeal protrusion absent (e.g., figs. 100,
105) . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2
Clypeal protrusion present, i.e., base of clypeus produced into variously modified tubercles, tubercle bending upward over fronto-clypeal portion of epistomal sulcus and
supraclypeal area, ending between antennal
sockets (e.g., figs. 8688, 9398) . . . . .
. . . . . . . . . . . . . . . . . . . Succinapis, n. gen.
2. Scutellum bulging, overhanging metanotum
and propodeum; apical margins of metasomal terga distinctly lighter than remainder of metasoma, thus metasoma appears
banded; anterior and posterior margins of
metabasitarus distinctly converging toward
apex (fig. 107) . . . . . . . . . . . . . . . . . . . . 3
Scutellum not bulging, not overhanging metanotum or propodeum (fig. 101); metasomal terga uniformly colored; anterior and
posterior margins of metabasitarsus approximately parallel (fig. 102) . . . . . . . .
. . . . . . . . . . . . . . . . . . . . Melikertes Engel
3. Anterior margin of first submarginal cell inside marginal cell approximately equal to
length of anterior margin of second submarginal cell inside marginal cell (i.e., vein
r-rs as long as immediately succeeding abscissa of Rs) (fig. 109); basal vein confluent
with cu-a; compound eyes converging below (fig. 105); F2 distinctly shorter than F3,
F1 distinctly shorter than combined lengths
2001
of F23 (fig. 105) . . . . . . . . . . . . . . . . . .

. . . . . . . . . . . . . . . . . . . Melissites, n. gen.
Anterior margin of first submarginal cell inside marginal cell many times longer than
length of anterior margin of second submarginal cell inside marginal cell (i.e., vein
r-rs many times longer than immediately
succeeding abscissa of Rs); basal vein confluent with cu-a; compound eyes approximately parallel; F2 approximately equal in
length to F3, F1 approximately equal to
combined lengths of F23 . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . Roussyana Manning
Succinapis, new genus

TYPE SPECIES: Succinapis proboscidea Engel, new species.
DIAGNOSIS: This genus is most similar in
general habitus to Melikertes but differs by
the presence of a basal clypeal protrusion.
DESCRIPTION: Malar space short, shorter
than basal mandibular width. Epistomal sulcus forming obtuse angle; clypeus with basal
clypeal protrusion (i.e., base of clypeus
slightly extending outward and upward over
fronto-clypeal sulcus, typically covering supraclypeal area and ending in a tubercle between antennal sockets) (figs. 8688, 9398).
F1 longer than F2; F2 equal to F3. Inner margins of compound eyes roughly parallel. Preoccipital ridge rounded. Mesoscutal anterior
border broadly rounded; tegula tear-drop
shaped; scutellum not bulging, not projecting
over metanotum. Basal area of propodeum
nearly horizontal, not declivitous. Keirotrichiate field not elevated (fig. 90); metatibial
spur serrate or minutely ciliate; auricular basket sparse, setae short; rastellum strong and
formed of stiff setae (fig. 90); metabasitarsus
parallel-sided. Three submarginal cells (fig.
91). Metasomal terga not banded.
is a combination of the Latin words succineus (meaning amber) and apis (meaning
bee). The name is feminine.
COMMENTS: The peculiar clypeal protrusion is a unique feature among the corbiculate Apinae. At present the function of such
a structure in workers is illusive.
Key to Species of SUCCINAPIS
1. Clypeal protrusion extending above level of
antennal sockets by an antennal socket diameter or less (figs. 8688, 9395) . . . . 2
115
Clypeal protrusion extending dorsally above

level of antennal sockets by more than the
diameter of an antennal socket (figs. 96
98) . . . . . . . . . . . . S. proboscidea, n. sp.
2. Clypeal protrusion straight, apex not bent anteriorly, apex with short setae (figs. 8688);
mesepisternum smooth . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . S. goeleti, n. sp.
Clypeal protrusion bent anteriorly at apex,
apex with long setae (figs. 9395); mesepisternum punctured . . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . S. micheneri, n. sp.
Succinapis goeleti, new species

Figures 8692
Plate 7d
DIAGNOSIS: Succinapis goeleti is the most

primitive species of the genus as it is presently understood (see phylogenetic discussion below; fig. 99). The species can be distinguished from all other Succinapis by the
short clypeal protrusion that does not extend
above the level of the antennae by more than
the diameter of an antennal socket (figs. 86
88), the greatly narrowed second submarginal cell (fig. 91), and the impunctate pleura,
among other features.
mm). Clypeal protrusion short, not extending
dorsally above level of antennal sockets by
more than diameter of an antennal socket
(figs. 8688), protrusion not bent anteriorly.
Upper interorbital distance 0.92 mm; lower
interorbital distance 0.82 mm. Interocellar
distance 0.27 mm; ocellocular distance 0.32
mm; median to lateral ocellus 0.08 mm. Intertegular distance 0.78 mm. Basal vein basad cu-a by 2.5 times vein width; 1m-cu basad 2rs-m by five times vein width; 2rs-m
distad 2m-cu by 1.5 times vein width; first
submarginal cell shorter than second and
third combined; length of anterior border of
second submarginal cell one-tenth that of
posterior border (i.e., second submarginal
cell strongly narrowed anteriorly); length of
posterior border of third submarginal cell 1.5
times that of anterior border, anterior border
seven times length of anterior border of second submarginal cell; six distal hamuli, arranged in a single, evenly spaced series.
Clypeus impunctate and smooth. Face,
116
NO. 259
Figs. 8688. Head of holotype female of Succinapis goeleti, new species. 86. Frontal view. 87.
Lateral view. 88. Frontodorsal, oblique view. Scale bar 5 0.5 mm.
2001
117
Figs. 8990. Metatibia and metatarsus of holotype female of Succinapis goeleti, new species. 89.
Outer surface. 90. Inner surface. Scale bar 5 0.5 mm.
vertex, and gena with sparse, minute punctures, integument between smooth. Pronotum
with sparse, faint, minute punctures, integument between smooth. Mesoscutum and scutellum sculptured as on mesoscutum. Tegula
impunctate and smooth. Metanotum impunctate and faintly imbricate. Pleura impunctate
and smooth. Propodeum glabrous. Metasoma
impunctate and faintly imbricate.
Head, mesosoma, and metasoma black
without maculations; legs dark brown. Wing
brown.
Pubescence fuscous. Labrum with moderate-length, simple, erect setae scattered on
surface. Clypeus with sparse, minute, suberect, simple setae and short, simple, suberect, anteriorly-directed setae on dorsal-facing surface of clypeal protrusion. Face with
sparse, suberect, minute, simple setae. Setae
of vertex and gena as described for face.

Postgena with short, simple, erect setae. Pubescence of pronotum as described for face.
Mesoscutum with short, sparse, simple, erect
setae. Scutellum with erect, long (2.54 OD)
setae on posterior border, setae minutely plumose. Pleural pubescence as described for
face. Lateral surface of propodeum with pubescence as described for face; basal area of
propodeum without pubescence. Inner surface of metafemur and metatrochanter without pubescence except apical sixth of metafemur with dense field of keirotrichiae; inner
surface of metatibia with keirotrichiate field,
field separated by its width from metatibial
apex; corbicula bordered by long, minutelyplumose setae; metatarsus with simple, stout
setae; seven comb rows on inner surface of
metabasitarsus. Metasomal terga with sparse,
simple, short, appressed setae, slightly more
118
NO. 259
Figs. 9192. Wings of holotype female of Succinapis goeleti, new species. 91. Forewing. 92. Hind
wing. Scale bar 5 0.5 mm.
numerous laterally; sterna with scattered,

simple, erect (2.53 OD) setae.
MATERIAL: Two specimens. Holotype. Female, worker caste, B-JH 90 (AMNH) labeled: Baltic amber: Eocene, Kaliningrad,
Yantarny // Holotype, Succinapis goeleti
Engel.
Paratype. Female, worker caste, B-W 162
Kaliningrad, Yantarny // Paratype, Succinapis goeleti Engel.
ETYMOLOGY: The specific epithet is a patronymic honoring Mr. Robert G. Goelet,
Chairman Emeritus of the AMNH Board of
Trustees. Mr. Goelet has generously supported my work at the AMNH, while at the same
time greatly augmenting the collection of living and amber-fossil insects. It is only fitting
that one of these spectacular fossils bear his

name.
Succinapis micheneri, new species
Figures 9395
Plate 7c
Electrapis (Roussyana) palmnickenensis (Roussy); Zeuner and Manning, 1976: 233. [misidentification]
DIAGNOSIS: This species is similar to S.

goeleti in that the clypeal protrusion is relatively short (not extending above the level of
the antennae by more than the diameter of
an antennal socket) but differs in that the
apex of the process is bent anteriorly (fig.
9395). Moreover, the second submarginal
cell is not as strongly narrowed anteriorly
2001
(the anterior border is only one-fourth that of

the posterior border in S. micheneri by comparison to approximately one-tenth in S. goeleti) and the pleura are distinctly punctured.
mm). Clypeal protrusion short, not extending
dorsally above level of antennal sockets by
more than diameter of an antennal socket,
protrusion strongly bent anteriorly (figs. 93
95). Upper interorbital distance 0.83 mm;
Intertegular distance 0.75 mm. Basal vein basad cu-a by 1.5 times vein width; 1m-cu bisecting second submarginal cell; 2rs-m distad
2m-cu by two times vein width; first submarginal cell shorter than second and third
combined; length of anterior border of second submarginal cell one-fourth that of posterior border; length of posterior border of
third submarginal cell 1.5 times that of anterior border, anterior border twice as long as
that of second submarginal cell; six distal
hamuli, arranged in a single, evenly spaced
series.
Clypeus impunctate and smooth. Scape
with sparse, minute punctures, integument
between smooth. Face with minute punctures
separated by 24 times a puncture width, integument between smooth. Sculpturing of
vertex and gena as described for face. Postgena impunctate and smooth. Pronotum with
minute punctures separated by a puncture
width except medially punctures faint, integument between smooth. Mesoscutum with
faint, sparse, minute punctures, integument
between smooth. Scutellum sculptured as on
mesoscutum. Tegula with minute, sparse,
faint punctures, integument between smooth.
Pleura with small punctures separated by 1
2 times a puncture width except hypoepimeral area impunctate, integument otherwise
smooth between punctures. Sculpturing of
propodeal lateral surface as described for
mesepisternum; basal area of propodeum glabrous. Metasomal terga smooth and impunctate; sterna finely imbricate and impunctate.
Color, where evident, brown, without maculations. Wing membrane hyaline; veins
strong and brown.
119
Pubescence generally whitish. Labrum

with moderate-length, simple, erect setae
scattered on integumental surface. Clypeus
without setae except moderate-length, simple, suberect, anteriorly-directed setae on
dorsal-facing surface of clypeal protrusion.
Face with sparse, suberect, short, simple setae, setae more dense in lower paraocular
area but not obscuring integument. Setae of
vertex and gena as described for face except
minute. Postgena with short, simple, erect setae. Pubescence of pronotum as described for
face except setae of dorsal surface restricted
to posterior border and minute. Mesoscutum
with short, sparse, simple, erect setae. Scutellum with erect, long (13 OD) setae on
posterior quarter, setae each with several minute branches and slightly fuscous. Hypoepimeral area without setae; remainder of
pleura with pubescence as described for face.
Lateral surface of propodeum with pubescence as described for face; basal area of propodeum without setae. Inner surface of metafemur and metatrochanter without pubescence except apical quarter of metafemur
with dense field of keirotrichiae; inner surface of metatibia mostly covered by field of
keirotrichiae, field separated from metatibial
apex by length equivalent to its own width;
corbicula bordered by long, minutely-plumose setae; metatarsus with simple, stout setae; six comb rows on inner surface of metabasitarsus. Metasomal terga with scattered,
simple, short, appressed setae, sparse on central discs; sterna with scattered, simple, suberect setae that are short on S12 (0.51 OD)
then progressively longer on more apical
sterna (reaching 2.5 OD by S5).
MATERIAL: Twenty-one specimens. Holotype. Female, worker caste, B-JH 103
Kaliningrad, Yantarny // Holotype, Succinapis micheneri Engel.
Paratype. Female, worker caste, B-W 154
Kaliningrad, Yantarny // Paratype, Succinapis micheneri Engel.
Paratypes. Five females, worker caste
(CFEG) labeled: Paratype, Succinapis
micheneri Engel.
(CCGG) labeled: Paratype, Succinapis
micheneri Engel.
120
NO. 259
Figs. 9395. Head of holotype female of Succinapis micheneri, new species. 93. Fronal view. 94.
Ventrolateral, oblique view. 95. Dorsal view. Scale bars 5 0.5 mm (a 5 fig. 93; b 5 figs. 94, 95).
Paratypes. Eleven females, worker caste

(eight complete, three partials), Nr. 317
(CGHG) labeled: Nr. 317 // Paratype,
Succinapis micheneri Engel.
(GPUH) labeled: Nr. 80 (Scheele) // Erdbiene, Apidae, genus Andrena? // Paratype, Succinapis micheneri Engel. This is
the specimen misidentified as Electrapis
(Roussyana) palmnickenensis by Zeuner and
2001
Manning (1976: see Comments under Roussyana palmnickenensis, below).

Paratype. Female, worker caste (PMUZ)
labeled: Paratype, Succinapis micheneri
Engel.
ETYMOLOGY: The specific epithet is a patronymic honoring my mentor and friend, Dr.
Charles D. Michener, worlds authority on
bees.
Succinapis proboscidea, new species
Figures 9698
Plate 7b
DIAGNOSIS: This is the most distinctive of

all the Succinapis species. The grossly enlarged clypeal protrusion that is both bent anteriorly as well as extending above the level
of the antennae by more than the diameter of
an antennal socket (figs. 9698) serves to immediately differentiate S. proboscidea from
both S. micheneri and S. goeleti.
mm). Clypeal protrusion greatly enlarged,
extending above level of antennal sockets by
more than diameter of an antennal socket,
protrusion bent anteriorly (figs. 9698). Upper interorbital distance 0.98 mm; lower interorbital distance 0.93 mm. Interocellar distance 0.40 mm; ocellocular distance 0.30
mm; median to lateral ocellus 0.10 mm. Intertegular distance 0.98 mm. Basal vein basad cu-a by vein width; 1m-cu bisecting second submarginal cell; 2rs-m distad 2m-cu by
two times vein width; first submarginal cell
shorter than second and third combined;
length of anterior border of second submarginal cell one-fourth that of posterior border;
length of posterior border of third submarginal cell 1.75 times that of anterior border,
anterior border two times length of that of
second submarginal cell; six distal hamuli,
arranged in a single, evenly spaced series.
Clypeus with sparse, minute punctures, integument between smooth; anterior-facing
surface of clypeal protrusion with a few
weak longitudinal striae. Scape with minute
punctures separated by a puncture width, integument between smooth. Face with minute
width, integument between smooth; punc-
121
tures becoming gradually more faint and

more sparse toward vertex and on gena.
by a puncture width except medially punctures faint, integument between smooth. Mesoscutum mostly obscured by a thin layer of
Schimmel; where evident, integument with
faint, minute punctures separated by 12
smooth. Scutellum sculptured as on mesoscutum. Tegula with minute, faint punctures
on outer half, otherwise integument smooth.
Metanotum obscured from view by fracture
planes and flows from scutellum. Pleura with
puncture width, becoming faint on central
disc of mesepisternum and separated by 24
times a puncture width, and hypoepimeral
area with faint and exceedingly sparse punctures, integument of pleura smooth between
punctures. Propodeal lateral surface with minute, faint punctures separated by 23 times
smooth; basal area of propodeum glabrous.
Metasoma faintly imbricate and impunctate.
Color, where evident, dark brown, without
Pubescence generally whitish. Labrum
with moderate-length, simple, erect setae
widely scattered over surface. Clypeus with
minute, suberect, simple setae restricted to
lateral borders and short, simple, suberect,
anteriorly-directed setae on dorsal-facing
surface of clypeal protrusion. Face with
sparse, suberect, minute, simple setae, such
setae becoming slightly more dense in lower
paraocular area near clypeal lateral border.
Setae of vertex and gena as described for
face. Postgena with short, simple, erect setae.
Pubescence of pronotum as described for
face except setae of dorsal surface restricted
to posterior border. Mesoscutum with short,
sparse, simple, erect setae evident through
thin layer of Schimmel. Scutellum with erect,
long (13 OD) setae on posterior third, setae
each with several minute branches. Hypoepimeral area without setae; remainder of
pleura with pubescence as described for face.
Lateral surface of propodeum with pubescence as described for face; basal area of propodeum without setae. Inner surface of metafemur and metatrochanter without pubes-
122
NO. 259
Figs. 9698. Head of holotype female of Succinapis proboscidea, new species. 96. Frontal view.
97. Oblique, lateral view. 98. Dorsal view. Scale bar 5 0.5 mm.
2001
cence except apical quarter of metafemur

with dense field of keirotrichiae; inner surface of metatibia with keirotrichiate zone;
corbicula bordered by long, minutely-plumose setae; metatarsus with simple, stout setae; six comb rows on inner surface of metabasitarsus. Metasomal terga with scattered,
simple, short, appressed setae, sparse on central discs; sterna with scattered, simple, erect,
slightly fuscous setae.
Female, worker caste, B-JH 96 (AMNH) labeled: Baltic amber: Eocene, Kaliningrad,
Yantarny // Holotype, Succinapis proboscidea Engel.
Paratype. Female, worker caste, B-JH 99
Kaliningrad, Yantarny // Paratype, Succinapis proboscidea Engel.
(CSUL) labeled: Paratype, Succinapis proboscidea Engel.
ETYMOLOGY: The specific epithet is derived from the Greek word proboskidos, referring to the trunk of an elephant, and is a
reference to the structure of the large clypeal
protrusion of this species.
123
applies if the polarities are reversed for each

of the discussed characters. Fortunately, the
sculpturing of the mesepisternum provides
some information that can be compared to
the related melikertine genera and for polarizing the species in Succinapis. The mesepisternum in all species of Melikertes is
smooth and impunctate. Within Succinapis
smooth pleural integument is only found in
S. goeleti, while both S. micheneri and S.
proboscidea have punctured mesepisterna. It
would therefore appear that S. goeleti is the
most plesiomorphic species of the genus by
comparison to Melikertes. This hierarchy polarizes the clypeal protrusion morphology
(fig. 99). The shortened clypeal protrusion
that does not project more than an antennal
socket diameter above the antennal sockets
observed in S. goeleti and S. micheneri is
presumably plesiomorphic. The apical bend
of the clypeal protrusion is an apomorphic
feature uniting S. micheneri and S. proboscidea, while the greatly enlarged process is
an autapomorphy of S. proboscidea (fig. 99).
Genus Melikertes Engel, status novus
Internal Phylogeny of Succinapis
Electrapis (Melikertes) Engel, 1998a: 95. Type

species: Electrapis (Melikertes) stilbonota Engel, 1998a, original designation.
The morphology of the clypeal protrusion

suggests a hierarchy of relationship among
the species of Succinapis (fig. 99). By comparison to other melikertine genera as outgroups it is possible to build a preliminary
cladistic hypothesis for the three known Succinapis species. The presence of the clypeal
protrusion is a unique apomorphy among the
corbiculate Apinae and nicely serves to support the monophyly of the genus. The polarity of the different morphotypes of clypeal
protrusions is impossible to determine in isolation from other characters as there is no
corresponding structure in the outgroups.
The apically bent clypeal protrusions of S.
micheneri and S. proboscidea could be interpreted as plesiomorphic with S. goeleti and
S. micheneri perhaps clustered on the basis
of shortened protrusion and S. goeleti autapomorphic for a straight process, or if the
polarity of the protrusions size is removed,
then our three-taxon statement collapses to a
trichotomy. The same scenario of difficulties
DIAGNOSIS: This genus is similar to Succinapis but lacks the basal clypeal protrusion.
than basal mandibular width. Epistomal sulcus forming obtuse angle or nearly linear
(e.g., M. stilbonotus: fig. 100); clypeus without basal clypeal protrusion (fig. 100). F1
longer than F2; F2 equal to F3. Inner margins of compound eyes roughly parallel. Preoccipital ridge rounded or weakly carinate.
Mesoscutal anterior border broadly rounded;
tegula tear-drop shaped; scutellum not bulging, not projecting over metanotum (fig.
101). Basal area of propodeum nearly horizontal, not declivitous. Keirotrichiate field
not elevated (fig. 102); metatibial spur serrate
or minutely ciliate; auricular basket sparse,
setae short; rastellum strong and formed of
stiff setae (fig. 102); metabasitarsus parallelsided (fig. 102). Three submarginal cells (fig.
103). Metasomal terga not banded.
COMMENTS: I previously (Engel, 1998a) erroneously reported an absence of metatibial
124
Fig. 99.
NO. 259
Phylogeny of Succinapis based on morphology of the clypeal protrusion.
spurs for Melikertes. Subsequent study of the

