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 A GUIDE TO

NATIVE BEES
OF AUSTR ALIA

TERRY HOUSTON
© Terry Houston 2018
All rights reserved. Except under the conditions described in the Australian Copyright Act 1968 and
subsequent amendments, no part of this publication may be reproduced, stored in a retrieval system or
transmitted in any form or by any means, electronic, mechanical, photocopying, recording, duplicating
or otherwise, without the prior permission of the copyright owner. Contact CSIRO Publishing for all
permission requests.
A catalogue record for this book is available from the National Library of Australia.
Published by
CSIRO Publishing
Locked Bag 10
Clayton South VIC 3169
Australia
Telephone: +61 3 9545 8400
Email: publishing.sales@csiro.au
Website: www.publish.csiro.au
Front cover: (main image) Blue-banded bee, Amegilla chlorocyanea, on Verticordia flowers, Jiri
Lochman; (top, left to right) female of Dawson’s Bee, Amegilla dawsoni, on flower of Rough Blue-bell,
Janine Guenther; males of Mellitidia tomentifera on their night-time roost, Alan Henderson; female of
Palaeorhiza disrupta, Andrew MacDougall.
Back cover: (left to right) Female of Hyleoides zonalis foraging on eucalypt flower, Kerry Stuart;
Amegilla sp. clearly grasping stem by mandibles alone, Jean and Fred Hort; Leioproctus conospermi with
proboscis inserted into smoke-bush flower, Kerry Stuart.
Title page: Two male carpenter bees at the entrance to their nest, Donna Sanders.
Photographs are by the author unless otherwise noted.
Set in 9.5/12.5 Adobe Minion Pro and Myriad Pro
Edited by Joy Window (Living Language)
Cover design by James Kelly
Typeset by Desktop Concepts Pty Ltd, Melbourne
Printed in China by Toppan Leefung Printing Limited
CSIRO Publishing publishes and distributes scientific, technical and health science books, magazines
and journals from Australia to a worldwide audience and conducts these activities autonomously from
the research activities of the Commonwealth Scientific and Industrial Research Organisation (CSIRO).
The views expressed in this publication are those of the author(s) and do not necessarily represent those
of, and should not be attributed to, the publisher or CSIRO. The copyright owner shall not be liable for
technical or other errors or omissions contained herein. The reader/user accepts all risks and
responsibility for losses, damages, costs and other consequences resulting directly or indirectly from
using this information.
Original print edition:
The paper this book is printed on is in accordance
with the standards of the Forest Stewardship
Council ®. The FSC® promotes environmentally
responsible, socially beneficial and economically
viable management of the world’s forests.
Contents

Preface v
Acknowledgements vii

PART I OVERVIEW OF BEES AND THEIR BIOLOGY 1

What is a bee? 2
Form and function 3
Origin and evolution of bees 17
Australian bee fauna 19
Importance of native bees 21
Bee life-cycle 22
Sexing bees 26
About males and mating 27
Stings 35
Bees’ glandular products 36
Size range 39
Colour patterns, mimicry and crypsis 40
Sociality 46
Nests and nesting behaviour 47
Cuckoo bees 57
Flower visitation and feeding 59
Seasonality and flight times 69
Associated organisms 71
Conservation of bees 76
Historical account 79
Collecting and preserving bees 81
Encouraging native bees in the garden 84

iii
A G U I D E T O N AT I V E B E E S O F A U S T R A L I A

PART II IDENTIFICATION OF BEES 87

Identifying bees 88
Is it a bee? 89
Native bee or honeybee? 91
Regarding names: scientific versus common 92
Identification of Australian bees to family 93
Family Colletidae 96
Family Stenotritidae 168
Family Halictidae 174
Family Megachilidae 197
Family Apidae 214
Bees introduced to Australia 246

Glossary 249
Further reading 252
Bibliography 253
Index 265

Photo: Kerry Stuart.