type material for M. stilbonotus as well as
newly discovered material of other melikertine bees has revealed a single, reduced metatibial spur. This structure is difficult to observe in the holotype and paratype of M. stilbonotus but is distinctly present (fig. 102).
Key to Species of MELIKERTES
1. Tegula without posterior lamella, oval or teardrop shaped; body size small (ca. 34.5
mm) . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2
Tegula with short, posterior lamella; body
size moderate (ca. 8 mm) . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . M. proavus (Menge)
2. Apical margin of clypeus unmodified,

straight, and flat as on remainder of clypeus
. . . . . . . . . . . . . . . M. stilbonotus (Engel)
Apical margin of clypeus strongly projecting
outward from face with deep medial cleft,
remainder of clypeus relatively flat . . . .
. . . . . . . . . . . . . . . . . . M. clypeatus, n. sp.
Melikertes proavus (Menge),

new combination
Apis proava Menge, 1856: 26.
Electrapis (Roussyana) proava (Menge); Zeuner
and Manning, 1976: 236.
Trigona (Roussyana) proava (Menge); Kerr and
Cunha, 1976: 39.
2001
125
Figs. 100101. Holotype female of Melikertes stilbonotus (Engel). 100. Frontal view of head. 101.
Dorsal view. Scale bars 5 0.5 mm (a 5 fig. 101; b 5 fig. 100).
Electrapis (Melikertes) proava (Menge); Engel,
1998a: 95.
DIAGNOSIS: This is the largest and least understood species of Melikertes (simply owing
to the fact that the two specimens known are
in poor condition by comparison to material
of other species). The species is exceptional
for its large size compared to other Melikertes.
8.30 mm; forewing length 6.0 mm. Apical
margin of clypeus straight, not bent apically.
Intertegular distance 2.15 mm. Tegula with
short, posterior lamella. Basal vein basad cua by two times vein width; 1m-cu bisecting
second submarginal cell; 2rs-m distad 2m-cu;
first submarginal cell only slightly shorter
than second and third combined; second submarginal cell narrowed anteriorly, not very
strongly so; length of anterior border of third
submarginal cell half of that of posterior border, anterior border about as long as that of
anterior border of second submarginal cell;

six distal hamuli, arranged in a single, evenly
spaced series.
Integument of head, mesosoma, and metasoma apparently smooth and impunctate
except metanotum rugulose and sterna imbricate.
dark brown to black and shining, without
Pubescence generally pale. Setae of face
and vertex widely scattered, simple, and
short. Gena with simple, short, suberect setae
very sparsely scattered; postgena with long,
simple setae sparsely scattered over integument. Pronotal collar without pubescence;
pronotal border with mesoscutum with short,
simple setae; lateral surface with similar minute setae. Mesoscutum with scattered, minute setae, more sparsely scattered on central
disc. Pubescence of scutellum as described
126
Fig. 102. Inner surface of metatibia and metatarsus of holotype female of Melikertes stilbonotus (Engel).
NO. 259
Paralectotype, Transf. [transferred] from

Zool. Dept. [Zoology Department], 15 Dec.
[December] 1904, (In. 18757), 9214 [74?]
// Neotype, Apis proava Menge, 1856, desig. ICZN (M. S. Engel). Formerly the paralectotype designated by Zeuner and Manning (1976: 236; see Comments below); designated as neotype by ICZN Opinion Nr.
1964 (ICZN, 2000). The amber piece is set
into Canadian balsam and mounted onto a
microscope slide.
Non-type. Female, worker caste, In.
43592 (BMNH) labeled: In. 43592, E. proava, Lectotype // M. proava? (Menge),
non-type, desig. ICZN (M. S. Engel) //
Melikertes proavus? (Menge), det. M. S.
Engel. Formerly lectotype; designation by
Zeuner and Manning (1976: 236; see Comments below). This individual is assigned to
M. proavus with great hesitation owing to its
extremely poor condition. Zeuner and Manning (1976), who examined the specimen before it was damaged, considered it to be conspecific with the above specimen.
COMMENTS: The type designations of Zeuner and Manning (1976) have been suppressed owing to the poor condition of their
chosen lectotype (Engel, 1999a; ICZN,
2000).
Melikertes stilbonotus (Engel),
new combination
Figures 100104
for mesoscutum. Hypoepimeral area without

pubescence; remainder of mesepisternum
with widely scattered, minute setae. Basal
area and posterior surface of propodeum
without pubescence. Inner surface of metafemur and metatrochanter without pubescence except apical quarter of metafemur
with dense field of keirotrichiae; inner surface of metatibia with keirotrichiate zone. T1
without pubescence on central disc, a few,
simple setae on lateral borders; pubescence
of T2T6 as described for T1 except a few
simple setae on posterolateral borders; sterna
with sparse, simple, subappressed setae.
MATERIAL: Two specimens. Neotype. Female, worker caste, Nr. 13781 (BMNH) labeled: No. 13781, Bee, Museum Stantien &
Becker // 515, Dr. R. Klebs, 9214 [74?]
// XII B781 [on the amber block] // Electrapis proava (Menge), Amber, East Prussia,
Electrapis (Melikertes) stilbonota Engel, 1998a:

95.
DIAGNOSIS: Refer to diagnosis for M. clypeatus (see below).

wider than long (length 1.12, width 1.32:
measurements from paratype). Upper interorbital distance 0.86 mm; lower interorbital
distance 0.82 mm (both metrics from paratype). Apical margin of clypeus straight, not
bent apically. Intertegular distance 0.82 mm.
Basal vein basad cu-a by two times vein
width; 1m-cu bisecting second submarginal
cell; 2rs-m distad 2m-cu by 1.5 times vein
width; first submarginal cell shorter than second and third combined; length of anterior
border of second submarginal cell one-tenth
that of posterior border (i.e., second submar-
2001
127
Figs. 103104. Wings of holotype female of Melikertes stilbonotus (Engel). 103. Forewing. 104.
Hind wing. Scale bar 5 0.5 mm.
ginal cell strongly narrowed anteriorly);

cell half of that of posterior border, just over
three times length of anterior border of second submarginal cell; six distal hamuli, arranged in a single, evenly spaced series; cua of hind wing orthogonal to M1Cu.
Integument of head, mesosoma, and metasoma smooth and impunctate except metanotum rugulose. S36 weakly and sparsely
nodulate, integument between smooth.
dark brown to black and shining, without
widely scattered, simple, and short; such
hairs becoming gradually longer toward vertex. Gena with simple, short, suberect setae;
postgena with long, simple setae sparsely
scattered over integument. Pronotal collar

without pubescence; pronotal border with
mesoscutum with short, simple setae; lateral
surface with similar minute setae, such setae
appressed but not obscuring integument. Mesoscutum with scattered, simple setae, more
sparsely scattered and shorter on central disc,
those setae on anterolateral borders with a
few short branches. Pubescence of scutellum
as described for mesoscutum except longer
and restricted to posterior border. Metanotum
with dense, minute, simple setae, not obscuring integument. Hypoepimeral area without pubescence; remainder of mesepisternum
with scattered, simple setae, such setae gradually become longer ventrally; central disc,
however, without pubescence. Basal area and
posterior surface of propodeum without pubescence; lateral surface with scattered, long,
simple setae intermixed with shorter, ap-
128
pressed setae, such setae partially obscuring

integument. Pubescence of legs generally
simple and scattered except inner surfaces of
mesotrochanter and mesofemur without pubescence and outer surface of mesotibia with
dense, branched setae; inner surface of metafemur and metatrochanter without pubescence except apical quarter of metafemur
with dense field of keirotrichiae; inner surface of metatibia with keirotrichiate zone;
eight comb rows on inner surface of metabasitarsus, each composed of stiff, elongate,
simple setae; outer surface with scattered,
long, simple setae. T1 without pubescence on
central disc, a few, simple setae on lateral
borders; pubescence of T2 as described for
T1 except a few simple setae on posterolateral borders; T36 with simple setae longer
than those of T12, sparsely scattered over
central disc, more concentrated on lateral
margins; sterna with sparsely scattered, simple setae.
MATERIAL: Five specimens. Holotype. Female, worker caste, In. 17778 (BMNH). The
specimen bears labels reading Holotype,
Electrapis stilbonota, desig. M. S. Engel [red
label] // Amber Inclusa, Hymenoptera, In.
17778, Samland, No. 67 [white label]. Amber piece preserved in Canadian balsam on a
slide mount.
Paratype. Female, worker caste, In. 17778
(BMNH); same block of amber as holotype.
Paratype total body length 3.72 mm; forewing length 3.16 mm. Refer to Engel
(1998a) for more measurements.
Kaliningrad, Yantarny // Melikertes stilbonotus (Engel), det. M. S. Engel.
Kaliningrad, Yantarny // Melikertes stilbonotus (Engel), det. M. S. Engel.
(CJDL) labeled: Nr. 63 // Melikertes stilbonotus (Engel), det. M. S. Engel.
Melikertes clypeatus, new species
Plate 7a

M. stilbonotus, the two species looking remarkably similar. Melikertes clypeatus, how-
NO. 259
ever, is immediately recognizable by the peculiar modification of the clypeal apical margin that is strongly flexed forward below the
level of the lower tangent of the compound
eyes. Moreover, the projecting portion of the
clypeal apical margin possesses a strong medial cleft.
Apical margin of clypeus strongly bent apically, with distinct medial cleft. Intertegular
distance 0.83 mm. Basal vein basad cu-a by
two times vein width; 1rs-m distad 1m-cu by
six times vein width; 2rs-m distad 2m-cu by
three times vein width; first submarginal cell
shorter than second and third combined;
length of anterior border of second submarginal cell one-third that of posterior border;
cell half of that of posterior border, twice
length of anterior border of second submarginal cell; five distal hamuli, arranged in a
Integument of head, mesosoma, and metasoma smooth and impunctate.
Coloration not preserved. Wing membrane
hyaline; veins strong and brown.
sparse, simple, suberect, and short, although
such setae becoming erect on vertex. Mesoscutum with sparse, simple, short setae. Pubescence of scutellum as described for mesoscutum except longer on posterior border.
Pleura with sparse, simple, short setae. Pubescence of legs generally simple and scattered except inner surfaces of mesotrochanter
and mesofemur without pubescence and outer surface of mesotibia with dense, branched
setae; inner surface of metafemur and metatrochanter without pubescence except apical
quarter of metafemur with dense field of keirotrichiae; inner surface of metatibia with
keirotrichiate zone; comb rows on inner surface of metabasitarsus, each composed of
stiff, elongate, simple setae; outer surface
with scattered, long, simple setae. Terga and
sterna with exceedingly sparse, minute, simple, appressed or subappressed setae.
2001
129
M ATERIAL : Holotype. Female, worker

caste, B-JH 77 (AMNH) labeled: Baltic
Holotype, Melikertes clypeatus Engel.
The only known specimen has a great deal
of Schimmel on the mesosoma that entirely
obscures the propodeum and metanotum.
The scutellum, mesoscutum, tegula, and
pleura can be seen with some difficulty. The
posterior portion of the head is also covered
with some Schimmel, as are lateral portions
of T1.
ETYMOLOGY: The specific epithet is a reference to the modified apical margin of the
clypeus.
Melissites, new genus
TYPE SPECIES: Melissites trigona Engel,
new species.
DIAGNOSIS: See Diagnosis for Roussyana
(below).
than basal mandibular width (fig. 105). Epistomal sulcus forming obtuse angle (fig.
105); clypeus without basal clypeal protrusion. F1 longer than F2; F2 shorter than F3.
Compound eyes strongly converging below
and emarginate (fig. 105). Preoccipital ridge
rounded. Mesoscutal anterior border broadly
rounded; tegula oval; scutellum bulging, projecting over metanotum and basal area of
propodeum. Basal area of propodeum declivitous. Keirotrichiate field not elevated (fig.
107); metatibial spur ciliate; auricular basket
sparse, setae short (fig. 106); rastellum strong
and formed of stiff setae; lateral margins of
metabasitarsus converging toward apex (figs.
106, 107). Three submarginal cells; anterior
border of second submarginal cell as long as
r-rs (fig. 109). Metasomal terga banded.
is a combination of melissa (Greek, meaning
bee) and the suffix -ites (Greek, meaning
nature of or likeness, a common suffix
for fossil genera). The name is feminine.
Melissites trigona, new species
Figures 105110
Plate 6d

Fig. 105. Frontal view of holotype female

head of Melissites trigona, new species. Scale bar
5 0.5 mm.

Intertegular distance 1.00 mm. Basal vein
confluent with cu-a; 1rs-m distad 1m-cu by
eight times vein width; 2rs-m distad 2m-cu
by 1.5 times vein width; first submarginal
and third submarginal cells; length of posterior border of second submarginal cell
slightly less than twice anterior border;
cell three-quarters that of posterior border,
1.25 times length of anterior border of second submarginal cell; six distal hamuli arranged in a single, evenly spaced series (fig.
110).
Integument of head and mesosoma smooth
and impunctate. Terga and sterna imbricate
and impunctate.
Head and mesosoma black. Antennae,
legs, and metasoma dark brown except apical
margins of terga light brown to pale yellow.
130
NO. 259
Figs. 106108. Leg structures of holotype female of Melissites trigona, new species. 106. Outer
surface of metatibia and metabasitarsus. 107. Inner surface of metatibia and metabasitarsus. 108. Claw
and arolium. Scale bar 5 0.5 mm (figs. 106, 107); 0.25 mm (fig. 108).

dark brown.
Pubescence lightly infuscated. Setae of
face widely scattered, simple, and short; such
setae longer on vertex. Gena with simple,
short, appressed setae; postgena with long,
simple setae sparsely scattered. Mesoscutum
with sparse, simple setae, more sparsely scattered and shorter on central disc. Pubescence
of scutellum as described for mesoscutum
except two or more times longer, plumose,
and dense on posterior half. Metanotum
without pubescence. Hypoepimeral area
without pubescence; remainder of mesepisternum and metepisternum with scattered,
simple, long setae. Basal area and posterior
surface of propodeum without pubescence;
lateral surface with scattered, minute, suberect, simple setae. Pubescence of legs generally simple and scattered except inner surfaces of mesotrochanter and mesofemur
without pubescence and outer surface of mesotibia with dense, branched setae; inner surface of metafemur and metatrochanter without pubescence except apical quarter of metafemur with dense field of keirotrichiae; inner surface of metatibia with keirotrichiate
zone; corbicular setae simple; seven comb
rows on inner surface of metabasitarsus, each
composed of stiff, elongate, simple setae;
outer surface with scattered, long, simple setae. Terga and sterna with short, simple,
sparse setae.
Yantarny // Holotype, Melissites trigona
Engel.
ETYMOLOGY: The specific epithet is taken
(as a noun in apposition) from the genusgroup name Trigona (Meliponini) and is a
2001
131
Figs. 109110. Wings of holotype female of Melissites trigona, new species. 109. Forewing. 110.
Hind wing. Scale bars 5 1 mm (a 5 fig. 109; b 5 fig. 110).
reference to the overall, superficial resemblance of this species to some stingless bees.
Genus Roussyana Manning, status novus
Electrapis (Roussyana) Manning, 1960: 306.
Type species: Apis palmnickenensis Roussy,
1937, original designation. Zeuner and Manning, 1976: 233.
Russyana Petrov, 1992: 361. Lapsus calami.
DIAGNOSIS: Roussyana most closely resembles Melissites; both genera have the distinctive banding of the metasoma owing to
the apical margins of the terga being very
pale brown in comparison to the remainder
of the terga, which are dark brown. The genus can be separated from Melissites by the
strongly narrowed second submarginal cell,
basal vein basal to cu-a, and the approximately parallel compound eyes.
than basal mandibular width. Epistomal sulcus forming obtuse angle; clypeus without
basal clypeal protrusion. F1 longer than F2;

F2 equal to F3. Inner margins of compound
eyes roughly parallel. Preoccipital ridge
rounded. Mesoscutal anterior border broadly
rounded; tegula oval; scutellum bulging, projecting over metanotum and basal area of
propodeum. Basal area of propodeum declivitous. Keirotrichiate field not elevated; metatibial spur serrate; auricular basket sparse,
setae short; rastellum strong and formed of
stiff setae; lateral margins of metabasitarsus
converging toward apex. Three submarginal
cells; anterior border of second submarginal
cell much shorter than r-rs. Metasomal terga
banded.
Roussyana palmnickenensis (Roussy)
Apis palmnickenensis Roussy, 1937: 66.
Electrapis (Roussyana) palmnickenensis (Roussy); Manning, 1960: 306. Zeuner and Manning,
1976: 233 [misidentification: see Succinapis
micheneri, above]
132
Electrapis minuta Kelner-Pillault, 1970a: 16. NEW

SYNONYMY.
Trigona (Roussyana) palmnickenensis (Roussy);
Kerr and Cunha, 1976: 39.
Roussyana palmuickenensis Petrov, 1992: 361.
Lapsus calami.

3.08 mm; forewing length 2.3 mm. Head longer than wide (length 0.93 mm, width 0.88
mm). Mandible with a single small tooth on
upper one-fifth of apical margin. Intertegular
distance 0.82 mm. Basal vein basad cu-a by
two times vein width; 1rs-m distad 1m-cu by
seven times vein width; 2rs-m distad 2m-cu
by two times vein width; first submarginal
and third submarginal cells; second submarginal cell strongly narrowed anteriorly, anterior border shorter than r-rs; anterior border
of third submarginal cell approximately nine
times longer than anterior border of second
submarginal cell; six distal hamuli arranged
in a single, evenly spaced series.
Head and mesosoma smooth and impunctate except corbicula finely imbricate. Terga
and sterna finely and faintly imbricate except
lighter apical margins of terga glabrous.
Head and mesosoma black. Antennae,
legs, tegulae, and metasoma dark brown except apical margins of terga light brown.
Pubescence generally pale. Labrum with
several, long, simple setae widely scattered.
Setae of face widely scattered, simple, and
short, although setae becoming slightly longer on vertex. Mesoscutum with scattered,
long, simple setae. Pubescence of scutellum
as described for mesoscutum except somewhat longer. Metanotum without pubescence.
Hypoepimeral area without pubescence; remainder of mesepisternum with scattered,
simple setae, although particularly sparse on
central disc; setae slightly longer ventrally.
Basal area of propodeum without pubescence. Pubescence of legs generally simple
and scattered except inner surfaces of mesotrochanter and mesofemur without pubescence and outer surface of mesotibia with
dense, branched setae; inner surface of metafemur and metatrochanter without pubescence; inner surface of metatibia with keirotrichiate zone; distinct comb rows on inner
surface of metabasitarsus, each composed of
NO. 259
stiff, elongate, simple setae; outer surface

with scattered, long, simple setae. T15 with
sparse, minute, suberect, simple setae; T6
with numerous, short, simple setae uniformly
covering surface; sterna with sparsely scattered, short, simple setae and with dense,
subapical rows of long setae.
M ATERIAL : Three specimens. Neotype
(palmnickenensis; here designated). Female, worker caste, NB.I.1945 [Berendt Collection] (ZMHB) labeled: Neotype, Apis
palmnickenensis Roussy, desig. M. S. Engel // 50a [Berendt handwriting] //
NB.I.1945. The types for Roussys species
were in his private collection. A diligent
search for their whereabouts failed to locate
them. I have thus here designated a neotype
for the species. The limited description Roussy (1937) presented for the species, however,
best agrees to Kelner-Pillaults specimens
among all of the corbiculate bees in Baltic
amber. I, therefore, believe Roussys and
Kelner-Pillaults specimens to be conspecific
and have chosen the most completely preserved of the two individuals in the piece to
serve as the neotype for Apis palmnickenensis Roussy as well as the lectotype for Electrapis minuta Kelner-Pillault. The two paralectotype individuals (designated below) are
clustered together on one end of the piece,
while the neotype/lectotype is alone on the
opposite end.
Lectotype (minuta; here designated). Female, worker caste, NB.I.1945 (ZMHB) labeled: Lectotype, Electrapis minuta KelnerPillault, desig. M. S. Engel // Electrapis
minuta S.K.P., syntypes, S. Kelner-Pillault,
det. [Kelner-Pillault handwriting] // Roussyana minuta (Kelner-Pillault), det. M. S.
Engel, 1999. Since this is the same specimen designated above as the neotype for A.
palmnickenensis, the remaining labels are
identical to those listed above for A. palmnickenensis (refer to preceding paragraph).
Paralectotypes (minuta; here designated). Two females, worker caste, NB.I.1945
(ZMHB): these specimens are in the same
amber block as the lectotype and bear the
same labels. They are distinguished from the
lectotype by the poorer state of preservation,
both are heavily covered in Schimmel and
surrounded by numerous small fracture
planes.
2001
COMMENTS: The specimen figured as a

non-type of Roussyana palmnickenensis by
Zeuner and Manning (1976: their pl. 3 as
Electrapis palmnickenensis) is not actually a
specimen of Roussyana. The specimen is in
GPUH and upon examination it is actually S.
micheneri. The specimen lacks the defining
features of not only the species but also the
genus. Moreover, the specimen has a distinct
clypeal protrusion (a character of Succinapis)
and sparse mesoscutal setae. A thin layer of
pollen and mold on the specimen was apparently interpreted as copious pubescence by
Zeuner and Manning. The description they
presented for R. palmnickenensis, therefore,
does not apply to this species.
Kelner-Pillault (1970a) designated the two
specimens discussed above as syntypes but
did not select one to be the name-bearing
type. I have, therefore, in the interest of nomenclatural stability, selected a lectotype and
paralectotype from her syntypes. Kelner-Pillault (1970a) enigmatically figured the hind
wing without a jugal lobe, which is not only
present but deeply incised and visible in the
specimens.
Tribe MELIPONINI Lepeletier
de Saint Fargeau
Meliponites Lepeletier de Saint Fargeau, 1836:
407. Type genus: Melipona Illiger, 1806.
Trigonini Moure, 1946: 611. Type genus: Trigona
Jurine, 1807.
Lestrimelittini Moure, 1946: 611. Type genus:
Lestrimelitta Friese, 1903.
DIAGNOSIS: Among living taxa the Meliponini are allied to the honey bees (tribe Apini); both tribes share the complete loss of
metatibial spurs and outer mandibular
grooves and possess a jugal lobe in the hind
wing. The meliponines differ from Apini by
the absence of an auricle, absence of an inner
tooth on the claw, the reduced forewing venation, and the reduction of the sting, among
other characters. Numerous ethological traits
similarly serve to differentiate the tribes (reviewed by Michener, 1990: as subfamilies).
Among the fossil taxa meliponines most
closely resemble the tribe Melikertini but differ notably by the aforementioned characters
as well as the complete loss of metatibial
spurs (melikertines retain a single metatibial
spur).
133

Kelneriapis eocenica (Kelner-Pillault). Scale bar
5 0.5 mm.
DESCRIPTION: Minute to moderate size (ca.

1.513 mm long), sparsely to moderately pubescent bees. Mandible without outer mandibular grooves. Labral width three to four
times length. Clypeus variously produced,
typically gently convex and not protuberant
in profile. Compound eyes typically bare. Supraalar carina absent; scutellum broadly
rounded posteriorly and variously produced
(i.e., ranging from projecting over metanotum and propodeum to not projecting at all:
e.g., see Michener, 1990). Claws of female
simple (e.g., fig. 115); arolium strong and
present; metatibial spurs absent; malus of
strigilis without anterior velum; metabasitarsus without auricle; metatibia with penicillum (e.g., fig. 114). Distal venation of forewing weakened (e.g., figs. 111, 113); marginal cell apex typically open (e.g., figs. 111,
113); pterostigma present, moderate to large
in size, much longer than prestigma, r-rs arising near midpoint, margin within marginal
cell convex; 1m-cu, when present, angled;
hind wing with distinct jugal lobe, lobe
broadly and deeply incised; hamuli reduced;
wing membrane without alar papillae. Sting
reduced.
COMMENTS: The tribe Meliponini contains
the familiar stingless bees. The group is today worldwide in the tropics, with a particularly high diversity in the neotropics. Michener (2000a) recognized 23 Recent genera
and subgenera worldwide. Although no meliponines today occur in Europe, at least two
species were present in the middle Eocene
fauna of this region. Both seem closely allied
to Recent sub-Saharan African genera.
134
Key to Genera of Meliponini

in Baltic Amber
1. Apical bend in M (at point where vein would
meet 1m-cu) present (fig. 111); antennal
sockets at least one antennal socket diameter above basal clypeal margin; posterior
corner of corbicula weakly rounded; scutellum projecting over metanotum and propodeum; basal area of propodeum shorter
than scutellum and declivitous . . . . . . .
. . . . . . . . . . . . . . . . Kelneriapis Sakagami
Apical bend in M (at point where vein would
meet 1m-cu) absent (fig. 113); antennal
sockets less than one antennal socket diameter from basal clypeal margin (fig.
112); posterior corner of corbicula distinctly pointed (fig. 114); scutellum not projecting over metanotum; basal area of propodeum horiztonal and as long as scutellum
. . . . . . . . . . . . . . . Liotrigonopsis, n. gen.
Genus Kelneriapis Sakagami

Tetragonula (Kelneriapis) Sakagami, 1978: 232.
Type species: Hypotrigona eocenica Kelner-Pillault, 1969a, monobasic. Michener, 1990: 106
[as a genus].
Kelnermelia Moure In Moure and Camargo,
1978: 565. Type species: Hypotrigona eocenica
Kelner-Pillault, 1969a, monobasic and original
designation, isotypic with Kelneriapis Sakagami, 1978.
DIAGNOSIS: This genus differs from the

only other Baltic amber meliponine genus,
Liotrigonopsis, by the separation of the antennal sockets from the basal margin of the
clypeus more than 1 OD, the rounded apical
corner of the metatibia, the scutellum projecting over the metanotum and propodeum,
and the strongly declivitous and shortened
basal area of the propodeum. Among living
genera Kelneriapis is most similar to, and
perhaps sister to, Hypotrigona from Africa;
both genera have the posterior apical corner
of the metatibia rounded, although slightly
less so in Kelneriapis. Kelneriapis differs,
however, by the scutellum that projects over
the metanotum and propodeum, the absence
of mandibular dentition, and the shortened
and declivitous propodeum.
DESCRIPTION: Mandible without dentition.
Malar space shorter than basal mandibular
width. Antennal sockets set above base of
clypeus by more than an antennal socket di-
NO. 259
ameter. F1 approximately equal in length to

pedicel; F1 shorter than F2; F2 approximately equal in length to F3; flagellomeres with
numerous, minute sensillar plates. Inner margin of compound eyes straight, eyes apparently parallel (not converging below). Preoccipital area rounded. Anterior border of
mesoscutum broadly rounded; tegula oval;
scutellum strongly projecting over metanotum and propodeum, without medioapical Vshaped notch, apical margin rounded. Basal
area of propodeum much shorter than scutellum, strongly declivitous. Corbicula only
weakly concave along apical third of metatibia; posterior angle of corbicula angled but
not sharply (intermediate between Hypotrigona and Liotrigona); inner surface of metatibia with narrow keirotrichiate field, keirotrichiate field weakly elevated, bordered by
broad, weakly depressed zone (as in Hypotrigona); inner surface of metabasitarsus
without basal sericeous area. Forewing with
distal bend at point where nebulous 1m-cu
meets M present (fig. 111); no indication of
submarginal cells (not even by nebulous
veins); marginal cell apex open; hind wing
difficult to see, without closed cells.
COMMENTS: The authorship of the name
Kelnermelia was indicated as Moure and
Camargo by Michener (1990, 1997); however, the name is attributed solely to Padre
Moure in the original publication.
Unlike the description by Kelner-Pillault
(1970b), which suggests that the specimen
was a male (i.e., that there was no corbicula,
13 antennomeres, absence of penicillum), the
specimen on which her description was
based (labeled in her own hand as the holotype and matching the photograph she presented) is in fact a female with a corbicula,
12 antennomeres, and a penicillum.
Kelneriapis eocenica (Kelner-Pillault),
new combination
Figure 111
Plate 7e
Hypotrigona eocenica Kelner-Pillault, 1969a: 87.

Trigona (Hypotrigona) eocenica (Kelner-Pillault);
Kelner-Pillault, 1970b: 437.
Tetragonula (Kelneriapis) eocenica (Kelner-Pillault); Sakagami, 1978: 232.
Kelnermelia eocenica (Kelner-Pillault); Moure
and Camargo, 1978: 565.
2001

mm). Interocellar distance 0.22 mm; ocellocular distance 0.17 mm; median to lateral
ocellus 0.07 mm. Labrum much broader than
long, apical margin straight. Glossa relatively
short; flabellum minute; setae of labial palp
segments identical to that described for Liotrigona mahafalya Brooks and Michener
(Michener, 1990: his fig. 61). Scape relatively straight; flagellum with 10 flagellomeres
(original description by Kelner-Pillault is in
error). Forewings with identical venation
(original description by Kelner-Pillault is in
error by mentioning they are different!); basal vein basad cu-a crossvein.
Mandible smooth and impunctate, without
outer grooves. Labrum, clypeus, supraclypeal
area, face, vertex, and gena smooth and impunctate. Pronotum smooth and impunctate;
mesoscutum smooth with a few, faint, small
punctures separated by five times a puncture
width or more; tegula and scutellum smooth
and impunctate; metanotum apparently finely
imbricate; pleura smooth and impunctate.
Corbicula and outer surface of metabasitarsus finely imbricate. Basal area of propodeum glabrous (lateral and posterior surfaces
not visible). Metasoma finely and faintly imbricate.
Head and mesosoma (where preserved)
dark brown to black, without maculations.
Legs dark brown. Wing membrane hyaline;
veins light brown. Metasoma apparently dark
brown.
Pubescence generally white. Mandible
with widely scattered, minute, subappressed,
simple setae. Labrum with minute, simple,
subappressed setae evenly scattered over surface. Clypeus, supraclypeal area, and face to
slightly above level of antennal sockets as
described for labrum. Face above level of antennal sockets with setae as on clypeus but
noticeably more sparse, erect, and intermixed
with simple, erect, slightly longer setae; such
erect setae also on vertex but disappearing
on gena where pubescence is as described for
clypeus. Mesoscutum with widely scattered,
short, simple setae, setae becoming more
dense toward anterolateral corners; tegula
with scattered, minute, erect setae restricted
135
to inner margin; scutellum as described for

mesoscutum except setae slightly longer, setae noticeably more dense along posterior
margin and slightly longer; metanotum and
basal area of propodeum without pubescence. Corbicula bordered by long, simple
setae; apex with penicillum; outer surface of
metabasitarsus without pubescence, laterally
with short, stiff, simple setae, inner surface
with stiff comb rows. Metasoma with sparse,
minute, simple, subappressed setae.
MATERIAL: One specimen. Holotype. Female, worker caste, MB.I.1946 (ZMHB) labeled: 15 [old Berendt collection number]
// Holotyp [sic], Palaontologisches Museum
Berlin, Trigona (Hypotrigona) eocenica Kelner-Pillault, 1970, Baltischer Bernstein //
Holotype, Trigona eocenica Kelner-Pillault // Hypotrigona eocenica S.K.P., Holotype, S. Kelner-Pillault det.[in Kelner-Pillaults handwriting].
C OMMENTS : Although Kelner-Pillaults
1970 paper (1970b) is commonly cited as the
original proposal of the species, the name
was apparently made available by Kelner-Pillault a year earlier (1969a), where a one-sentence diagnosis is presented to separate the
species from extant Hypotrigona; the name
is given as Hypotrigona eocenica n. sp.,
and two figures are provided to distinguish
the fossil from living meliponines. Although
meeting abstracts are excluded from availability (ICZN, 1999b: Art. 9.9), published
proceedings [as is the case for Kelner-Pillault
(1969a)] are available. I therefore consider
the name as having been made available in
1969 rather than 1970, and that the original
combination was Hypotrigona eocenica rather than Trigona (Hypotrigona) eocenica.
Although Michener (1990) was the first to
use Kelneriapis at the generic level (Sakagami treated this group as a subgenus of Tetragonula), he did not make the associated
taxonomic combination for the type species
and I have therefore considered my usage
above to be a new combination.
Liotrigonopsis, new genus
TYPE SPECIES: Liotrigonopsis rozeni Engel,
new species.
DIAGNOSIS: In keys to the genera of Meliponini this group runs to the African and
136
Malagasy genus Liotrigona Moure (1961).

This fossil differs, however, in the following
generic characteristics: antennae inserted low
on face (separated from clypeus by less than
1 OD) and not far above lower tangent of
compound eyes as well as vein M without
apical bend among other, more trivial, features. Liotrigonopsis can be separated from
Kelneriapis, also in Baltic amber, by the
characters presented above in the Diagnosis
for Kelneriapis.
DESCRIPTION: Malar space shorter than basal mandibular width (fig. 112). Antennal
sockets set near base of clypeus, separated
by less than an antennal socket diameter (fig.
112). F1 approximately equal in length to F2;
F2 approximately equal in length to F3; flagellomeres with numerous, minute sensillar
plates. Preoccipital area rounded. Anterior
border of mesoscutum broadly rounded; tegula oval; scutellum clearly not projecting
over metanotum, without medioapical Vshaped notch, apical margin rounded. Basal
area of propodeum as long as scutellum, horizontal. Posterior angle of corbicula sharply
angled (as in Liotrigona) (fig. 114); inner
surface of metatibia with narrow keirotrichiate field, keirotrichiate field not elevated,
bordered by broad, glabrous zone; inner surface of metabasitarsus without basal sericeous area. Forewing without distal bend at
point where 1m-cu would meet M (fig. 113);
without indication of submarginal cells (fig.
113); marginal cell apex open; hind wing difficult to see, apparently without closed cells.
is a combination of Liotrigona and the suffix
-opsis (Greek, meaning appearing like).
The name is a reference to the similarity between the fossil and Liotrigona, a genus of
African and Malagasy stingless bees. The
name is feminine.
Liotrigonopsis rozeni, new species
Figures 112115
Plate 7f

length 0.87 mm (width indeterminate from
specimen). Basal vein distad cu-a by three
times vein width; minute basal stub of first
NO. 259
Fig. 112. Right lateral view of head of holotype female of Liotrigonopsis rozeni, new species.
Scale bar 5 0.5 mm.
abcissa of Rs after separation from M present, M terminating shortly thereafter without apical bend.
Integument of head, mesosoma, and legs
smooth and impunctate. Terga and sterna
finely imbricate and impunctate.
Head and mesosoma dark brown except
scape light brown (remainder of antenna dark
brown). Legs and metasoma light brown or
ferruginous. Wing membrane hyaline; veins,
when present, light brown.
Pubesence golden, sparse, and simple. Setae not apparent on head. Mesosoma with
sparse, simple, short, erect setae. Setae of
legs simple and scattered; inner surface of
metatibia densely covered by keirotrichiae
except narrow (ca. 0.75 OD) glabrous posterior border; rastellar bristles of moderate
length, tapering at apices, not flattened. Terga
with exceedingly sparse, short, suberect setae; setae of sterna sparse, restricted to apical
thirds, and moderately long.
Yantarny // Holotype, Liotrigonopsis rozeni Engel. The specimen is partially preserved along the edge of the amber piece
with the left half of the head and left anterior
portion of the mesosoma missing. Enough of
2001
137
Figs. 113115. Mesosomal structures of Liotrigonopsis rozeni, new species. 113. Forewing. 114.
Outer surface of metatibia and metatarsus. 115. Metadistitarsus, claw, and arolium. Scale bars 5 0.5
mm (a: for fig. 113) (b: for fig. 114); 0.25 mm (b: for fig. 115).
the specimen is preserved to allow for not

only identification but for meaningful comparison to living species.
ETYMOLOGY: The specific epithet is a patronymic honoring Dr. Jerome G. Rozen, Jr.
for his friendship and support of my studies
on bees.
APOIDEA INCERTAE SEDIS
Genus and Species Indeterminate
The following specimens are too poorly
preserved to allow identification below the
level of family. I have listed them here simply as a record of the families and for the
total number of individual bees recognized
in Baltic amber.
APIDAE gen. et sp. indet.: One female,
MB.I.1938 (ZMHB) labeled: Museum fur
Naturkunde Berlin, Palaontologisches Museum, Inv. Nr. MB.I.1938 (No. 53) // 53
// Apidae indet., det. M. S. Engel. One
female, Nr. 2616 (CCGG) labeled: Nr.
2616 // Apidae indet., det. M. S. Engel.
MEGACHILIDAE gen. et sp. indet.: One
138
female, Nr. 8 (CSUL) labeled: Megachilidae indet., det. M. S. Engel.