iv
Preface
The natural world in all its various facets
can be a source of interest and wonder-
ment for many of us. Certainly that was
the case for me from my earliest years. I
grew up in suburban Adelaide at a time
when there were still paddocks, swamps,
coastal dunes and open drains and when
children were allowed to roam and play
freely, as long as they turned up for the
next meal-time. My father was a keen gar-
dener who grew fruit trees, grape vines,
vegetables and a variety of flowers. He
maintained a very neat garden with rock-
edged beds with gravel paths between and
no weed ever lasted more than a day or two
before getting the flick from his pocket-
knife. In this orderly environment there
was still some of ‘the wild’ – a variety of
native birds, reptiles, spiders and insects –
and I spent hours watching them to see
what they were doing and, as I grew older,
seeking out their correct names.
Honeybees were ever present because of
the abundant flowers my father grew. The
hum of their wings among the poppy flow-
ers was one of the most delicious sounds of
summer days. But it was probably my
mother’s little herb garden that first
opened my eyes to the existence of native
bees. When some of her parsley plants pro-
duced heads of tiny white flowers, I hap-
pened to notice one day how they were
visited periodically by tiny black insects
with an iridescent green sheen that rum-
maged around in the manner of honey-
bees. When, on close inspection, I saw that
these insects were packing loads of white
pollen beneath their abdomens, it became
clear that they really were bees, just very
tiny ones. I didn’t know it then but I had
just met Homalictus urbanus.
Subsequently, I became aware of
another kind of native bee in our garden.
This bee was almost as large as the honey-
bee but had a more rounded body and
strong black and white bands on its abdo-
men, and its flight was different. The
females worked frenetically, dashing from
flower to flower and, intermittently, they
would stop to hover while they had a quick
clean up. In those brief moments, I noticed
with amusement how their bodies waggled
vigorously. This species, I learned much
later, was a blue-banded bee, Amegilla
chlorocyanea. Where did they come from
and where were they taking their loads of
pollen, I wondered. The seeds of an abid-
ing interest in native bees began to
germinate.
As an undergraduate student at the
University of Adelaide, I began to consider
seriously what kind of career I wanted.
Entomology, the study of insects, had a
strong appeal and seemed to offer good
prospects for employment. By this time, I
had learned that there were hundreds of
species of native bees in Australia and that
they were poorly studied. I was excited by
the thought that perhaps I could rectify
this situation, at least to some extent. My
v
A G U I D E T O N AT I V E B E E S O F A U S T R A L I A
desired course was set from that time on,
and my interest in native bees has persisted
to this day, more than 50 years later. The
reader might think that I could have
learned all that there is to know about Aus-
tralia’s native bees in far fewer years and
that I might have tired of them by now. Not
so! The diversity of our bees (an estimated
2000 species with extremely varied form
and behaviour) has yielded surprise after
surprise. It is still the case that the life-
cycles and behaviour of the majority of
species have not been subject to any in-
depth study, so more surprises may be in
store for students of the bees. It is all too
easy to assume that because some mem-
bers of a group behave in a certain way, all
members do likewise. Successive and com-
paratively recent discoveries of novel forms
vi
and behaviours among our bees should
encourage further study of our bee fauna.
The first part of this guide provides a
general introduction to the bees and should
give the reader at least a basic grasp of their
morphology, evolution, behaviour and
ecology. The second part is intended to
enable the reader to identify any Australian
bee to at least genus. As well, it features
some of the most common species and
some of the most unusual or remarkable
species, which may be recognisable from
the illustrations. For those desiring to go
further with bee identification and study,
there will be found within the pages of this
work references to sources of more detailed
information. Above all, I hope this work
will stimulate interest in and appreciation
of Australia’s wonderful native bee fauna.
Acknowledgements
Photography of live bees in nature usually
requires considerable time, patience and
skill and, for the many fine bee portraits
that grace the pages of this book, I am
deeply indebted to the following contribu-
tors: Mark Berkery, James Dorey, Bryony
Fremlin, Janine Guenther, Alan Hender-
son, Jean and Fred Hort, Bernhard Jacobi,
Tony Kirkby, Remko Leijs, Andrea Lim,
Jiri and Marie Lochman, Andrew Mac-
Dougall, Marc Newman, David Pike,
David Rentz, Linda Rogan, Laurence and
Donna Sanders, Tobias Smith, Kerry
Stuart, Malcolm Tattersall and Jenny
Thynne. Michael Batley and Ken Walker
kindly provided images of museum speci-
mens. For technical assistance with image
handling and manipulation, I thank Evan
Rogers and Robert Fleming.
Preparation of this guide has been a
long and complex task and, throughout the
process, I have very much appreciated the
guidance, patience and encouragement
provided by CSIRO Publishing editors
Briana Melideo and Lauren Webb. They
helped get me over the finish line. I could
not have produced this book, either, with-
out the support of the Western Australian
Museum, its staff and facilities.
The system of classification of Aus-
tralian native bees has been developed
over centuries by various specialists and
no one has done more to provide a stable
classification of Australia’s and the
world’s bees than the late Professor
Charles Michener of Kansas University.
The scheme used in this book is based
largely on that outlined in his monumen-
tal work, The Bees of the World (Michener
2000, 2007). Furthermore, my identifica-
tion keys are based on his with some
modifications.
vii
This page intentionally left blank
 Part I