Missing Taxa of Uncertain Identity
The following named taxa are known only
on the basis of their fragmentary original descriptions. No specimens are known to be extant for any of these species. Interestingly,
most were considered to be bumble bees, or
bumble bee ancestors, by their authors. It is
likely that none were actually bumble bees
or even closely related to the Bombini, but
this may never be known with certainty.
Three of these names (Bombus carbonarius,
B. pusillus, Bombusoides mengei) were erroneously listed as nomina nuda by Keilbach
(1982).
Andrena wrisleyi Salt
Andrena wrisleyi Salt, 1931: 141.
COMMENTS: The description and illustrations of this specimen are adequate to determine that the species was not an Andrena or
even an andrenid. It is possible that the species was a melittid of some sort that has not
subsequently been discovered among newer
material. Although the melittid hypothesis is
perhaps the best possibility, the original description and particularly Salts figure of the
wing venation are enigmatically suggestive
of an electrapine. Although the figure of the
hind tibia does not show a corbicula, the description provided of the hind leg by Salt
does not match his figure and is somewhat
suggestive: . . . tibiae triangular, much widened apically, . . . (Salt, 1931: 142). The
wing venation is quite similar to species of
both Electrapis and Protobombus (if an electrapine, Andrena wrisleyi would fall close to,
or in, Electrapis owing to the elongate metabasitarsus). It seems hard to imagine but it
is possible that Salts specimen was actually
an unknown corbiculate apine in the Electrapini! The holotype was deposited in the illfated University of Konigsberg and was not
subsequently located in any other institution.
None of the bees before me can be readily
associated with Salts Andrena and so I
have not designated a neotype. The name
must be relegated to incertae sedis.
NO. 259
Bombus carbonarius Menge

Bombus carbonarius Menge, 1856: 27.
COMMENTS: The size and general habitus

of B. carbonarius from Menges description
suggest a species of Electrapis; however, this
is entirely conjectural. There is not enough
information to confidently assign this species
to any of the several Bombus-like genera.
Bombus muscorum Roussy
Bombus muscorum Roussy, 1937: 58. Nomen
praeoccupatum (nec Linnaeus, 1758).
COMMENTS: The name for this species

must eventually be changed as it is a junior
secondary homonym of Linnaeuss Apis
muscorum, which has been considered a valid species of Bombus for well over a century.
The type was deposited in Roussys private
collection and is now presumably lost. As
mentioned above, this species is likely not a
true bombine at all; Roussy (1937) himself
considered this specimen to be representative
of a solitary bee, while all bombines are
known to be social, a fact universally known
in Roussys time.
Bombus pusillus Menge
Bombus pusillus Menge, 1856: 27.
COMMENTS: A supposed bumble bee that

was described as . . . ist kaum 3 mill. lang
und 1 mill. breit. Behaarung gelblich weisz,
auf dem brustrucken zottig. (Transl. . . .
is scarcely three millimeters long and 1 millimeter wide. Pilosity yellowish-white, of the
thoracic dorsum shaggy; translation of the
author). It is hard to imagine what this species might have been (perhaps a spheciform
or other aculeate?). If indeed a bee, then it
was certainly one of the smallest in Baltic
amber.
Genus Bombusoides Motschulsky
Bombusoides Motschulsky, 1856: 28. Type species: Bombusoides mengei Motschulsky, 1856,
monobasic.
COMMENTS: Refer to Comments provided

for the type species (below).
2001
Bombusoides mengei Motschulsky

Bombusoides mengei Motschulsky, 1856: 28.
COMMENTS: Motschulskys treatment of

this specimen is exceedingly short and provides no clue as to what the bee may have
actually been. The entire reference to this bee
consists of the clause . . . entre autres une
tre`s belle Andraena [sic] et un Bombus nain
da peine 1 ligne de longueur, que je me
permets de dedier a` M. Menge, en le nommant Bombusoides Mengei. (Transl. . . .
among others a very beautiful Andrena and
a dwarf Bombus scarcely 1.5 lines in length,
which I allow myself to dedicate to Monsieur
Menge, with the name Bombusoides mengei; translation courtesy M. G. Rightmyer).
There is no way to associate this name with
any of the smaller bees presently known
from Baltic amber and thus the name must
be relegated to incertae sedis.
CLADISTIC ANALYSES
Cladistic analyses were undertaken in order to both acquire an understanding of the
phylogenetic position of particular fossil taxa
and to explore the potential effects on hypotheses of relationships previously based
solely on Recent species. Data matrices were
constructed in WINCLADA (Nixon, 1999)
and submitted from there for analysis by
NONA (Goloboff, 1993). An initial search
was made on each matrix using the wh*
command to identify a single tree or set of
trees upon which more exhaustive branch
swapping was undertaken using the max*
command. Topologies were visualized and
printed using WINCLADA (Nixon, op. cit.).
LITHURGINAE
No previous attempt has been made to cladistically reconstruct lithurgine relationships.
Three recent genera have generally been recognized: Trichothurgus, Microthurge, and
Lithurgus; the last has two subgenera, Lithurgopsis and Lithurgus proper (Michener,
1983). Michener (1983) discussed characters
supporting the monophyly of the subfamily
and presented an argument for the relationships among these three genera as Trichothurgus (Microthurge 1 Lithurgus). Herein I
have examined representatives of each ge-
139
TABLE 10
Characters Used in Cladistic Analysis of

Lithurginae
nus-group taxon inclusive of both subgenera

of Lithurgus and Lithurgomma, a junior synonym of Trichothurgus (sensu Michener,
1983, 2000a). Like Michener (1983), I could
find no qualitative character to differentiate
Lithurgomma from Trichothurgus and therefore, when coding the matrix, chose to retain
these taxa as synonymous. The subgenera of
Lithurgus, however, could be qualitatively
diagnosed, as could the enigmatic Australian
species Lithurgus rubricatus Smith (1853),
and each was coded individually in the matrix. Ten characters of adult external morphology were identified and coded for the recent lithurgines, the fossil genus Protolithurgus, and the basal megachilid subfamily Fideliinae (used as an outgroup). The resulting
data matrix is presented in table 11, while
character definitions are provided in table 10.
Analysis of the data matrix resulted in a single topology of length 11, CI 0.88, and RI
0.90 (fig. 116).
The relationships proposed by Michener
(1983) are generally in accord with those
presented here (fig. 116). Protolithurgus is
placed as sister to all other lithurgines owing
to the plesiomorphic retention of a flattened
metabasitarsus (typical of other bees), two
mandibular teeth, and absence of spicules on
the tibiae. Another feature apparently uniting
the living genera is the elongation of the proboscis; however, this quantitative character
showed little discrete differentiation from
140
TABLE 11
Data Matrix for Analysis of Generic

Relationships of Lithurginae
(Character descriptions presented in table 10)
some fideliines and alternative states could

not be consistently and unambiguously assigned. For this reason the elongation of the
proboscis was not included in the data analysis but the general congruence of this feature with the resulting topology is interesting
and noteworthy. Although Protolithurgus
seems to possess an enigmatic combination
of characters unique among megachilids, the
genus does share with other Lithurginae the
distinctive flattened, first metasomal tergum
with a rounded apical margin, a feature
found only in lithurgines and which unambiguously unites the fossil with this clade.
NO. 259
The genus Trichothurgus, as in Micheners

system, is sister to all other extant lithurgine
genera owing to the lowest mandibular tooth
being longer than those above it and the nonprojecting pygidial plate in females. Unexpectedly, the genus Microthurge, although itself likely monophyletic, was sister to Lithurgus exclusive of Lithurgopsis and L. rubricatus. The monophyly of Lithurgus should
be carefully scrutinized as this preliminary
analysis could not support the group as it has
been traditionally defined. The subgenus Lithurgopsis should perhaps be accorded generic rank and L. rubricatus transferred to its
own genus. It must be noted that the phylogeny presented here is preliminary. A more
thorough analysis of lithurgine relationships
is needed before major classificatory changes
are made to the living genera. I have placed
all extant genera in a nominate tribe (with
Trichothurgini Moure as a junior synonym),
while the fossil is sufficiently distinct to warrant the recognition of a monogeneric tribe
sister to Lithurgini (see Protolithurgini,
above).
XYLOCOPINAE
Sakagami and Michener (1987) investigated tribal relationships with the Xylocopinae,
Fig. 116. Phylogeny of Lithurginae based on cladistic anlaysis of data presented in table 11 (Length
11, CI 0.88, RI 0.90). Black dots are unreversed changes; white dots homoplastic character transitions.
The character number is indicated above the branch and the state change is indicated below. Fideliinae
is the outgroup.
2001
141
TABLE 12
Characters Used in Cladistic Analysis of Xylocopinae
although they assumed this group to be sister

to the corbiculate Apinae (Apidae in their
system) and used this group, particularly the
basal tribe Euglossini (see below), for a
priori character polarizations. They concluded that the monogeneric tribe Manueliini was
sister to all other xylocopines and that Xylocopa (Xylocopini) was intermediate between Manuelia and a Ceratinini 1 Allodapini clade. As part of a more exhaustive
study on long-tongued bee tribal relationships, Roig-Alsina and Michener (1993)
demonstrated that the xylocopines were more
basal in apid phylogeny and not closely allied to the corbiculates. Furthermore, they
concluded that the relative positions of XyTABLE 13
Data Matrix for Analysis of Tribal

Relationships of Xylocopinae
(Character descriptions presented in table 12;
the one multistate is nonadditive)
locopini and Manueliini from the earlier

study were in fact reversed. Thus, the topology of xylocopine relationships was Xylocopini (Manueliini (Ceratinini 1 Allodapini)).
In order to determine the phylogenetic
placement of the enigmatic Baltic amber xylocopines (i.e., Boreallodape) a character
matrix was constructed to explore relationships among the tribes of the subfamily. Fifteen characters were identified and coded for
the sole genus of Xylocopini (Xylocopa sensu Minckley, 1998), the sole genus of Manueliini (Manuelia), one of the two genera of
Ceratinini (Ceratina), and two genera of Allodapini (Macrogalea and Allodape). The
two genera of allodapines were selected to
represent phylogenetic diversity within the
tribe based on the cladistic studies of Michener (1977) and Reyes (1998). The genus Exomalopsis was used as an outgroup. Characters identified for analysis are presented in
table 12, while the complete matrix is presented in table 13. Analysis of the data produces a single topology of length 16, CI
1.00, and RI 1.00. The topology is depicted
in figure 117.
The results of this analysis are in complete
accord with the analysis of Roig-Alsina and
Michener (1993) in that Xylocopini is basal
within the subfamily owing to the plesiomorphic retention of alar papillae and an elon-
142
NO. 259
Fig. 117. Phylogeny of Xylocopinae based on cladistic analysis of data presented in table 13 (Length
16, CI 1.00, RI 1.00). Black dots are unreversed changes; white dots homoplastic character transitions.
The character number is indicated above the branch and the state change is indicated below. Exomalopsis
is the outgroup.
gate first flagellomere. The Ceratinini, Allodapini, and Boreallodape are grouped together in a well-supported clade (fig. 117). This
group is most notable for the abruptly narrowed mandibular structure (e.g., fig. 54).
The analysis demonstrates that Boreallodape
is most closely related to the Allodapini, both
groups sharing the presence of two submarginal cells and the absence of the metabasitibial plate. As discussed above (under the
description of the tribe Boreallodapini) differences between the Allodapini and Boreallodape are significant and inclusion of the
latter as the basalmost member of the former
would mask the enigmatic features of the
fossil among xylocopines. Boreallodape is as
distinctive as any of the other tribes of the
subfamily and recognition of a separate tribe
for this group is well supported. It is biogeographically interesting to note that Boreallodape is sister to the Allodapini, a group
today primarily distributed in sub-Saharan
Africa, Asia, and Australia. Among the allodapines only the genus Exoneuridia occurs
in the West Palearctic and extends as far
north as the Mediterranean region in the
southeastern half of Turkey (Terzo, 1999).
However, a historical biogeographic analysis
of the allodapines derived from the recent
cladogram of Reyes (1998) suggests that the
extant groups of the tribe as a whole are per-
haps of a sub-Saharan African origin and that

Exoneuridia represents a subsequent dispersal into Asia Minor. Most likely, the protoallodapines (perhaps together with proto-boreallodapines) were of an original Gondwanan origin (as is likely the same for several
bee tribes and subfamilies) and the group as
we recognize it today diversified in sub-Saharan Africa, subsequently radiating further
as they dispersed in the Old World. Under
either scenario, Boreallodape serves as another example of Baltic amber fossils related
to groups now found in Asia or sub-Saharan
Africa (see below under Discussion).
CORBICULATE APINAE
Relationships among the corbiculate bees
have been controversial as have their implications for understanding the evolution of
eusocial behavior in this group of bees. As
was briefly discussed above, in the modern
fauna there are four well-defined tribes: Apini (honey bees), Bombini (bumble bees),
Euglossini (orchid bees), and Meliponini
(stingless bees). Of these four, the Bombini,
Apini, and Meliponini are all eusocial, although advanced eusociality (i.e., those societies with morphologically differentiated
queen and worker castes) is found only in
Apini and Meliponini (not just among cor-
2001
biculates, but among all of the Apoidea). Of

the 15 possible rooted topologies for four
taxa, eight have been proposed at one time
or another for the living corbiculates based
on either a priori interpretations of the same
1020 morphological characters, single character analyses, or studies of limited DNA sequences. Outgroup-based cladistic analysis
of morphological evidence produced a single, fully resolved topology (Prentice, 1991;
Roig-Alsina and Michener, 1993; Chavarra
and Carpenter, 1994; Schultz et al., 1999).
This topology supports the single origins
hypothesis whereby both general eusocial behavior and advanced eusocial behavior arose
once in a hierarchical fashion. Thus, eusociality arose in the common ancestor of the
Bombini, Meliponini, and Apini, while advanced eusociality arose in the common ancestor of Meliponini and Apini. Molecular
analyses have produced alternative cladograms that are not consistent with this scenario, nor are they always consistent with
themselves (e.g., Cameron, 1991, 1993).
These analyses imply either dual origins of
advanced eusocial behavior or ambiguous reconstructions by uniting the bumble bees and
stingless bees. A simultaneous analysis of
these DNA sequences with morphological
data produces topologies that support the single origin for advanced eusocial behavior by
reuniting the Meliponini and Apini (Chavarra and Carpenter, 1994; Schultz et al., 1999).
However, while these combined studies typically also support a single origin for general
eusociality, the support is not as strong.
Thus, a clear resolution has yet to be
achieved concerning relationships among
these tribes and the addition of more characters and more taxa to analyses is needed.
Since most characters used in morphological
studies of the tribes are fixed across taxa
within each tribe, added taxa of living representatives provide little information for resolution of relationships outside of more accurately reconstructing the basal node of
each respective tribe. Most of the Baltic amber corbiculates as demonstrated above,
however, do not fall into the living tribes and
possess combinations of characters unknown
in any of the living lineages. Thus, the addition of these groups to analyses may have
important implications for understanding
143
corbiculate relationships. Herein I have both

expanded the available morphological data as
well as the taxon representation (tables 14,
15). The character-state matrix is presented
in table 15. Analysis of these data produces
two most parsimonious topologies (length
65, CI 0.81, RI 0.93), the strict consensus of
which is depicted in figure 118. Of the two
trees, only one of the resultant topologies is
strictly supported (Nixon and Carpenter,
1996) and is identical to the consensus (fig.
118). The analysis produces a nested set of
relationships, with numerous groups breaking up some of the long branches between
the living tribes. More importantly, these results produce a hierarchy of intermediary lineages separating the advanced eusocial tribes
(i.e., Apini and Meliponini) from the bumble
bees and orchid bees. Topologies attempting
to unite Bombini with Meliponini cannot remotely account for the character combinations exhibited by the fossil taxa. That this
hierarchy is congruent with behavioral characters and other biological traits (e.g., Noll,
1998) is noteworthy. Thus, quite contrary to
the view asserted by Patterson (1981), paleontological data has had a significant impact
on our understanding of corbiculate bee relationships. Interestingly, the corbiculate
bees are a nice example of the type of group
wherein fossils are hypothesized to be most
important for phylogeny reconstruction.
Gauthier et al. (1989: 193) hypothesized,
. . . fossils should be most important in
phylogenetic inference when the group of interest is old and only a few, highly modified,
terminal taxa are extant. The extant tribes
of corbiculate bees are highly derived with
numerous synapomorphies supporting each,
and the tribes are undoubtedly ancientextending into the Late Cretaceous, as shown
by the presence of a true meliponine in amber of probable Maastrichtian age (Michener
and Grimaldi, 1998a; Engel, 2000b; see also
below under Origin of Bees). Under such
conditions intermediary fossil lineages are
critical for evaluating relationships among
the surviving lineages.
The reconstruction of social behavior on
the cladogram may, at first glance, appear to
be problematic owing to the absence of direct
observations for the social state of the fossil
taxa. It is possible, however, to infer behav-
144
TABLE 14
Characters Used in Cladistic Analysis of Corbiculate Apinae

(Characters 35 and 48 additive; all others nonadditive)
NO. 259
2001
145
TABLE 15
Data Matrix for Analysis of Tribal Relationships Within the Corbiculate Apinae
(Character descriptions presented in table 14)
ioral character information from fossils either

through 1. particular morphologies that are
tightly correlated or unique to a given ethological trait (e.g., Michener and Grimaldi,
1988b; Poinar, 1991; Engel, 1998c, 2000b;
Simmons and Geisler, 1998); 2. concrete
behaviors (e.g., nest architecture, footprints), whereby the products of a behavioral
repertoire are preserved as trace fossils either
in association with the remains of the organism or in isolation (e.g., Carpenter, 1992;
Genise and Hazeldine, 1998; Lockley and
Matsukawa, 1999); 3. preservation of the individual while in the act of performing some
portion of a particular behavioral repertoire
(e.g., Norell et al., 1995; Clark et al., 1999);
or, 4. phylogenetic inference (e.g., Michener
and Grimaldi, 1988b; Engel, 1998c, 2000b).
As mentioned above, advanced eusocial societies are recognized by morphologically
differentiated queen and worker castes, and
this attribute is known only for these societies. Among the fossils, all except the single
individual of Electrobombus exhibit the re-
duced metasomal structure (and to a lesser

degree a barbed sting) typical of workers; indicating that they were members of advanced
eusocial societies (fig. 119). Thus, although
the fossils were coded and analyzed as unknown for their social state, all except Electrobombus could essentially have been coded
as advanced eusocial. The fact that those
taxa with worker-caste morphologies cladistically group with the Apini and Meliponini
(fig. 119) reinforces the reconstruction of a
single origin for advanced eusocial behavior
in the bees. Moreover, the position of Electrobombus within a clade of already eusocial
(either primitive or advanced eusocial) lineages implies, via phylogenetic inference, eusocial behavior to some degree in this taxon
(fig. 119). In summary, taking all of this information into account produces a single
possible reconstruction of eusocial evolution
within the corbiculate bees. The available evidence supports only the single origins
scenario wherein general eusociality arose
once in the common ancestor of the Bom-
Fig. 118. Phylogeny of corbiculate Apinae based on cladistic analysis of data presented in table 15 (Length 65, CI 0.81, RI 0.93). Black dots
are unreversed changes; white dots homoplastic character transitions. The character number is indicated above the branch and the state change is
indicated below. Anthophora, Centris, and Xylocopa are the outgroups.
146
NO. 259
2001
147
Fig. 119. Distribution of social ethotypes among the corbiculate apines as well as fossils exhibiting
worker morphologies. The euglossine genera (Euglossa, Eufriesea, and Eulaema) are generally communal or solitary, while the outgroups (not depicted here) are solitary. Extinct genera are italicized.
bini, Electrobombini, Electrapini, Melikertini, Apini, and Meliponini with a subsequent,

single evolutionary elaboration of this trait to
advanced eusociality in the common ancestor
of at least the Electrapini, Melikertini, Apini,
and Meliponini (fig. 119) (Engel, in press-b).
Recently, new studies of DNA sequences
have attempted to provide further support for
a Meliponini 1 Bombini clade (Koulianos et
al., 1999; Mardulyn and Cameron, 1999). Interestingly, a simultaneous analysis of morphological characters and of the sequence
data published by Koulianos et al. (1999)
produces a single origins topology. These
sequence data are particularly problematic in
that they employ mitochondrial DNA regions
(like the earlier studies of Cameron, 1991,
1993) commonly used in species-level anal-
148
yses owing to high levels of interspecific variation. For example, analysis of the earlier
mitochondrial DNA data for relationships
among species within a given genus were
more internally congruent than when examining older relationships (Engel and Schultz,
1997; Schultz et al., 1999). That the corbiculate tribes are undoubtedly ancient only
serves to reinforce the need for careful selection of gene regions when undertaking an
analysis, particularly in a group that extends
back into the Mesozoic [the divergence
among the tribes being at least 75 Ma owing
to the presence of a true meliponine in Late
Cretaceous amber (Engel, 2000b)]. The data
of Mardulyn and Cameron (1999) are an improvement over those of previous studies by
attempting to select relatively conserved
gene regions, but Ascher and Danforth (in
rev.) have demonstrated that there are numerous other critical problems with these sequence data and analyses. Moreover, all of
these studies have suffered from poor taxon
sampling and choice of outgroups. Thus, the
available sequence data are not at all reliable
and a well approached (e.g., choice of ingroup taxa, choice of outgroup taxa, number
of taxa examined, number of informative
characters, choice of gene region) molecular
analysis remains to be undertaken for the
corbiculate Apinae.
Perhaps one of the more interesting morphological implications of the fossil corbiculate taxa and the cladistic analysis presented here concerns the evolution of the metatibial spurs. The apomorphic absence of
metatibial spurs in the Apini and Meliponini
has often been used as one of several traits
uniting these two tribes in analyses of the
living taxa; the presence of two metatibial
spurs is plesiomorphically found in the outgroups, euglossines, and bombines. Those
Baltic amber corbiculate genera that do not
cladistically fall into any of the extant tribes
have a single, often reduced, metatibial spur
(difficult to see in many specimens). The
taxa exhibiting this one spur characterstate form a paraphyletic assemblage derived
from a two-spurred ancestor and subsequently giving rise to the Apini and Meliponini,
neither of which have metatibial spurs. This
pattern itself is intuitively pleasing in that it
implies a gradual reduction in the number of
NO. 259
metatibial spurs. Surprisingly, however, the

Apini and Meliponini are separated by a
group of extinct taxa that are sister to the
Meliponini (i.e., the Melikertini) and that
have a single metatibial spur. Thus, the implication is that the loss of spurs in the honey
bees and stingless bees is not a feature shared
through common ancestry (i.e., not a synapomorphy for the two tribes); nonetheless,
support for their close relationship was not
degraded by the loss of this supposed synapomorphy. The apines and meliponines independently lost tibial spurs from a singlespurred ancestor. From the viewpoint of reciprocal illumination this interpretation of
independent loss makes sense. A reexamination of the metatibia-metabasitarsus junction in living stingless bees and honey bees
reveals two, somewhat different morphologies, suggesting at an observational level the
possibility that the homology of absence is
incorrect in this case. The inner apex of the
metatibia and the junction with the metabasitarsus in honey bees is essentially unmodified; the only significant difference by comparison to spurred corbiculate bees is the
simple absence of the metatibial spurs. On
the other hand, in stingless bees the entire
junction between these two leg segments is
grossly transmogrified, perhaps partly
through the loss of the auricle. In most stingless bee taxa the inner apical region of the
metatibia where the metatibial spurs would
articulate is modified with a slight impression leading down to the articulation with the
metabasitarsus, which itself is highly modified by the complete absence of an auricle.
Likewise, the inner apical margin of the metatibia extending posteriorly from this impression where the metatibial spurs would
otherwise articulate is weakened; in some instances this is further manifested through a
weakening of the rastellum. These two different morphologies of absence suggest
from examination of living taxa alone that
the conditions seen in honey bees and stingless bees are perhaps not homologous and, in
fact, represent two separate character states.
A paleontological perspective has thus provided illumination toward understanding not
only cladistic relationships among the living
taxa but also for interpreting their morphol-
2001
149
Fig. 120. Hypothesis of relationship for the corbiculate bees as envisioned by Cockerell (reproduced
from Cockerell, 1908b). He did not include tribe Euglossini and the ages he used for particular deposits
are now understood to be as follows: Calyptapis [considered as a junior subjective synonym of Bombus
by Zeuner and Manning (1976)] is from early Oligocene deposits of Florissant, while the genera Sophrobombus, Chalcobombus, Protobombus, and Electrapis, all in Baltic amber, are middle Eocene in
age. Most species of Synapis (a subgenus of Apis) are from Oligocene strata but at least two species
are from early Miocene deposits (Engel, 1999c, unpubl. data). Meliponorytes (described by Tosi, 1896)
is either a true fossil in Miocene amber from Sicily or perhaps misidentified in African copal and
congeneric with living meliponines (see appendix 1).
ogy via a character-state known only from

the fossil record.
The analysis presented here is the first cladistic analysis combining fossil corbiculate
genera with living genera. It is, however, not
the first phylogenetic attempt to unite the fossil record of corbiculate apines with their extinct counterparts. Cockerell (1908b) pre-
sented a phylogeny for this group of bees

(excluding the Euglossini) derived from a
few characters and a great deal of intuition.
His interpretation (fig. 120) is not too different from the one presented here; i.e., Cockerell united the Apini and Meliponini as sister taxa. My interpretation of the fossils differs from that of Cockerell in that he also
150
used plesiomorphic traits to unite genera; for

instance, he brought together those Bombuslike groups (e.g., Electrapis, Protobombus)
on the primitive absence of elongate setae on
the compound eyes (fig. 120). A cladistic interpretation of these same taxa based on outgroup comparison (both with the Euglossini
as well as non-corbiculate apine tribes) demonstrates that although fossils such as Electrapis and Protobombus are plesiomorphically similar to bumble bees, these genera
share apomorphic traits that unite them more
closely to honey bees and stingless bees (fig.
120). Thus, the Bombus-like genera form a
paraphyletic grade leading to the Apini, Meliponini, and Melikertini (species of the latter
were not known to Cockerell, as he had not
seen the only described melikertine in his
era; i.e., M. proavus). Cockerell also considered some of the fossils to be directly ancestral to modern lineages (e.g., the single species of Synapis was the ancestor of living
Apis species). While many of these taxa are
indeed plesiomorphic in most respects by
comparison to living relatives, none can be
interpreted as ancestors of extant lineages.
Paraphyletic stem groups that are probably
ancestral to large extant families are known
in the amber fossil record, a good example
being Prioriphora (Phoridae), known only in
Cretaceous amber (Grimaldi and Cumming,
1999).
DISCUSSION
ORIGIN
OF
BEES
The bees are a derived, monophyletic

group of the spheciform wasps (Muller,
1872; Michener, 1944, 2000a; Brothers,
1975, 1999; Lomholdt, 1982; Alexander,
1992; Brothers and Carpenter, 1993; Alexander and Michener, 1995; Melo, 1999), perhaps most closely related to the family Crabronidae (Alexander, 1992; Melo, 1999). Although alternative arguments have been presented that suggest bees to be related to
mutillids and formicids (e.g., Borner, 1919;
Lanham, 1979, 1980), these hypotheses were
based primarily on two characters taken in
isolation and have not been supported by any
rigorous analysis of aculeate relationships.
Reports of bees from pre-Cretaceous deposits and arising prior to the origin of an-
NO. 259
giosperms have generated a great deal of

controversy (and rightly so) but are, unfortunately, entirely unfounded. The accounts
by Hong (1984) and Hong and Miao (1992)
of a bee from the earliest Cretaceous of China can be immediately dismissed since this
fossil is easily identifiable as a sphecid wasp
(Darling and Sharkey, 1990; Michener, 1997;
Engel, 1998c, 1999c, 2000b) and in fact has
been recently synonymized with the genus
Archisphex (Rasnitsyn et al., 1998). Similarly, the fossil identified as a native bee by
Carroll (1962) from the Jurassic of Australia
is not actually a bee. Carrolls specimen is a
very poorly preserved compression fossil
lacking wings and a complete head (see her
fig. 1) and cannot therefore be assigned to
any insect group with great certainty. It must
be noted that no single character is preserved
that could place this fossil among the Apoidea (nor even among the Aculeata!).
The reports by Hasiotis and Demko
(1996), Hasiotis (1997), Hasiotis et al. (1995,
1998), and Kay et al. (1999) are not based
on body fossils but instead on trace fossils
from the Jurassic and Triassic of North
America. These accounts are fraught with errors, with statements like, . . . Jurassic
bees, like their modern homologs, may have
also used carrion for food and nutrients. . .
(Hasiotis and Demko, 1996: 366) and Triassic hymenopterans may also have scavenged carrion. . . as do some modern species
of halictid, anthophorid, trigoniid [sic], and
meliponine bees7 (Hasiotis et al., 1998:
116) Per common knowledge, bees are exceptional for their use of pollen; they collect
and provision their young with a mixture of
pollen, nectar, and/or plant oils. Even cleptoparasitic bees feed on pollen, although the
mother does not collect the pollen herself. Of
the nearly 20,000 described species of living
bees, only three species of the highly derived
7 A small taxonomic note: Enigmatically, Hasiotis et
al. (1998: 116) have removed the necrophagous Trigona
species from the tribe Meliponini and accorded this meliponine genus familial rank as trigoniids. Trigona is
a genus of Meliponini; there is no such thing as a trigoniid. An additional taxonomic problem arises in that
Hasiotis and Demko (1996: 366) refer to their nest as
the product of . . . sweat bees in the Anthophoridae and
Halictidae. Sweat bees are species of the subfamily
Halictinae; other halictids (i.e., rophitines, nomiines) and
anthophorines are not sweat bees.
2001
tribe Meliponini are obligate necrophages

(Camargo and Roubik, 1991). Despite the
claim by Hasiotis et al. (1998), no halictid or
anthophorine provisions its nests with carrion. The three Trigona necrophages certainly
do not represent the groundplan feeding behavior for bees, as they are from a derived
genus, of a derived tribe, of the most derived
subfamily, of the most derived family of
bees. Even if one wanted to argue that the
fossil nests represented an extinct species of
this derived clade, the position would still be
unfounded, as all three construct resinous
nests and combs in tree or ground hollows
and not branching burrows/galleries in the
soil or in wood (Roubik, 1983; Camargo and
Roubik, 1991). The large number of cells in
the fossil nest described by Hasiotis and
Demko (1996) was taken as evidence for eusocial behavior. As has been repeatedly demonstrated for ground-nesting bees (e.g., Sakagami and Michener, 1962; Eickwort and
Sakagami, 1979; Engel, 1998b), there is no
correlation between nest architecture and level of sociality. Solitary and communal species can make nests just as large and elaborate as semisocial and eusocial species. Hasiotis and Demkos fossil nest shows no single
character indicative of bee activity. The presence of flask-shaped cells, the slightly constricted opening, and shafts/corridors are
generalized features common to many
ground-nesting Hymenoptera, as well as
some insects in other orders, so such characters are not diagnostic for bees as erroneously suggested by Hasiotis and Demko
(1996). The presence of a spiral cell closure
is one of the few features suggestive of bee
activity (although it also occurs in some other insects) and such a structure was not found
with their trace fossils. Otherwise, bee nests
in the ground are similar in general plan to
many groups of Hymenoptera. Researchers
should be far more hesitant to attribute such
trace fossils to bees. These same difficulties
apply equally to the Triassic trace fossil discovered in a petrified tree in Arizona and
popularized by Hasiotis (1997; see also Wilford, 1995), Hasiotis et al. (1995, 1998), and
Kay et al. (1999). None of these nests are the
products of bees; not only do they lack any
characters suggestive of bees, they actually
show evidence of having been beetle galler-
151
ies (see also critique by Grimaldi, 1999, who

arrived at the same conclusion). Several
modern beetle groups make galleries that are
similar to the traces they have recovered. The
chemical characteristics presented by Kay et
al. (1999) to be diagnostic for primitive bees
are in fact not diagnostic for bees at all. Phenetic comparisons in chemical composition
are meaningless for a positive identification;
what is needed would be an apomorphic
chemical trait unique (i.e., fixed and diagnostic) to bees. No such chemical or blend
has been identified to date.
Some bee species have been observed to
collect pollen from gymnospermsTrigona
carbonaria Smith on Cycas media Brown
(Cycadaceae) (Ornduff, 1991) and Colletes
sp. on Juniperus virginiana Linnaeus (Cupressaceae) (Burnham, 1978)and this
might be seen to lend credence to a pre-angiosperm origin for the bees. Such forgaing
behavior led both Burnham (1978) and Ornduff (1991) to speculate that bees might be
older than flowering plants; however, to truly
support such a conclusion one must demonstrate that the groundplan foraging behavior
for the bees was on gymnosperm pollen with
a subsequent host-shift to angiosperms.
The foraging activity of the polylectic T. carbonaria does not reflect any groundplan foraging behavior for the bees owing to its apomorphic position in the Meliponini (fig. 123).
The behavior of the unidentified Colletes
species is more tantalizing owing to the basal
position of this subfamily (albeit not necessarily this genus). However, this foraging behavior is not distributed across a series of
basal species in Colletes, the Colletinae, or
even the Colletidae (or other bee families, for
that matter, as would be required for a cladistic reconstruction of gymnosperm foraging in the bee groundplan). Colletes, in fact,
shares credible larval synapomorphies with
the Hylaeinae, Xeromelissinae, and Euryglossinae while the remainder of the Colletinae as it is currently conceived (i.e., the
Paracolletini) is more basal (McGinley,
1981). Thus, such an observation in a single
species of Colletes does not reflect a groundplan trait for the bees. Instead, the foraging
activities of a variety of genera across the
Paracolletini (as well as numerous other colletid groups) and basal genera of other basic
152
short-tongued bee families would need to be

observed before any such conclusion could
be drawn. Until an identifiable body fossil of
a bee is discovered from pre-Cretaceous sediments (or in association with nest remains),
there exists no evidence that bees arose prior
to the origin and early diversification of the
angiosperms.
A particular problem with claims for a Jurassic or Triassic origin is that they hypothesize, in the absence of any bee fossil, that
bees arose prior to Aculeata (in the case of
the Jurassic) and before the Apocrita (in the
case of the Triassic)! The entire order Hymenoptera does not appear in the fossil record until the Triassic and then only on the
basis of the basalmost symphytan family
Xyelidae. No apocritan lineage has been
found prior to the Jurassic. Bees simply
could not have been present in the Triassic
since no apocritans or even advanced wood
wasps had yet originated. Likewise, in Jurassic deposits the known hymenopterans are
basal apocritan lineages, a diversity of symphytans, and a single plesiomorphic family
of aculeates. The earliest aculeate fossils are
of the extinct family Bethylonymidae from
the Uppermost Jurassic of Kazakhstan (Rasnitsyn, 1975) considered to be the sister
group to all other aculeates (Rasnitsyn, 1988;
Ronquist et al., 1999). All other aculeates in
the fossil record have their first appearance
in the Cretaceous. Moreover, the earliest
members of the Apoidea are plesiomorphic
spheciform wasps in Lower Cretaceous sediments of Brazil, Europe, and Central Asia.
No single specimen has been found of a
more advanced spheciform lineage (e.g.,
Crabronidae) prior to the Early Cretaceous.
As mentioned above, bees are a derived,
monophyletic group arising from the paraphyletic assemblage of spheciform families
in the Apoidea. Bees, therefore, must have
originated after the earliest spheciforms.
Since spheciforms (and thereby the entire superfamily Apoidea) did not appear until the
Cretaceous, bees must have originated in the
Cretaceous. If one prefers to argue for the
oldest possible age of bees, then the uppermost boundary for their age is the earliest
part of the mid-Cretaceous, or about 125 Ma.
This automatically places the origin of bees
after the origin of angiosperms, which them-
NO. 259
selves either originated in the earliest Cretaceous (Crane et al., 1995) or the uppermost
Jurassic (Sun et al., 1998). Thus, the available evidence suggests quite the opposite
scenario from those of the aforementioned
authors. Bees arose sometime after the origin
of flowering plantsa long-standing conclusion arrived at by numerous authors (e.g.,
Michener, 1979; Engel, 1996, 2000b; Grimaldi, 1999) and the only one with available
evidence.
Interestingly, the radiation of bees coincides nicely with the period in which angiosperm diversity increased dramatically. The
oldest fossil bee presently recorded is Cretotrigona in Late Cretaceous (Maastrichtian)
amber from New Jersey8 (Engel, 2000b).
Cretotrigona is a derived apine of the tribe
Meliponini (Apidae: Apinae). Thus, the cladogenetic events that produced all of the lineages between the origin of bees and the
branch that eventually gave rise to the meliponines must have taken place prior to the
Late Cretaceous. If a cladogram of bee families is overlayed on the geological column
and the oldest possible age for bees assumed
(fig. 121), then the diversification of bees
into higher lineages must have coincided
with the angiosperm radiations that took
place from 13090 Ma (Crane et al., 1995),
with a particularly dramatic increase in generic diversity 11590 Ma (Lidgard and
Crane, 1988; Crane and Lidgard, 1990).
Even if the bees are somewhat younger than
the 125 Ma date hypothesized here, they still
must have radiated prior to the Late Cretaceous [simply owing to the presence of Cretotrigona and the existence of plants today
tightly associated with Apinae in 90 Ma deposits (Crepet and Nixon, 1998)]. Thus, the
window for both the origin and early diversification of bees into higher lineages lies between ca. 125 Ma and 100 Ma. This period
of diversification for bees accords nicely
with similar periods in the evolution of derived floral characters associated with insect
pollination (Crepet, 1996). There is, at pre8 The Cretaceous age of this amber was questioned by
Rasnitsyn (In Rasnitsyn and Michener, 1991) and considered to be Paleocene. Grimaldi (1999), however, has
argued convincingly for a Cretaceous age of this deposit
and the Late Maastrichtian age (6570 Ma) is adopted
here [see also Engel (2000b)].
2001
153
Fig. 121. Phylogeny of bee families overlaid on the geological column. The shaded area indicates
the period of angiosperm diversification. Although flowering plants originated earlier than the shaded
area indicates, the geological record does not show a dramatic increase in angiosperm diversity until
when first indicated in the figure.
sent, no escape from the conclusion that bees

radiated along with angiosperms in the Cretaceous.
As just discussed, bee body fossils are
presently known only as far back as the
Maastrichtian (Engel, 2000b). Trace fossils
believed to be the product of bees are known
from a variety of localities but the most notable for the discussion here are those from
desposits ranging from the latest Cenomani-
an (Elliott and Nations, 1998) to the earliest

Paleocene (e.g., Genise and Bown, 1996;
Genise and Hazeldine, 1998; Genise, 1999;
Genise and Verde, 2000). The depth and total
number of cells in some of these nests were
taken as evidence for social behavior (e.g.,
Elliott and Nations, 1998) but as mentioned
above such attributes are not correlated with
sociality in living bees and solitary bees can
dig remarkably deep and large nests. It
154
would be impossible to state, based on nest

design, whether any of these presumed tracemakers (if indeed bees) were social. The
identity of most of these traces is suspect but
a fewsuch as the ichnogenus Uruguay
(Roselli, 1938)are clearly the product of
bee activity. Thus, trace fossils do provide
some, albeit limited, information for documenting Cretaceous bees. The Late Maastrichtian (although perhaps earliest Paleocene) ichnogenus Uruguay is remarkably
similar to cell-cluster nests of halictid bees,
and those of the tribe Augochlorini in particular [this based on a cladistic reconstruction
of nest architecture known for living Augochlorini Engel (1998b: also summarized briefly in Engel, 2000a), which reconstructed a
groundplan design for the tribe that included
cell-clusters]. Although another feature concluded to be plesiomorphic to the tribe was
semisocial behavior, this determination was
based merely on the distribution of sociobiological attributes of Recent species and not
on correlated aspects of their nest architecture (Engel, 1998b, 2000a). At that time the
augochlorines were hypothesized to be at
least 7580 Ma and to have originated from
regions today in southern South America;
thus, there is a tantalizing correlation between the reconstructed groundplan nest design for Augochlorini and the structure of
Uruguay. It is possible that the tracemakers
may not have been primitive Augochlorini
but neither should augochlorines be excluded
as a likely candidate. Genise and Verde
(2000) dismissed the augochlorine hypothesis for these cells, concluding that . . . Uruguay cannot be attributed to sweat bees because of its size, thickened walls, spiral closures and other features. In actuality, none
of these can exclude the possibility of sweat
bees for the following reasons: 1. it is true
that for typical sweat bees, such as Dialictus
(tribe Gastrohalictini), the cell sizes of Uruguay are too large; however, the cell sizes are
ample for the more robust augochlorines
(e.g., Pseudaugochlora, Megommation, some
Augochloropsis, some Rhinocorynura, &c.);
moreover, several of these species are known
to produce cell clusters [(of cell-cluster-producing Augochlorini, cell sizes can range
from 8 mm up to at least 21 mm (Sakagami
and Michener, 1962), while the range of cell
NO. 259
sizes in Uruguay auroranormae Roselli are