Overview of bees and their biology

1
What is a bee?
For those people who have no knowledge
of bees other than the European honeybee,
Apis mellifera, this book should be a revela-
tion. The usual concept of bees as medium-
sized, hairy, highly social insects that live
in hives and produce honey will be chal-
lenged. Australia’s native bees are
extremely diverse in size, form, coloura-
tion and behaviour. Certainly, some of
them look rather like honeybees, but many
others are very wasp-like in appearance.
Add to that the fact that some wasps (and
even some flies and beetles) appear very
bee-like and you can begin to appreciate
the need for a definition.
Bees, in essence, are wasps. They are
an offshoot of the wasp clan and have
become specialised for a diet of pollen
and honey in both adult and larval stages.
Their closest relatives are hunting wasps
The pollen load on this insect clearly identifies it as a female bee, rather than a wasp.
Lasioglossum lanarium (Halictidae). Photo: Jenny Thynne.
2
that capture insects and spiders, sting
them into paralysis and store them as
food for their larvae in specially made
brood cells (see more in Origin and evolu-
tion of bees). Bees are structurally very
similar to their wasp relatives and it is
chiefly their behaviour that sets them
apart. They have become vegetarians and
the change-over involved relatively minor
structural adaptations. The sting has been
retained in most bees but is now used
only for defence. Body hairs (setae)
became branched, perhaps as an adapta-
tion for holding pollen grains, and
females of most bees have dense sets of
hairs (called scopae) for carrying loads of
pollen. Bees are closely associated with
flowers, but so too are many hunting
wasps that depend on nectar (but not
pollen) for their energy requirements.
Form and function
The classification of bees into groups such
as families and genera is largely based on
characters of external and internal anat-
omy. These characters frequently involve
structures that are highly functional and
many of them can be considered part of a
bee’s tool-kit. They can be involved in
sensing, locomotion, feeding, grooming,
nest-building and mating. This section
outlines the external anatomical struc-
tures of bees and their functions, with
emphasis on those of most importance to
making identifications.
Head
The head (Figs 1, 2) connects to the thorax
by a flexible neck so that it can swivel in
most directions. On each side are the com-
pound eyes formed of thousands of tiny
fixed lenses (or facets). In combination
with the nervous system, the compound
eyes provide bees with a visual system that
evidently functions extremely well. The
eyes are bare in most bees but, in some,
they are quite hairy, the hairs being
inserted between facets. Situated on or
close to the top of the head, the vertex, are
three simple eyes, the ocelli. They are
believed to help the bee orientate itself in
relation to the horizon during flight. Ocelli
are unusually large in bees that habitually
fly at night or in dim light.
The face between the ocelli and the
antennal sockets (the frons) often has a
pair of broad depressions, deep grooves or
pits either side. These are the facial foveae
and mark the openings of wax glands. Sev-
eral sutures divide the lower face into sepa-
rate areas. The clypeus is fairly level to
gently convex in most bees but is variously
modified in others. Females of many meg-
achilids, for example, have the clypeus
excavated, protuberant or developed into
stout tubercles – modifications which
assist them to carry building materials.
The areas immediately behind the eyes,
the genae, occasionally also bear spines
assisting the transport of materials. The
concavity in the back of the head, the
occiput, accommodates the anterior end of
the thorax. The occipital surface may
round onto the adjoining vertex and genae
or be sharply delineated from them by a
crescentic rim, the preoccipital carina. A
large median cavity in the lower posterior
part of the head capsule, the proboscidial
fossa, accommodates the proboscis when
the latter retracts and folds.
Antennae
Often referred to as feelers, the antennae of
bees (Fig. 1) serve a tactile function but,
more importantly, they provide bees with
their sense of smell. Each comprises several
segments (12 in females, 13 in males). The
first and usually longest segment is the
scape, its base forming a ball and socket
joint in the face, so the antennae are quite
mobile. The second segment is the relatively
small, rounded pedicel and the remaining
10 or 11 segments form the flagellum. Hun-
dreds of microscopic sensory organs
3
A G U I D E T O N AT I V E B E E S O F A U S T R A L I A