from 1215 mm (Genise and Bown, 1996)];
2. the thickness of cell walls is quite variable
in halictines, and the most distinct thickenening of the walls in Uruguay are at the cell
apex, a feature typical in nests where the
cells are reused; for example, in Pseudaugochlora where cells in a cell-cluster are reused, the bees fill in more soil mostly at the
apex of the old cell and to a lesser degree on
the sides before applying a new cell lining
(Michener and Lange, 1958), resulting in a
similar cell construction to that of the tracefossil; and 3. spiral closures are well known
in halictines, albeit less conspicuous in some
species (Sakagami and Michener, 1962). The
possibility remains that Uruguay is the oldest
evidence of sweat bee activity. If indeed the
products of bees (perhaps also Halictinae),
then the fossilized nests described by Elliott
and Nations (1998) would be the oldest record of bees presently known (ca. 92 Ma),
as well as superceding Uruguay as the oldest
record for Halictinae. Ichnology contributes
important information for understanding the
evolutionary history of particular taxa. The
unification of paleontology and paleoichnology will undoubtedly be increasingly important in the future for elucidating the evolution
of bees (Genise and Engel, 2000).
Although the region of bee origination and
diversification is speculative, the xeric interior of western Gondwanaland (southern
South America, Africa, and portions of Antarctica) is most likely to have been the area
from which they arose. Today, bees and
spheciform wasps, particularly the basal
groups of most families, are most abundant
and diverse in such xeric habitats (although
clear exceptions do occur in some subfamilies). This pattern of xeric-concentrated diversity is also true for the Masarinae and major pollinating flies (Gess, 1996; Grimaldi,
1999). A rudimentary cladistic reconstruction of habitat preference among bees would
predict a distributional restriction to xeric regions for the ancestor of all Apiformes. This
is congruent with the hypothesis that the first
angiosperms were decumbent herbaceous
plants (Taylor and Hickey, 1992) and that the
climate in the Cretaceous was generally drier
than today (Spicer et al., 1993). Flowering
plants themselves likely originated (although
2001
at an earlier date) in these xeric regions (Raven and Axelrod, 1974; Taylor and Hickey,
1992). Associating this hypothesized ancestral bee habitat with the supposed area of
origination for angiosperms implicates western Gondwanaland as the most likely regional candidate. As hypothesized by Grimaldi
(1999), the significant xeric diversity of pollinating insects (including bees) may represent Cretaceous refugia. That the most
plesiomorphic and presumably ancient bees
of the paraphyletic subfamily Colletinae are
most diverse in temperate South America
and Australia is perhaps indicative of this
origination in Gondwanaland (colletines are
also represented in southern Africa by Colletes and Scrapter, but to a much lesser degree than elsewhere). It is perhaps also significant that many basal groups of the various
families show globally disjunct or, in some
cases, primarily Gondwanan distributions.
For example, aside from Colletinae, there are
the subfamilies of Halictidae (themselves
each likely ancient) that perhaps originated
early and subsequently diversified in different regions. The Andreninae are found mostly in the Laurasian continents and perhaps
diversified there after having moved out of
western Gondwanaland. The remaining basal
subfamilies of Andrenidae and basal tribes of
Panurginae are southern in distribution (the
subfamily Euherbstiinae is the sole representative of non-panurgine andreninds in southern South America with Alocandreninae
slightly farther north in Peru). The Panurginae are diverse in xeric regions and the Old
World tribes (i.e., Panurgini, Melitturgini,
and Meliturgulini) perhaps invaded through
Africa during the breakup of the continents
and then subsequently returned to western
North America in the Tertiary (i.e., within
the Panurgini). Primitive megachilids of the
Fideliinae (which occur in southern South
America and southern Africa except for one
species in Morocco) and Xylocopinae in the
Apidae also demonstrate similar patterns of
diversity. These patterns, however, must be
taken with caution since some of these
groups may have been once more diverse
than they are today. For instance, on the basis
of living taxa alone, the Lithurginae, with
basal living groups most diverse in temperate
South America, would appear to hold this
155
pattern. However, the most plesiomorphic lithurgine, P. ditomeus (see Cladistic Analyses
above), is present in Baltic amber indicating
extinct, basal lineages that were once more
globally distributed. Extinction has certainly
been a significant factor in bee evolution and
must be taken into consideration when attempting to explain present day distributions
(refer also to Grande, 1985; Grimaldi, 1992).
There is a possible taphonomic outcome
of the original habitat preference of bees that
may pose difficulties when trying to locate
mid-Cretaceous fossils. If bees originated in
xeric regions and subsequently invaded more
forested and tropical habitats, then it may become increasingly difficult to find older bee
fossils in amber since these are not the paleoenvironments thought to eventually produce amber deposits. The original bees may
simply not have occurred in regions where
amber was being formed. If this is the case,
then there will be a heavy reliance on compression fossils for locating the earliest bees.
Furthermore, the minute details needed to
distinguish an early bee from a spheciform
wasp would perhaps be hard to discern in a
mid-Cretaceous amber inclusion. Such a distinction among taxa may be overwhelmingly
difficult even in the finest of compression
fossils. Therefore, it may become increasingly challenging to piece together progressively
more ancient bee faunas.
How the transition from a prey-hunting
spheciform wasp to vegetarian bee took
place is, of course, entirely conjectural. Despite our inability to ever know how such a
significant evolutionary transition might
have unfolded, it is interesting to speculate
on this event as well as those groundplan features most likely to have characterized the
Urbiene or proto-bee. Several authors
have hypothesized about the traits of the
proto-bee with varying degrees of speculation on processes (e.g., Malyshev, 1968;
Radchenko and Pesenko, 1994a, 1994b,
1996; Michener, 2000a, 2000b). The following account presented herein is generally in
accord with most of those prior views and I
have therefore kept this discussion and my
speculative scenarios to a minimum.
By undertaking a cladistic reconstruction
for the Apiformes (figs. 122, 123) it is possible to obtain a generalized picture of the
156
Fig. 122.
NO. 259
Phylogeny of the bees (short-tongued bee families).
groundplan morphological and biological attributes that were likely present in the ancestor of all bees. The Colletidae are indeed the
most basal family of the Apiformes (figs.
121, 122) but the clade consisting of the
wasp-like subfamilies Hylaeinae, Eury-
glossinae, and Xeromelissinae is likely not to

be plesiomorphic for this family nor from
where groundplan traits of all bees should be
sought. As alluded to above, the subfamily
Colletinae as it is presently constituted is
very likely paraphyletic not only to Stenotri-
2001
Fig. 123.
Phylogeny of the bees (long-tongued bee families).
157
158
tinae and Diphaglossinae but also to the Hylaeinae 1 Euryglossinae 1 Xeromelissinae

clade. Presently, relationships among the
several genera of Colletinae are not entirely
clear so I have avoided splitting the subfamily apart for the time being. Some genera,
such as Scrapter and Callomelitta, may
prove to be more closely related to the latter
three subfamilies as well as Colletes, while
the stenotritines may be sister to a clade of
Australian colletines (e.g., some Paracolletini), thereby rendering Colletinae with its current composition paraphyletic to all other
colletids (see also McGinley, 1981). The internal phylogeny of Colletidae, however, is
in need of clarification and any discussion
(including the one presented herein) of the
proto-bee must be considered as tentative.
By this account, however, the general wasplike features of hylaeines and euryglossines
(e.g., reduced scopae) are apomorphic for
this colletid clade and not plesiomorphic for
bees. Thus, groundplan features for the bees
should be derived from plesiomorphic traits
of the Colletinae, Halictidae, and Andrenidae
as well as the remaining families. From these
considerations the proto-bee was a relatively robust, moderately pubescent, shorttongued apoid with a broad glossa. Pollen
would have been carried externally on the
body and not within the crop (as is done by
species of Hylaeinae and Euryglossinae).
Ground-nesting is a groundplan feature of
each family and thereby of the bees as a
whole; the proto-bee would have therefore
constructed simple burrows in the soil. The
stem-group wasps from which the protobee itself arose would have nested in the
soil (such a reconstruction based on the features of the spheciform wasps is well supported). Such wasps would perhaps have
brought back pollen-covered prey that females had captured on early flowering plants.
Once the wasp larva had by chance become
able to digest and make use of the pollen as
a protein resource, it was perhaps not long
before pollen became the primary, if not sole,
source of food. A female wasp would have
a decreased search-time for provisions owing
to the fact that flowers advertise themselves
for pollination and are stationary while prey
attempt to hide and are mobile. Moreover,
negative consequences such as injury and
NO. 259
risk of death to the provisioning female from

those prey that might fight back would be
immediately given up. Such presumably
strong selective pressures in this scenario
would quickly result in any such lineage of
wasps to become vegetarian. The plumose
setae, a distinguishing trait of bees, were
likely not an original adaptation for the transport of pollen. Plumose setae were perhaps,
as discussed by Michener (2000a), a means
of conserving water in an arid environment.
Branched setae are not a necessity for the
transport of pollen since many bees have
simple scopal setae. As mentioned above,
bees likely arose in the xeric interior of
Gondwanaland and plumose setae would potentially have been a novel mechanism for
decreasing integumental evaporation (in contrast to simply increasing the total number of
setae). As also pointed out by Michener
(2000a), that such setae are generally pale
(white or off-white) and are a groundplan
feature for the bees is also suggestive since
many xeric-environment insects presumably
use pale hue to reflect heat and as protective
coloration. As just mentioned, pilosity serves
a thermoregulatory function (e.g., Church,
1960). In hovering and other strong flying,
pollinating insects, like bees and beeflies,
thick pilosity also serves to maintain a high
body temperature, especially as flight begins
after torpor from cold, desert nights. Clearly,
some desert bees have minimal setae (e.g.,
some Xylocopa) but those that fly earlier in
the morning tend to be more setose. Plumose
setae in the proto-bee likely played a role
in thermoregulation and water conservation
rather than in pollen transport.
The major lineages of bees (i.e., families,
perhaps some subfamilies) perhaps diversified rapidly shortly following the origin of
bees as a whole (figs. 121123). The division
into those ancestral lineages that eventually
gave rise to the various families recognized
today must have taken place sometime between 12590 Ma. These families then progressed along independent evolutionary
tracks, undoubtedly with numerous groups
that exhibited interesting character combinations becoming extinct at various stages
(and as is documented above, several by the
latest Eocene at least since only modern
groups are known in the Oligocene and
2001
younger deposits), leaving the six families

we recognize today. The family Paleomelittidae represents one such intermingled lineage whose extinction left large character
gaps between the surviving lineages. Such
extinct lineages were perhaps dwindling by
middle Eocene times. At present there should
be little or no concern for the absence of a
tight correlation between the stratigraphic
ranks and phylogenetic ranks of bee taxa.
Significant correlations have been found in a
variety of taxa where the fossil record is relatively well represented and cladograms are
well resolved (e.g., Norell and Novacek,
1992a, b; Borkent, 1995; Grimaldi and Cumming, 1999). For the bees, however, the fossil record is comparatively sparse, with geological data points based on limitations: 1. in
number of well preserved specimens upon
which to base character analyses; 2. in chronological diversity and number of horizons
with identifiable bee specimens (only a few,
scattered deposits have at present revealed
fossil bees); 3. in diversity and number of
higher-level lineages (e.g., tribes) with a fossil record of any sort [e.g., see appendix 2,
where of 83 suprageneric groups (not including subtribes) only 28 are represented in the
fossil record, and most of these are based on
dubiously assigned compression fossils (personal obs.)]; and 4. in number of cladistic
analyses for bee groups.
Although bees are quite rare as amber inclusions (except for the common Proplebeia
dominicana in Dominican amber), the exploration of Cretaceous amber sites for older
bees is imperative, no matter how seemingly
futile. Compression fossils will also play an
important, albeit somewhat more limited,
role in understanding bee evolution and may
eventually prove to be critical for understanding the earliest of bees. Such exploration of the Cretaceous will hopefully bring
to science additional lineages that close the
gaps in our knowledge and help us to further
refine the scenario of bee diversification I
have presented. It is from such Mesozoic fossils (particularly those preserved in amber)
that the greatest contributions will come for
not only understanding bee origins but for
reconstructing the pattern of bee phylogeny.
159
THE EOCENE BEE FAUNA

A general pattern of affinity between the
Eocene fauna of Europe and the present day
faunas of Asia and sub-Saharan Africa is
documented for various groups (e.g., Ander,
1942; Larsson, 1978). For example, the famous Baltic amber lizard has affinities to
groups in Africa and was even assigned, until
recently, to the living African genus Nucras
(Bohme and Weitschat, 1998). Likewise, the
Baltic amber scorpion Palaeolychras balticus Lourenco and Weitschat has as its nearest
relative a genus presently occurring in Africa
and southern Asia (Lourenco and Weitschat,
1996). A similar pattern certainly holds true
for the Baltic amber bees. The Boreallodapini is allied to the Allodapini, a tribe that
has its greatest diversity in sub-Saharan Africa, southeast Asia, and Australia (see also
discussion above under Cladistic Analyses).
The genera Kelneriapis, Liotrigonopsis, and
Electrolictus are all most similar and certainly related to genera today occurring in Africa
and southeast Asia (namely Hypotrigona,
Liotrigona, and Patellapis, respectively).
Similarly, the Melikertini is sister to Meliponini, a group that is today pantropical in
distribution and the Glyptapina have affinities with the Old World (principally African)
Anthidiini.
The bee fauna of the Baltic amber is
unique, with a wider array of bees than any
other Lagerstatte in the world. The fauna is
further important for the preservation of a
number of primitive, higher groups that
failed to survive to the present day. This is
in sharp contrast to younger fossil faunas
from the Oligocene or Miocene where species belong to extant genera or extinct genera
closely related to modern groups and in all
cases to living tribes, subfamilies, and families (e.g., lowest Miocene Dominican amber:
table 1; Oligocene of Germany: Engel,
1998c). It is possible that some of the extinct
lineages from the Baltic amber fauna are the
final Tertiary survivors of originally Mesozoic bee lineages. Owing to the presence of
a meliponine in latest Cretaceous amber (Engel, 2000b), this is almost certainly the case
for the tribes Electrobombini, Electrapini,
and Melikertini as well as the Paleomelittidae. The almost modern character of the only
160
halictine in the Baltic amber fauna, Electrolictus, is indicative of not only this tribes antiquity, but of that for the whole family.
From the available paleomelittological information it would appear that the bees underwent at least two periods of diversification.
The first was either at their origin or shortly
thereafter and produced the array of taxa that
eventually gave rise to families we recognize
today. The second radiation appears to have
been in the Tertiary. As mentioned repeatedly above, those taxa from the Eocene and
earlier all belong to extinct genera and,
where information is available, most belong
to higher categories (i.e., tribes, subfamilies,
and families) that are no longer extant. Immediately following the Eocene and by the
earliest Miocene the bee fauna becomes remarkably modern in appearance, with most
fossils belonging to living genera. There are
no extinct suprageneric lineages in Oligocene
or younger deposits. The bees in the Eocene
and perhaps older deposits were presumably
remnants of originally Cretaceous diversifications. At the end of the Eocene those enigmatic higher lineages were perhaps dwindling and, at least in Europe, their eventual
disappearance was possibly aided by the
changing global climate at the time. Thus, at
the Eocene-Oligocene transition there was a
dramatic shift in the tribal composition of the
bee fauna to entirely modern tribes and essentially modern genera that either originated
or first experienced their greatest radiation at
that time. Thus, the bee fauna we are familiar
with today appears to have had its compositional origin in the Oligocene Epoch.
It is significant that there is such a remarkable diversity of advanced eusocial corbiculate bees in the Eocene fauna and more
so that most of these taxa represent higher
lineages that have since become extinct (Engel, in press-b). A general, epoch-by-epoch
overview of the geological history of the corbiculate Apinae (e.g., Engel, 2000c) shows a
decrease in diversity at higher categorical
levels (e.g., genera and tribes), particularly
between the Eocene and Oligocene Epochs.
During this transition period numerous tribes
and genera of advanced eusocial lineages
disappeared so that by the Oligocene the corbiculate bee fauna was essentially modern in
character and, in fact, the bee fauna as a
NO. 259
whole became very modern at this period.

This is noteworthy since the success, as measured by ecological dominance and specific
diversity, of groups such as the termites (ca.
3,000 species), ants (ca. 16,000 species), and
social wasps (ca. 1,000 species) has been directly attributed to their advanced eusocial
behavior. The bees have been understandably
included with these lineages since bees are
undoubtedly more diverse than all three combined and arguably more ecologically dominant owing to their role as the most significant pollinators of angiosperms. Bees as a
whole, however, are mostly solitary, while
the advanced eusocial lineages (all restricted
to the corbiculate Apinae) account for merely
ca. 380 species. Not surprisingly, pollination
has likely driven the overall specific diversity
of bees but counter-intuitively, it appears that
advanced eusocial behavior has not further
fueled diversity in those lineages where it has
arisen. In fact, it could even be argued that
such highly organized and ecologically aggressive societies have hindered the specific
diversity of the corbiculate bees over geological time. Advanced eusocial species, like the
western honey bee, Apis mellifera, are
known to influence the foraging patterns and
abundance of other species through competition for resources (Eickwort and Ginsberg,
1980; Wenner and Thorp, 1994; Buchmann,
1996). The diversity of native bees has been
negatively impacted, including that of other
advanced eusocial lineages, in areas where A.
mellifera has been introduced (e.g., Roubik
et al., 1986; Sugden et al., 1996). This negative interaction is intense enough that extinction of stingless bee colonies was one of
the predicted likely outcomes of the competitive interactions between introduced honey
bees and native Meliponini (Roubik et al.,
1986). Introduced honey bees not only negatively affect native bees but also nectarfeeding bird populations (Paton, 1993). Even
amongst the living honey bees it has been
observed that introduced populations of A.
mellifera aggressively displace A. cerana
(e.g., Sakagami, 1959). Interestingly, in Apis,
the most ecologically dominant and aggressive eusocial group, the specific diversity is
at its lowest for any advanced eusocial lineage (Engel, 1999c). The generally less aggressive Meliponini are not only more spe-
2001
ciose than Apini, but the tribe is most diverse

in regions where Apis species are not native
(i.e., South and Central America). It is possible that over more expansive reaches of
time such ecologically dominant foragers
have had a more devastating impact on not
only non-social species but also and perhaps
especially on other advanced eusocial lineages that were not as efficient competitors.
Certainly, extrinsic factors have played a role
in the diversity and distribution of bees and
continued paleontological work throughout
the world will contribute to either supporting
or refuting this hypothesis.
EPILOGUE
The study of amber inclusions allows the
scientist to peer directly into past eons with
far greater clarity than is possible from other
kinds of fossils. In the case of the present
work, these tiny bees, perhaps guards from
some ancient colonies, have waited attentatively while 45 million years of history
passed in review. Arriving in life-like form,
surviving untold geological turmoil, these
tiny mummies have opened a window onto
a world since lost, utterly reshaping our
knowledge of bee evolution. I must admit to
sharing the sentiment expressed by Rev.
Hope (1836), who, in regard to his own study
of Baltic amber insects, wrote (p. 137):
Let then the geologist boast of his Mastodon and Megatherium, his wondrous Saurian Reptiles, and numerous genera of unknown beings; the entomologist also
prides himself on his Amber Insects, . . . , surpassing
them in beauty and in colouring. Let the former add
room to room, gallery to gallery, and fill a city with
his relics and his casts, the latter may also rest contented with his stores, less bulky indeed although not
less interesting. The earth is a study for them both, an
inexhaustible field of inquiry, and it may be a matter
of question which will most contribute to illustrate the
condition and character of its earliest tenants.
Both the Age of Dinosaurs and Age

of Mammals, periods frequently referred to
by scientists and laymen alike, are eclipsed
by an Age of Insects. From the late Paleozoic
to the present day, insects have dominated
terrestrial and freshwater ecosystems. A paleontological perspective of insect diversity
will, therefore, provide a powerful complement, if not in some instances a more significant component, to the reconstruction of
161
ancient ecologies, patterns of evolution, and

past epochs of life on Earth.
ACKNOWLEDGMENTS
This work would not have been possible
without the generous support of Robert G.
Goelet, Chairman Emeritus of the AMNH
Board of Trustees. I am sincerely grateful for
his patronage and admirable support of collections-based research. Jerome G. Rozen,
Jr., has been a close mentor and friend as
well as serving as a sounding board during
the past two years for ideas presented herein.
Charles D. Michener continually provided
support and advice; I am particularly grateful
for his time and patience during my visits to
discuss fossils, phylogeny, and evolutionary
trends in bees. David A. Grimaldi has enthusiastically supported my work in paleoentomology, giving access to equipment, funds,
and supplies, as well as providing valuable
advice and friendship. Molly G. Rightmyer
masterfully executed the numerous line illustrations in this monograph; her patient and
meticulously careful work has greatly enhanced the presentation of this material.
My work also has been greatly facilitated
by assistance and/or advice received either
formally, during visits to foreign institutions,
or during casual discussions from Alger G.
Engel, Donna G. Engel, Hans Jahnke, Jan
Koteja, Kumar Krishna, Valerie Krishna,
Malcolm C. McKenna, Paul Nascimbene,
Tam C. Nguyen, Michael Ohl, Erika Pietrzeniuk, Andrew J. Ross, and Wolfgang
Weitschat. This monograph would have been
far less fruitful had it not been for the generosity of the private collectors listed on pp.
1718 (Material and Methods) who patiently
allowed me to study their specimens; I am
sincerely grateful to each. Barbara Kosmowska-Ceranowicz kindly supplied copies of her
work and Wolfgang Weitschat copies of recent work on Baltic amber stratigraphy and
dating; the latter also acted as courier for numerous specimens from private collections.
Although I could not agree with his conclusions, I am appreciative to Stephen T. Hasiotis for generously supplying copies of his
work concerning trace fossils in North America. Jorg Wunderlich, Jens von Holt, and
Bernard Striebich made Baltic amber bees
162
available to the AMNH and myself; their

support and interest was of great value. Lastly, I am thankful to Michael Ohl and his family for kindly hosting my stay in Berlin, Erika Pietrzeniuk for assistance while in the Institut fur Palaontologie (Berlin), Hans Jahnke
for providing lodging and assistance in the
Institut und Museum fur Geologie und Palaontologie (Gottingen), and to Kumar and
Valerie Krishna for graciously hosting me
during my stay in London. Oliver C. Phillips
and Stanley Lombardo, of the University of
Kansas Classics Department, kindly helped
with Latin translations.
I am enormously indebted to Lee Herman,
Brenda Jones, and Robert Voss for their help
with the publication of my manuscript. The
manuscript received critiques from David A.
Grimaldi, Lee Herman, Zhiwei Liu, Charles
D. Michener, Molly G. Rightmyer, Jerome G.
Rozen, Jr., and two anonymous reviewers. I
am grateful to each for their time and patience. Not all agreed with my conclusions
and any remaining errors or peculiar interpretations are, naturally, my own.
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APPENDIX 1
Bees in Amber and Copal