Fig. 1. Terminology of parts of the head and its appendages (based on Callohesma megachlora).

­ istributed over the flagellum detect floral

d fied in males of certain bee species.
fragrances, pheromones, and other odours. Megachile (Thaumatosoma) species, for
Antennae, though generally quite uni- example, have antennae somewhat like
form among females, are variously modi- those of butterflies – extremely slender,

a b

Fig. 2. Terminology of posterior parts of the head (based on Hylaeus females). (a) Head lacking a
preoccipital carina (H. chrysaspis; occiput rounds onto vertex and genae); (b) head with
preoccipital carina (p.o. carina; H. globuliferus).

4

almost as long as the head and body, and
with the apical two segments expanded
(see p. 212). Capitate or spatulate antennae
crop up in several distantly related groups
(e.g. Euhesma semaphore (p. 33) and
Trichocolletes dives (p. 119). How the
expansions serve the males is unknown.
The most elaborate male antennal flagella
are those of Leioproctus (Cladocerapis)
bipectinatus (p. 107), each segment pos-
sessing antler-like branches. Leioproctus
(L.) macmillani (p. 107) also has elaborately
modified flagella, each segment being
bipectinate. Scapes, rather than flagella,
are modified in males of some other bees.
In several cases, the scapes are greatly
broadened, though rather flat (e.g. species
of Neopasiphae (p. 112) and Hylaeus
(Xenohylaeus) (p. 158) while in others they
are globose, reaching an extreme form in
the two species of Hylaeus (Sphaerhylaeus)
(p. 155). These enlarged scapes accommo-
date glands that release their secretions
(probably pheromones) via pores on the
rear of the scapes. In Sphaerhylaeus, males
have hair-filled facial cavities behind the
scapes that evidently receive the secretion.
Labrum
Set on the lower margin of the face between
the mandibles, this usually small, hinged
flap folds back to protect the end of the
proboscis when it is retracted (Fig. 1). The
shape and size of the labrum and its vari-
ous embellishments provide useful charac-
ters for distinguishing different kinds of
bees.
Mandibles
Commonly referred to as nippers or jaws,
the mandibles are a pair of simple, hard-
ened appendages either side of the mouth
F o r m a n d f u n c tio n
(Fig. 1). They are employed in a wide vari-
ety of tasks such as loosening soil (burrow-
ing bees), boring into pith or wood
(carpenter bees), tearing away detritus
(lodger bees), cutting leaf pieces or gather-
ing leaf pulp, resin, plant hairs and other
materials (megachilids), snipping open
flower buds and grasping opponents
during fighting. Their form varies consid-
erably but mandibles commonly bear two
or more teeth at their apices.
Proboscis
This is a complex feeding organ that func-
tions primarily to enable bees to suck
nectar from flowers and, in females, may
function also in applying secretions during
nest construction (Figs 3–5). Being a
jointed organ, it can be extended or
retracted and, when folded into a recess in
the back of the head (the proboscidial
fossa), it may scarcely be visible. The pro-
boscis has evolved from two pairs of jointed
appendages possessed by the insect ances-
tor. Known as maxillae (singular, maxilla),
these appendages are found in most insects,
though the second (posterior) pair have
partially fused and are known as the
labium. Each original maxilla possessed a
jointed appendage known as a palpus or
palp and these persist in most insects. In
those insects that feed by chewing, the
maxillae and their palps assist in manipu-
lating food into the mouth. In bees, there is
a pair of maxillary palpi (primitively six-
segmented) and a pair of labial palpi
(primitively four-segmented) but they
serve an essentially sensory role.
One of the most important parts of the
proboscis is the tongue (or glossa): a hairy,
flexible extension of the labium and one
that takes a variety of forms among the
5
A G U I D E T O N AT I V E B E E S O F A U S T R A L I A

Fig. 3. Terminology of proboscis of a long-tongued bee (based on Megachile aurifrons).

(a) Whole proboscis, lateral view; (b) same, dorsal view; (c) labium, ventral view.

bees. Its primary function is to mop up long-tongued bees) or move it from side to
nectar from the nectaries of flowers, and side. The duct of a bee’s salivary glands
muscles within the labial prementum can opens at the base of its glossa, and the
retract the tongue (at least partially in glossa may serve as a brush to apply the

6
 F o r m a n d f u n c tio n

a b
Fig. 4. Terminology of proboscis of a short-tongued bee (based on Lasioglossum mirandum,
female, Halictidae). (a) Whole proboscis, ventral view; (b) maxilla lateral view (cardo removed;
broken line divides galea into relatively long pre-palpal portion (b) and much shorter post-palpal
portion (a)).

salivary secretion during nest construc-
tion, especially in the family Colletidae.
Thorax
All of a bee’s locomotory equipment is
located on and in the middle section of the
body, the thorax (Fig. 6). Three pairs of
legs and two pairs of wings attach to it.
Derived from three body segments of the
insect ancestor, each carrying one pair of
legs, this section is divisible into the pro-
thorax, mesothorax and metathorax. The
prothorax, carrying the fore legs, may be
quite inconspicuous, although sometimes
the upper part is strongly developed into a
pronotal collar between the head and
meso­thorax. The rounded pronotal lobes
wrap around onto the sides of the meso-
thorax. Containing the muscles that power
the fore wings, the mesothorax is the larg-
est thoracic segment. Dorsally, it is divided
into four shields: the large scutum, the
scutellum and, on each side of the latter,
the axillae. The sides of the mesothorax
are divided by two sutures: the horizontal
scrobal suture and the vertical episternal
suture (strongly reduced in certain
groups). Between the bases of the hind
wings on the metathorax sits the dorsal
plate, the metanotum. During the evolu-
tion of the wasps from sawfly-like ances-
tors, a constriction developed between the
first and second segments of the abdomen,
the first segment becoming closely associ-
ated with the metathorax and being known
as the propodeum. The propodeum is
developed to greater or lesser degrees in
different groups of bees but always has a
demarcated area, the propodeal enclosure
(sometimes referred to as the propodeal
triangle). This enclosure is broad dorsally
just behind the metanotum and tapers
posteroventrally to the insertion of the
abdomen. Usually its sculpture differs
from that of adjacent areas.
Wings
Flight becomes more efficient with just
one pair of wings and, in bees (and their
wasp relatives), the fore and hind wing of

7
A G U I D E T O N AT I V E B E E S O F A U S T R A L I A

a b

c
Fig. 5. Terminology of proboscis of a short-tongued bee (based on Leioproctus plumosus,
female, Colletidae). (a) Head in lateral view to show proboscis extended; (b) proboscis in situ,
posterior view; (c) maxilla lateral view (broken line divides galea into pre-palpal portion (b) and
relatively longer post-palpal portion (a)).

each side are coupled: a row of micro- so that, in flight, the two wings act as one
scopic hooks (hamuli) on the leading edge (Fig. 7).
of the hind wing engages with a raised Stiffening the wing membranes is a net-
vein on the hind margin of the fore wing, work of veins. They form a pattern that is so

8
 F o r m a n d f u n c tio n

Fig. 6. Terminology of parts of the mid-body or mesosoma (thorax + propodeum) (based on

Callohesma megachlora). Wings have been removed in (b).