The following list enumerates those valid species presently recognized as inclusions in both amber and copal (synonymous names
are not listed). The list excludes incertae sedis taxa. All species known from amber are extinct while only one of those recognized
from copal is considered an extinct species (Liotrigona vetula).
The two meliponine species listed as in Sicilian amber (Tosi, 1896) may actually be in East African copal and synonymous with
Recent stingless bee species. Unfortunately, Tosis specimens were destroyed by a bomb during World War II and their true identity
will, therefore, forever remain a mystery. Apis catanensis, also from Sicilian amber, is listed, As discussed by Engel (1999c), A.
catanensis is a nomen dubium and was perhaps not a bee at all (as suspected by Kohring and Schluter, 1989); or, it may have been
a living bee (A. mellifera) in African copal. Owing to the uncertainty over their identities, I have retained all of the Sicilian specimens
in the list as bees occurring in true amber.
The Eocene amber of Arkansas is rich in insert inclusions. It will be interesting to see if bees are ever discovered from these
deposits. Owing to the relative contemporaneous age with the Baltic amber fauna, the possibility of discovering similar taxa is
tantalizing.
An annotated, general catalog of the worlds species of fossil bees (both amber inclusions and compression fossils) as well as trace
fossils attributed to bees will be presented elsewhere (Engel, in prep.)
176
APPENDIX 1(Continued )
NO. 259
2001
177
APPENDIX 2
Tribal Classification and Geological Distribution of Bees

The following outline presents the tribal classification of bees adopted by the present author based on continuing
cladistic investigations into the higher-level phylogeny of Apoidea (figs. 122, 123). Some of these analyses have been
presented above while others will be detailed in forthcoming papers. Those groups presently recorded from the
geological record are followed by the standard geological abbreviation for their oldest epoch/stage [e.g., although
numerous Miocene to Eocene fossils exist for Meliponini only Cretaceous/Maastrichtian (K/Maa) is listed owing to
a single, older fossil from this period]. Compression fossils are indicated in boldface by CF, amber inclusions by A.
Daggers () denote higher groups that are extinct.
178
NO. 259
2001
179
180
NO. 259
APPENDIX 3
Catalog of Family- and Genus-Group Names
for Fossil Bees
The following catalog presents all known family- and genus-group names proposed for fossil
bees (amber inclusions and compression fossils).
Not all of the names are currently recognized as
valid (e.g., Sophrobombus is a synonym of Protobombus; Paleoeuglossa is a synonym of Eufriesea) or are confidently assigned to higher categories (e.g., Protomelecta); thus this list should
not be taken as a classification. At the end of the
list I have included two genus-group names originally proposed as fossil bees but subsequently
recognized to be insects other than bees. A catalog of the worlds species of fossil bees (both amber inclusions and compression fossils) will be
presented elsewhere (Engel, in prep.).
The format is generally that of Michener (1986)
for family-group names and Michener (1997) for
genus-group names. For family-group names I
have indicated the higher group to which a given
taxon belongs. The number that indicates the location of the name in the system of Michener
(1986: with additions and corrections by Michener, 1997) is appended at the end of each entry.
The citation Engel (2001) in the following
list refers to the present monograph and those
names newly proposed in the body of the text;
thus, this reference will not be found in the bibliography.
Family-Group Names
(10 names)
Boreallodapini Engel, 2001: 77. Type genus:
Boreallodape Engel, 2001. Stem: Boreallodap-.
Apidae: Xylocopinae [M86/97120.2].
Chalicodomopsini Engel, 1999d: 4. Type genus: Chalicodomopsis Engel, 1999d. Stem: Chalicodomops-. Megachilidae: Megachilinae [M86/
9767.2]. Nomen nudum.
Ctenoplectrellina Engel, 2001: 54. Type genus: Ctenoplectrella Cockerell, 1909a. Stem:
Ctenoplectrell-. Megachilidae: Megachilinae
[M86/9767.3].
Electrapina Engel, 1998a: 99. Type genus:
Electrapis Cockerell, 1908b. Stem: Electrap-. Apidae: Apinae [M86/97129.1].
Electrobombini Engel, 2001: 88. Type genus:
Electrobombus Engel, 2001. Stem: Electrobomb-.
Apidae: Apinae [M86/97129.2].
Eomacropidini Engel, 2001: 46. Type genus:
Eomacropis Engel, 2001. Stem: Eomacropid-.
Melittidae: Macropidinae [M86/9757.2].
Glyptapinae Cockerell, 1909b: 13. Type ge-
nus: Glyptapis Cockerell, 1909a. Stem: Glyptap-.

Megachilidae: Megachilinae [M86/9767.1].
Melikertini Engel, 2001: 112. Type genus: Melikertes Engel, 1998a. Stem: Melikert-. Apidae:
Apinae [M86/97129.3].
Paleomelittidae Engel, 2001: 41. Type genus:
Paleomelitta Engel, 2001. Stem: Paleomelitt-. Paleomelittidae [M86/9757.1].
Protolithurgini Engel, 2001: 51. Type genus:
Protolithurgus Engel, 2001. Stem: Protolithug-.
Megachilidae: Lithurginae [M86/9761.1].
Genus-Group Names
(47 names)
Anthophorites Heer, 1849: 97. Type species:
Anthophorites mellona Heer, 1849, by designation
of Cockerell, 1909a. Apidae: Anthophorini?.
Apiaria Germar, 1839: 210. Type species: Apiaria dubia Germar, 1839, monobasic. Megachilidae: Osmiini?.
Bombusoides Motschulsky, 1856: 28. Type
species: Bombusoides mengei Motschulsky, 1856,
monobasic. Incertae sedis.
Boreallodape Engel, 2001: 78. Type species:
Boreallodape baltica Engel, 2001, original designation. Xylocopinae: Boreallodapini.
Calyptapis Cockerell, 1906: 41. Type species:
Calyptapis florissantensis Cockerell, 1906, monobasic. Apidae: Bombini.
Cascapis Engel, 1999c: 187. Type species: Apis
armbrusteri Zeuner, 1931, monobasic and original
designation, isotypic with Hauffapis. Proposed as
a subgenus of Apis.Apidae: Apini.
Chalcobombus Cockerell, 1908b: 326. Type
species: Chalcobombus humilis Cockerell, 1908b
[5 Protobombus indecisus Cockerell, 1908b], by
designation of Cockerell, 1909b; isotypic with
Protobombus Cockerell, 1908b, by synonymy.
Apidae: Apinae: Electrapini.
Chalicodomopsis Engel, 1999d: 3. Type species: Megachile (Chalicodomopsis) glaesaria Engel, 1999d, monobasic and original designation.
Proposed as a subgenus of Megachile. Megachilidae: Megachilini.
Cretotrigona Engel, 2000b: 3. Type species:
Trigona (Trigona) prisca Michener and Grimaldi,
1988a, monobasic and original designation. Apidae: Meliponini.
Ctenoplectrella Cockerell, 1909a: 314. Type
species: Ctenoplectrella viridiceps Cockerell,
1909a, monobasic. Megachilidae: Megachilinae:
Osmiini.
Cyrtapis Cockerell, 1908a: 339. Type species:
Cyrtapis anomalus Cockerell, 1908a, monobasic.
Halictidae: Halictini.
Eckfeldapis Lutz, 1993: 180. Type species:
2001
Eckfeldapis electrapoides Lutz, 1993, monobasic

and original designation. Apidae: Apinae: Electrapini.
Eickwortapis Michener and Poinar, 1996
[1997]: 354. Type species: Eickwortapis dominicana Michener and Poinar, 1996, monobasic and
original designation. Halictidae: Caenohalictini.
Electrapis Cockerell, 1908b: 326. Type species: Apis meliponoides Buttel-Reepen, 1906, by
designation of Cockerell, 1909b. Apidae: Apinae: Electrapini.
Electraugochlora Engel, 2000a: 32. Type species: Augochlora (Electraugochlora) leptoloba
Engel, 2000a. Proposed as a subgenus of Augochlora. Halictidae: Augochlorini.
Electrobombus Engel, 2001: 88. Type species:
Electrobombus samlandensis Engel, 2001, monobasic and original designation. Apidae: Apinae:
Electrobombini.
Electrolictus Engel, 2001: 38. Type species:
Electrolictus antiquus Engel, 2001, monobasic
and original designation. Halictidae: Halictini.
Eomacropis Engel, 2001: 46. Type species:
Eomacropis glaesaria Engel, 2001, monobasic
and original designation. Melittidae: Macropidinae: Eomacropidini.
Glaesosmia Engel, 2001: 75. Type species:
Glaesosmia genalis Engel, 2001, monobasic and
original designation. Megachilidae: Megachilinae: Osmiini.
Glyptapis Cockerell, 1909a: 314. Type species:
Glyptapis mirabilis Cockerell, 1909a, monobasic.
Megachilidae: Megachilinae: Osmiini.
Hauffapis Armbruster, 1938: 37. Type species:
Hauffapis scheuthlei Armbruster, 1938 [5Apis
armbrusteri Zeuner, 1931], by designation of Zeuner and Manning, 1976, isotypic with Cascapis.
Unavailable as no type species was originally designated (required after 1930 by ICZN, 1999b: Art.
13.3). Note: Zeuner and Mannings action does
not validate the name with themselves as authors
as it was in the synonymy of the genus. Apidae:
Apini.
Kelneriapis Sakagami, 1978 (June): 232. Type
species: Hypotrigona eocenica Kelner- Pillault,
1969a, monobasic, isotypic with Kelnermelia.
Proposed as a subgenus of Tetragonula. Apidae:
Meliponini.
Kelnermelia Moure In Moure and Camargo,
1978 (November): 565. Type species: Hypotrigona eocenica Kelner-Pillault, 1969a, monobasic
and original designation, isotypic with Kelneriapis. Apidae: Meliponini.
Libellulapis Cockerell, 1906: 42. Type species:
Libellulapis antiquorum Cockerell, 1906, monobasic. Andrenidae: Panurginae?.
Liotrigonopsis Engel, 2001: 135. Type species:
181
Liotrigonopsis rozeni Engel, 2001, monobasic and

original designation. Apidae: Meliponini.
Lithandrena Cockerell, 1906: 44. Type species: Lithandrena saxorum Cockerell, 1906,
monobasic. Andrenidae: Andreninae?.
Lithanthidium Cockerell, 1911b: 225. Type
species: Lithanthidium pertriste Cockerell, 1911b,
monobasic. Megachilidae: Anthidiini.
Melikertes Engel, 1998a: 95. Type species:
Electrapis (Melikertes) stilbonota Engel, 1998a,
original designation. Proposed as a subgenus of
Electrapis. Apidae: Apinae: Melikertini.
Meliponorytes Tosi, 1896: 352. Type species:
Meliponorytes succini Tosi, 1896, by designation
of Sandhouse, 1943. Apidae: Meliponini.
Melissites Engel, 2001: 129. Type species: Melissites trigona Engel, 2001, monobasic and original designation. Apidae: Apinae: Melikertini.
Oligochlora Engel, 1996 [1997]: 336. Type
species: Oligochlora eickworti Engel, 1996, original designation. Halictidae: Augochlorini.
Paleoeuglossa Poinar, 1998 [1999]: 30. Type
species: Paleoeuglossa melissiflora Poinar, 1998,
monobasic and original designation. Apidae:
Euglossini.
Paleomelitta Engel, 2001: 42. Type species:
Paleomelitta nigripennis Engel, 2001, monobasic
and original designation. Paleomelittidae.
Pelandrena Cockerell, 1909d: 159. Type species: Pelandrena reducta Cockerell, 1909d,
monobasic. Andrenidae: Andreninae?.
Priorapis Engel, 1999c: 188. Type species:
Apis vetusta Engel, 1998c, monobasic and original designation. Proposed as a subgenus of Apis.
Apidae: Apini.
Probombus Piton, 1940: 218. Type species:
Probombus hirsutus Piton, 1940, monobasic. Apidae: Bombini.
Prohalictus Armbruster, 1938: 48. Type species: Prohalictus schemppi Armbruster, 1938,
monobasic. Halictidae: Halictini.
Proplebeia Michener, 1982: 44. Type species:
Trigona (Liotrigona) dominicana Wille and Chandler, 1964, monobasic and original designation.
Proposed as a subgenus of Trigona. Apidae: Meliponini.
Protobombus Cockerell, 1908b: 326. Type species: Protobombus indecisus Cockerell, 1908b,
monobasic [also designated by Cockerell, 1909b],
isotypic with Chalcobombus Cockerell, 1908b, by
synonymy. Apidae: Apinae: Electrapini.
Protolithurgus Engel, 2001: 51. Type species:
Protolithurgus ditomeus Engel, 2001, monobasic
and original designation. Megachilidae: Lithurginae: Protolithurgini.
Protomelecta Cockerell, 1908a: 341. Type species: Protomelecta brevipennis Cockerell, 1908a,
monobasic. Apidae: Melectini?.
182
Roussyana Manning, 1960 [1961]: 306. Type

species: Apis palmnickenensis Roussy, 1937, original designation. Proposed as a subgenus of Electrapis. Apidae: Apinae: Melikertini.
Soliapis Engel, 2000a: 47. Type species: Oligochlora (Soliapis) rozeni Engel, 2000a, original
designation. Proposed as a subgenus of Oligochlora. Halictidae: Augochlorini.
Sophrobombus Cockerell, 1908b: 326. Type
species: Sophrobombus fatalis Cockerell, 1908b,
monobasic [also designated by Cockerell, 1909c].
Apidae: Apinae: Electrapini.
Succinapis Engel, 2001: 115. Type species:
Succinapis goeleti Engel, 2001, original designation. Apidae: Apinae: Melikertini.
Synapis Cockerell, 1907: 229. Type species:
Apis (Synapis) henshawi Cockerell, 1907, monobasic. Proposed as a subgenus of Apis. Apidae:
Apini.
NO. 259
Thaumastobombus Engel, 2001: 109. Type

species: Thaumastobombus andreniformis Engel,
2001, monobasic and original designation. Apidae: Apinae: Electrapini.
Non-Bees
Palaeapis Hong, 1984: 37. Type species: Palaeapis beiboziensis Hong, 1984, monobasic and
original designation. Originally proposed in Apidae; presently a junior synonym of Archisphex
(Rasnitsyn et al., 1998). Sphecidae.
Sinostigma Hong, 1983: 9. Type species: Sinostigma spinalata Hong, 1983, monobasic and
original designation. Originally proposed in Melittidae; later transferred to Megachilidae (Hong,
1985). Vespidae?.
APPENDIX 4
Full Journal Titles Abbreviated in References
The following list provides full journal titles for
abbreviations in the References section. Abbreviations are listed in alphabetical order in boldface, followed by the expanded title. Specific series or report numbers are replaced in this list by
#.
Acta Zootaxon. Sinica Acta Zootaxonomica
Sinica
Am. Chem. Soc. American Chemical Society
Am. J. Bot. American Journal of Botany
Am. Midl. Nat. American Midland Naturalist
Am. Mus. Novitates American Museum Novitates
Ann. Entomol. Soc. Am. Annals of the Entomological Society of America
Ann. Geol. Pays Helleniques Annales Geologiques des Pays Helleniques
Ann. Mag. Nat. Hist., ser. # Annals and Magazine of Natural History, series #
Ann. Missouri Bot. Gard. Annals of the Missouri Botanical Garden
Ann. Natal Mus. Annals of the Natal Museum
Ann. Soc. Entomol. Fr. Annales de la Societe
Entomologique de France
Annu. Rev. Ecol. Syst. Annual Review of Ecology and Systematics
Annu. Rev. Entomol. Annual Review of Entomology
Appl. Spectrosc. Rev. Applied Spectroscopy
Reviews
Arch. Bienenkd. Archiv fur Bienenkunde
Arq. Mus. Parana. Arquivos do Museu Paranaense
Aust. J. Sci. Australian Journal of Science
Aust. Zool. Australian Zoologist

Belg. J. Entomol. Belgian Journal of Entomology
Bernstein-Forsch. Bernstein-Forschungen
Biol. J. Linn. Soc. Biological Journal of the
Linnean Society
Biol. Zentbl. Biologisches Zentralblatt
Biul. Inst. Geol. Biuletyn Instytutu Geologicznego
Bol. Soc. Amig. Cienc. Nat. Kraglievich-Fontana Boletn de la Sociedad Amigos de las
Ciencias Naturales Kraglievich-Fontana
Bol. Univ. Fed. Parana, Zool. # Boletim da
Universidade Federal do Parana, Zoologia
Bot. Gaz. Botanical Gazette
Bull. Am. Mus. Nat. Hist. Bulletin of the
American Museum of Natural History
Bull. Br. Mus. Nat. Hist. (Entomol.) Bulletin
of the British Museum of Natural History (Entomology)
Bull. Br. Mus. Nat. Hist. (Geol.) Bulletin of
the British Museum of Natural History (Geology)
Bull. Mus. Comp. Zool. Bulletin of the Museum of Comparative Zoology
Bull. Nat. Hist. Mus., London (Geol.) Bulletin
of the Natural History Museum, London (Geology)
Bull. Pol. Acad. Sci. Bulletin of the Polish
Academy of Sciences
Bull. Soc. Entomol. France Bulletin de la Societe Entomologique de France
Bull. Tianjin Inst. Geol. Miner. Res. Bulletin
of the Tianjin Institute of Geology and Mineral
Resources
2001
Bull. Zool. Nomencl. The Bulletin of Zoological Nomenclature