constant that each vein and each ‘cell’ developed but occasionally may be virtually
formed by them can be named. A thicken- unrecognisable. Variation occurs in the rel-
ing in the leading edge of the fore wing, the ative sizes and shapes of the stigma, veins
stigma (or pterostigma), is usually well and cells, and some veins may be weaker

9
A G U I D E T O N AT I V E B E E S O F A U S T R A L I A

c
Fig. 7. Terminology of wings. (a, b) Fore wing; based on Lipotriches australica: (a) veins (numbers
identify first, second and third submarginal cross-veins; (b) cells (numbers identify first, second
and third submarginal cells). (c) Hind wing (based on Paracolletes (Anthoglossa) sp.).

than others, or lost altogether (particularly After alighting, a bee typically retracts
in the smallest bees). These differences can its wings so that they overlap above the
provide useful taxonomic characters. abdomen. Sometimes, however, certain

10

Fig. 8. Terminology of parts of the leg (based on hind leg of Megachile aurifrons, female; note
absence of scopal hairs).
kinds of bees maintain the wings out in a
V-shape after alighting on flowers. This is
true particularly of those bees that mimic
mud-nest wasps.
Legs
In addition to enabling bees to stand, walk
or grasp the substrate, legs (Figs 8, 9) can
assist bees to manipulate flowers, carry
pollen, scrape or push away soil, wood or
pith particles during burrowing, carry
nest-building materials to nest sites, groom
themselves and, in males, grasp mates.
The legs of bees have the same basic
form as those of other insects, consisting
of five articulated sections: coxa (attaching
to the body), trochanter, femur, tibia and
tarsus (Fig. 8). The tarsus (or foot) is five-
segmented, the first segment (basitarsus)
tending to be the longest. The last segment
F o r m a n d f u n c tio n
(distitarsus) bears a pair of claws and, in
the majority of bees, a retractable adhesive
pad, the arolium (Fig. 9c; lost in some
groups of bees).
The tibiae typically have one or, in the
case of the hind leg, two articulated apical
spurs. These play a role in grooming, that
of the fore tibia forming part of the
antenna cleaner, the spurs of the mid and
hind tibiae often being comb-like and
assisting the removal of pollen from the
legs and body.
At the very base of the hind tibia in
many bees (mainly the ground-dwelling
kinds) is a specialised area, the basitibial
plate (Fig. 9d, e). Often defined by raised
edges (carinae) and sometimes with short,
specialised hairs, these plates serve as knee
pads that are pressed against the walls of
the nest burrow as the bee moves along it.
11
A G U I D E T O N AT I V E B E E S O F A U S T R A L I A

d c

e f
Fig. 9. Terminology of some hind leg structures. (a–c) Lasioglossum mirandum, female: (a) whole
leg, outer view; (b) hind tarsus to show penicillus (arrowed); (c) apex of tarsus in ventral view to
show arolium (arrowed; note that arolia usually occur on all legs of both sexes); (d, e) Ctenocolletes
ordensis, female: (d) outer view of whole leg (except for coxa); (e) dorsal view of apex of femur and
base of tibia; (f) Tetragonula worker: outer view of hind leg to show location of corbicula (arrowed).

In many groups of bees, pollen loads be limited to the basal section of the leg
are transported on special sets of hairs on (Fig. 9a) or to the outside of the tibia, often
the hind legs of females. Known as the extending as well onto the basitarsus (Fig.
scopa, the set of pollen-carrying hairs may 9d). Scopal hairs are simple in some bees