Can. Entomol. The Canadian Entomologist
asopis Moravskeho
Cas. Morav. Mus. Brne C
musea v Brne
Chacaras Quint. (Sao Paulo) Chacaras e Quintais, Sao Paulo
Contrib. Am. Entomol. Inst. Contributions of
the American Entomological Institute
Contrib. Geol., Univ. Wyoming.
Contributions to Geology, University of Wyoming
Cretac. Res. Cretaceous Research
Dalle Mem. Soc. Entomol. Ital. Dalle Memorie
della Societa Entomologica Italiana
Denkschr. Kaiserl. Akad. Wiss., Math.-Natwiss. Cl. Denkschriften der Kaiserlichen
Akademie der Wissenschaften [Wien], Mathematisch-Naturwissenschaftliche Classe
Doc. Nat. Documenta Naturae
Dtsch. Entomol. Z. Detusche Entomologische
Zeitschrift
Entomol. Entomologist
Entomol. Mag. Entomological Magazine
Entomol. News Entomological News
Entomol. Obozr. Entomologicheskoye Obozreniye
Entomol. Rev. Entomological Review
Entomol. Scand. Entomologica Scandinavica
EPHE Biol. Evol. Insectes EPHE Biologie et
Evolution des Insectes
Etude Entomol. Etude Entomologiques
Fortschr. Geol. Palaeontol. Fortschritte der
Geologie und Palaeontologie
Gaz. Apic., Montfavet La Gazette Apicole,
Montfavet
J. Anim. Ecol. Journal of Animal Ecology
J. Baltic Stud. Journal of Baltic Studies
J. Evol. Biol. Journal of Evolutionary Biology
J. Exp. Biol. Journal of Experimental Biology
J. Fac. Sci., Hokkaido Univ., ser. #
(Zool.) Journal of the Faculty of Science,
Hokkaido University, series # (Zoology)
J. Hym. Res. Journal of Hymenoptera Research
J. Kansas Entomol. Soc. Journal of the Kansas
Entomological Society
J. Kansas Entomol. Soc., Suppl. Journal of the
Kansas Entomological Society, Supplement
J. New York Entomol. Soc. Journal of the New
York Entomological Society
J. Paleontol. Journal of Paleontology
J. Trop. Ecol. Journal of Tropical Ecology
Jahrb. Nassau. Ver. Naturkd. Jahrbucher des
Nassauischen Vereins fur Naturkunde
Jahrb. Ver. Naturkd. Herzogthum Nassau
Jahrbucher des Vereins fur Naturkunde im Herzogthum Nassau
Linzer Biol. Beitr. Linzer Biologische Beitrage
rskr., 2 Afd. Lunds Universitets
Lunds Univ. A
183
rsskrift, 2 Afdelning, Medicin samt MatemaA

mnen [Parallel
tiska och Naturvetenskapliga A
serial title: Acta Universitatis Lundensis]
Mag. Insektenkd. Magazin fur Insektenkunde
Mainzer Natwiss. Arch. Mainzer naturwissenschaftliches Archiv
Mem. Beijing Nat. Hist. Mus. Memoirs of the
Beijing Natural History Museum
Mitt. Geol.-Palaontol. Inst. Univ. Hambg. Mitteilungen aus dem Geologisch-Palaontologischen Institut der Universitat Hamburg
Mitt. Zool. Mus. Berl. Mitteilungen aus dem
Zoologischen Museum in Berlin
Mol. Phylogenet. Evol. Molecular Phylogenetics and Evolution
Munster. Forsch. Geol. Palaontol.
Munstersche Forschungen zur Geologie und Palaontologie
Mus. North. Arizona Bull. Museum of Northern Arizona Bulletin
Natl. Park Serv., Paleontol. Res. [Tech. Rep.
#] National Park Service, Paleontological Research [Technical Report]
Neue Denkschr. Allg. Schweiz. Gesell. Ges. Natwiss. Neue Denkschriften der Allgemeinen
Schweizerischen Gesellschaft fur die gesammten Naturwissenschaften
Nova Acta Acad. C. Leopold.-Carol. Germ.
Nat. Cur. Nova Acta Academiae Caesarea
Leopoldino-Carolinae Germanicum Naturae
Curiosorum
Nova Acta Leopold. (Kaiserl. Leopold.-Carol.
Dtsch. Akad. Naturf.) Nova Acta Leopoldina (Kaiserlich Leopoldinisch-Carolinisch
Deutschen Akademie der Naturforscher)
Orient. Insects Oriental Insects
Palaeogeogr.,
Palaeoclimatol.,
Palaeoecol. Palaeogeography, Palaeoclimatology, Palaeoecology
Palaeontol. Z. Palaeontologische Zeitschrift
Pan-Pac. Entomol. Pan-Pacific Entomologist
Pap. Avulsos Zool. (Sao Paulo) Papeis avulsos
de Zoologia, Sao Paulo
Philipp. Entomol. Philippine Entomologist
Philos. Trans. R. Soc. London, ser. B, Biol.
Sci. Philosophical Transactions of the Royal
Society of London, series B, Biological Sciences
Plant Syst. Evol.5Plant Systematics and Evolution
Plateau J., Mus. North. Arizona5Plateau Journal, Museum of Northern Arizona
Proc. Acad. Nat. Sci., Philadelphia Proceedings
of the Academy of Natural Sciences, Philadelphia
Proc. Natl. Acad. Sci., U.S.A. Proceedings of
the National Academy of Sciences of the United States of America
184
Proc. First Intl. Palaeoentomol. Conf.,

Moscow Proceedings of the First International Palaeoentomological Conference, Moscow
[Russia]
Proc. VI Congr. IUSSI, Bern Proceedings VI
Congress of the International Union for the
Study of Social Insects, Bern [Switzerland]
Proc. 13th Intl. Congr. IUSSI, AdelaideProceedings of the 13th International Congress
of the International Union for the Study of Social Insects, Adelaide [Australia]
Proc. U.S. Natl. Mus. Proceedings of the United States National Museum
Prof. Pap. Stratigr. Paleontol. Professional Papers of Stratigraphy and Paleontology
Rev. Biol. Trop. Revista de Biologa Tropical
Rev. Palaeobot. Palynol. Review of Palaeobotany and Palynology
Rev. Zool. Bot. Afr. Revue de Zoologie et de
Botanique Africaines
Riv. Ital. Paleontol., Bologna Rivista Italiana
di Paleontologia, Bologna [Italy]
Rocky Mountain Sec., Geol. Soc. Am. Reg.
Meeting Rocky Mountain Section, Geological Society of America Regional Meeting
Russian Acad. Sci. Russian Academy of Sciences
Schr. Naturf. Gesell. Danzig Schriften der Naturforschenden Gesellschaft in Danzig
kon. Gesell. Schriften der PhySchr. Phys.-O
sikalisch-okonomischen Gesellschaft
Sitzungsber. Akad. Wiss. Wien, Math.-Natwiss.
Kl. Sitzungsberichte der Akademie der Wissenschaften in Wien [Vienna], MathematischeNaturwissenschaftliche Klasse
Smithson. Misc. Coll. Smithsonian Miscellaneous Collections
Sociobiol. Sociobiology
Stettiner Entomol. Ztg. Stettiner Entomologische Zeitung
Stud. Entomol. Studia Entomologia
Stud. Geol. Salmanticensia (Stud. Palaeocheloniol.) Studia Geologica Salmanticensia
(Studia Palaeocheloniologica)
NO. 259
Syst. Entomol. Systematic Entomology

Syst. Zool. Systematic Zoology
Tr. Paleontol. Inst. Akad. Nauk SSSR Trudy
Paleontologicheskogo Instituta Akademii Nauk
SSSR
Trans. Am. Entomol. Soc. Transactions of the
American Entomological Society
Trans. Entomol. Soc. London Transactions of
the Entomological Society of London
Trans. Norfolk Norwich Nat. Soc. Transactions of the Norfolk and Norwich Naturalists
Society
Univ. California Publ. Entomol. University of
California Publications in Entomology
Univ. Colorado Stud., Ser. Bibliogr. University
of Colorado Studies, Series in Bibliography
Univ. Kansas Nat. Hist. Mus. Sci. Pap. Scientific Papers, Natural History Museum, University of Kansas
Univ. Kansas Nat. Hist. Mus. Spec.
Publ. University of Kansas Natural History
Museum Special Publication
Univ. Kansas Sci. Bull. University of Kansas
Science Bulletin
U.S. Geol. Surv. Bull., Washington United
States Geological Survey Bulletin, Washington
[D.C.]
Usp. Sovrem. Biol., Moscow Uspechi sovremennoi Biologii, Moscow
Verh. XI Intl. Kongr. Entomol., Wien Verhandlungen XI Internationaler Kongress fur Entomologie, Wien [Vienna]
Verh. Naturhist. Ver. Preuss. Rheinlande
Westphalen Verhandlungen des Naturhistorischen Vereines der Preussischen Rheinlande
und Westphalen
Wiener Ber. Natwiss. Kunst Wiener berichte
uber Naturwissenschaft in der Kunst
Zool. Inst., Acad. Sci. Zoological Institute,
Academy of Sciences
Zool. J. Linn. Soc. Zoological Journal of the
Linnean Society
Zool. Scr. Zoologica Scripta
Z. Syst. Hym. Dipt. Zeitschrift fur systematische Hymenopterologie und Dipterologie
2001
INDEX OF TAXONOMIC NAMES

Page of original reference in Roman typeface; figures
in italics. Specific names are cited individually, with current generic combination indicated in parentheses.
Names in the family-group are in capitals.
Andrena, 138
andreniformis (Thaumastobombus), 109,110114,190
antiquus (Electrolictus), 38,39,40,41,186
APIDAE, 76
APIFORMES, 35
APINAE, 86
Apis, 94,98,124,131
APOIDEA, 34
apoides (Protobombus; 5indecisus), 101
baltica (Boreallodape), 79,7880,189
basilaris (Protobombus), 105,106
Bombus, 138
Bombusoides, 138
bombusoides (Electrapis; 5tornquisti), 96
Boreallodape, 78
BOREALLODAPINI, 77
carbonarius (Bombus), 138
Chalcobombus (5Protobombus), 100
clypeatus (Melikertes), 128,192
cockerelli (Ctenoplectrella), 71,72
cockerelli (Electrapis; 5tornquisti), 96
Ctenoplectrella, 68
CTENOPLECTRELLINA, 54
Dasypoda, 64
densopunctata (Glyptapis), 62,63,188
dentata (Ctenoplectrella; 5viridiceps), 69
disareolata (Glyptapis), 59,6062,187
ditomeus (Protolithurgus), 51,52,187
Eckfeldapis (5Electrapis), 92
ELECTRAPINI, 90
Electrapis, 92
ELECTROBOMBINI, 88
Electrobombus, 88
Electrolictus, 38
eocenica (Kelneriapis), 133,134,192
EOMACROPIDINI, 46
Eomacropis, 46
fatalis (Protobombus), 103,104
fuscula (Glyptapis), 62,64,65,66,188
genalis (Glaesosmia), 75,75
glaesaria (Eomacropis), 47,47,48,186
Glaesosmia, 75
GLYPTAPINA, 54
Glyptapis, 56
goeleti (Succinapis), 115,116118,192
grimaldii (Ctenoplectrella), 73,74,188
HALICTIDAE, 36
HALICTINAE, 36
HALICTINI, 37
hermenaui (Electrapis; 5tornquisti), 96
hirsutus (Protobombus), 106,108,190

humilis (Protobombus; 5indecisus), 101
Hypotrigona, 134
indecisus (Protobombus), 100,101,102,191
Kelneriapis, 134
Kelnermelia (5Kelneriapis), 134
krishnorum (Electrapis), 94,94,96
Liotrigonopsis, 135
LITHURGINAE, 50
MACROPIDINAE, 46
martialis (Electrapis), 92,93
Megachile, 75
MEGACHILIDAE, 49
MEGACHILINAE, 53
Melikertes, 123
MELIKERTINI, 112
MELIPONINI, 133
meliponoides (Electrapis), 98,98,191
Melissites, 129
MELITTIDAE, 44
MELITURGULINI, 177,179
mengei (Bombusoides), 139
micheneri (Succinapis), 118,120,192
minuta (Roussyana; 5palmnickenensis), 132
mirabilis (Glyptapis), 2,55,57,58,187
mollyae (Boreallodape), 81,81,82,189,190
muscorum (Bombus), 138
neglecta (Glyptapis; 5fuscula), 64
nigripennis (Paleomelitta), 42,4244,186,187
OSMIINA, 54
OSMIINI, 54
Paleomelitta, 42
PALEOMELITTIDAE, 41
palmnickenensis (Roussyana), 131
PENAPINI, 37
proavus (Melikertes), 124
proboscidea (Succinapis), 121,122,192
Protobombus, 99
PROTOLITHURGINI, 51
Protolithurgus, 51
pusillus (Bombus), 138
REDIVIVINI, 45
reducta (Glyptapis; 5fuscula), 64
reticulata (Glyptapis; 5fuscula), 64
Roussyana, 131
rozeni (Liotrigonopsis), 136,136,137,192
samlandensis (Electrobombus), 88,89,90,190
Sophrobombus (5Protobombus), 100
splendens (Ctenoplectrella; 5viridiceps), 69
stilbonotus (Melikertes), 126,125127
striebichi (Boreallodape), 82,8386,189
Succinapis, 115
Tetragonula, 134
Thaumastobombus, 109
tornquisti (Electrapis), 96
Trigona, 124,132,134
trigona (Melissites), 129,129131,191
tristellus (Protobombus), 107
TRYPETINA, 54
viridiceps (Ctenoplectrella), 68,69,70,188
wrisleyi (Andrena), 138
XYLOCOPINAE, 77
185
186
NO. 259
Plate 1. Families Halictidae, Paleomelittidae, and Melittidae. a. Electrolictus antiquus Engel, holotype female (MB.I.1952 ZMHB), right lateral view. b. E. antiquus, holotype female, left lateral view.
c. Paleomelitta nigripennis Engel, holotype female (B-JH 101 AMNH), frontodorsal oblique view. d.
P. nigripennis, holotype female, dorsal view. e. Eomacropis glaesaria Engel, holotype female (SAMH),
frontal view of head. f. E. glaesaria, holotype female, left lateral view with arrow indicating staphyliniform larva on dorsum.
2001
187
Plate 2. Families Paleomelittidae and Megachilidae (Lithurginae and Glyptapina). a. Amber block
with inclusions of Paleomelitta nigripennis Engel (B-JH 101 AMNH), position of holotype indicated
by arrow (visible dimensions of block: width 5.5 cm, length 13.5 cm). b. Protolithurgus ditomeus Engel,
holotype female (B-W 157 AMNH), dorsal view. c. P. ditomeus, holotype female, right lateral view. d.
P. ditomeus, holotype female, frontal view of head. e. Glyptapis mirabilis Cockerell, holotype female
(K72 IMGP), dorsal view. f. G. disareolata Engel, holotype female (B-JH 104 AMNH), ventral view.
188
NO. 259
Plate 3. Family Megachilidae: Glyptapina and Ctenoplectrellina. a. Glyptapis fuscula Cockerell,

female (ZMUC), frontal view of face. b. G. densopunctata Engel, holotype female (B-W 161 AMNH),
left ventrolateral oblique view. c. Ctenoplectrella viridiceps Cockerell, male (NB.I.1949 ZMHB), left
lateral view. d. C. viridiceps, male, ventral view. e. C. viridiceps, neotype female (B-JH 95 AMNH),
right frontolateral oblique view. f. C. grimaldii Engel, holotype female (B-JH 88 AMNH), right lateral
view.
2001
189
Plate 4. Family Apidae: Xylocopinae. a. Boreallodape baltica Engel, holotype female (B-JH 152
AMNH), dorsal view. b. B. baltica, holotype female, ventral view. c. B. striebichi Engel, holotype
female (B-BS 153 AMNH), dorsal view. d. B. striebichi, holotype female, ventral view. e. B. mollyae
Engel, holotype female (B-JH 81 AMNH), right lateral view. f. Amber block with inclusions of B.
striebichi, holotype female is the uppermost specimen to the left of a flower fragment and highlighted
by light source (visible dimensions of block: maximal width 2.1 cm, length 3.7 cm).
190
NO. 259
Plate 5. Family Apidae: Xylocopinae, Electrobombini, and Electrapini. a. Boreallodape mollyae

Engel, paratype female (19948 ZMPA), dorsal view. b. B. baltica Engel, holotype female (B-JH 152
AMNH), left lateral view. c. Electrobombus samlandensis Engel, holotype female (B-JH 94 AMNH),
dorsal view. d. Protobombus hirsutus (Cockerell), neotype female (B-JH 92 AMNH), frontolateral
oblique view. e. E. samlandensis, holotype female, frontal view of head. f. Thaumastobombus andreniformis Engel, holotype female (B-JH 164 AMNH), left lateral view.
2001
191
Plate 6. Family Apidae: Electrapini and Melikertini. a. Protobombus indecisus Cockerell, neotype
female (B-JH 98 AMNH), right lateral view; neotype is individual in left of photograph, the head of a
second specimen is visible along the lower margin near the hind wing apex of the neotype. b. P.
indecisus, neotype female, frontal view of head. c. Electrapis meliponoides (Buttel-Reepen), neotype
female (B-JH 97 AMNH), dorsal view. d. Melissites trigona Engel, holotype female (B-JH 102 AMNH),
left lateral view.
192
NO. 259
Plate 7. Family Apidae: Melikertini and Meliponini. a. Melikertes clypeatus Engel, holotype female
(B-JH 77 AMNH), right lateral view. b. Succinapis proboscidea Engel, paratype female (B-JH 99
AMNH), left lateral view. c. S. micheneri Engel, holotype female (B-JH 103 AMNH), right lateral view
(arrow indicates clypeal protusion). d. S. goeleti Engel, holotype female (B-JH 90 AMNH), right lateral
view. e. Kelneriapis eocenica (Kelner-Pillault), holotype female (NB.I.1946 ZMHB), left lateral view.
f. Liotrigonopsis rozeni Engel, holotype female (B-JH 79 AMNH), right lateral view.

Evolution of The Apoidea

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A MONOGRAPH OF

THE BALTIC AMBER BEES

BULLETIN OF THE AMERICAN MUSEUM OF NATURAL HISTORY

Copyright q American Museum of Natural History 2001

Photomicrograph of the holotype (IMGP-K72) of Glyptapis mirabilis Cockerell.

BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY

Genus Melissites, new genus . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 129

ENGEL: BALTIC AMBER BEES

BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY

ENGEL: BALTIC AMBER BEES

Et latet, et lucet Phaetontide condita gutta,

Bees occupy a unique role in the world of

flowers teeming with the gentle murmur of

1 Translation: Now dim, now bright, trapped in its

2 Go to the bee, and learn how diligent she is, and

MONOGRAPHIA APUM SUCCINI BALTICI

BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY

(e.g., Moldenke, 1976; Arroyo et al., 1982;

ENGEL: BALTIC AMBER BEES

Bee Fauna of Miocene Dominican Amber

BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY

diligent efforts over the past three years I

Hierarchical Supraspecific Classification of

ENGEL: BALTIC AMBER BEES

fall nicely into extant genera or into genera

but the living fauna was considered on a

BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY

Comparison of Present Classification with Earlier Usage

ENGEL: BALTIC AMBER BEES

presently stands; more detailed studies and

diligent search it appears that only one of his

BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY

gen as indicated in Kelner-Pillaults paper;

ENGEL: BALTIC AMBER BEES

Although pockets of Baltic amber can be

BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY

cinic acid (e.g., glessite, gedanite), or possess

port. Among the thousands of inclusions in

ENGEL: BALTIC AMBER BEES

the United States (i.e., North Carolina to

It is not of minor significance that Hope

MATERIALS AND METHODS

BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY

burg, Germany; CHFG, Collection of C. and

When possible, specimens were vacuumembedded in epoxy following the procedure

In presenting taxonomic summaries for

ENGEL: BALTIC AMBER BEES

ber of specimens (immediately following the

future study (satisfying ICZN, 1999b: Art.

BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY

area (fig. 1) and is slightly set below the

ENGEL: BALTIC AMBER BEES

along the inner margin of the compound eye

ciput bears a large opening, the foramen

BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY

atomical details concerning mandibular

paired maxillae and the central labium. The

ENGEL: BALTIC AMBER BEES

glossa (fig. 2). As is discussed briefly below