12

while in others they are forked or plumose.
The density of the scopa hairs bears an
inverse relationship to the size of the pollen
grains that are normally carried.
Certain members of the family Apidae
lack scopal hairs and instead carry their
pollen loads on corbiculae (also referred to
as pollen baskets): these are the largely
bare, rather flat outer surfaces of the hind
tibiae that are fringed with erect hairs or
bear only scattered erect hairs (Fig. 9f).
A small brush of hairs (penicillus)
occurs on the apical end of the basitarsus
of females of most Halictidae (Fig. 9a, b)
and some Apidae (notably Amegilla).
Legs exhibit a great array of modifica-
tions, especially in males, and these may
assist in grasping females during mating.
In Lipotriches species and various other
bees, the hind femora are strongly swollen
Fig. 10. Terminology of some abdominal structures (based on Lipotriches australica, female,
lateral view; abdomen partially cleared and expanded). Legend: gr, gradulus; S1–S6, sternites; sp,
spiracle; sting, sting apparatus; T1–T6, tergites.
F o r m a n d f u n c tio n
and the hind tibiae appear deformed. Tarsi
(the feet) are especially prone to being
expanded and elaborated (see more under
About males and mating, p. 27).
Abdomen
As noted above under Thorax, the original
first section of the insect abdomen forms
the propodeum in wasps and bees (Figs 10,
11). Technically, the remaining part of the
abdomen is termed the gaster or meta­soma
but, for simplicity, the term abdomen will
be used in this work.
Externally, the abdomen is seen to con-
sist of six (female) or seven (male) seg-
ments. Each segment is covered by a pair of
large plates, a tergum (or tergite) above and
a sternum (or sternite) below, all plates
being joined by flexible, colourless interseg-
mental membranes. The terga overlap the
13
A G U I D E T O N AT I V E B E E S O F A U S T R A L I A

a b

c d e
Fig. 11. Terminology of some abdominal structures. (a) Dorsal view of apex of abdomen of
female bee (Stenotritus greavesi) showing fimbriae and basitibial plate; (b) hind part of of female
bee (Xanthesma sp.), lateral view, showing fovea of second abdominal tergum; (c–e) terminalia of
male bee (Leioproctus plumosus), dorsal view: (c) genital capsule; (d) sternum 7; (e) sternum 8.

sterna laterally and each pair overlaps the
bases of the following pair, so that the
abdomen can telescope in and out. This
telescoping can be observed when a bee
lands after a flight, the rhythmic move-
ments equating to the breathing move-
ments of mammals. Insects do not have
lungs but they do have a system of air tubes
(tracheae) that carry oxygen to the tissues
and permit escape of carbon dioxide to the
exterior. Numerous tracheae are expanded
to form air sacs, especially within the
abdomen, and the pumping movement of
the abdomen forces air through the system.
A pair of small openings (spiracles) occurs
on each body segment, permitting the
exchange of gases between the insect’s tra-
cheal system and the air.
In females of ground-nesting bees, the
last visible (sixth) tergum of the abdomen
has a special raised median plate, the
pygidial plate (Fig. 11a). Females use this
to tamp the soil, compressing it and form-
ing a smooth surface, during burrow and
cell construction. Males, too, occasionally
possess such a plate but on the seventh
tergum.
The second abdominal tergum of
females, especially in the family Colleti-
dae, often has a fovea each side. This may

14
 F o r m a n d f u n c tio n

be an impressed, dark spot or groove (Fig.

11b) or just an area of finer surface scultur-
ing, sometimes distinctively coloured.
The original insect abdomen consisted
of 10 segments. In wasps and bees, the more
apical segments have become invaginated
and form either part of the sting apparatus
in females (Fig. 10) or are associated with
the genitalia in males. Commonly, the sev-
enth and eighth sterna of males are invagi-
nated and strongly modified (Fig. 11d, e). a
Sometimes other apical sterna are modified
as well. The genitalia of males consist of a
sclerotised organ (Fig. 11c) which has two
pairs of appendages: the outer gonocoxites
and the medial penis valves.
In certain groups of bees (e.g. Homalic-
tus (Halictidae) and most Megachilidae),
the pollen-transporting hairs (scopa) are
located on the abdominal sterna. b

Hairs (setae)
What we loosely call hairs on bees and
other insects are not homologous with the
hairs of mammals and are more correctly
termed setae. A seta is a hollow outgrowth
of the cuticle, is composed of chitin and
articulates at its base in a socket. A sensory
organ at the base of each seta detects move-
ment, so hairs are important tactile organs. c
Setae develop in the pupal stage, and their Fig. 12. Branched hairs (setae) – the hallmark
size and form remain fixed for the life of of bees. (a) Long, branched hairs for holding
pollen on abdominal segment of Homalictus
an adult. They can occur on almost any dotatus (intermixed with some shorter, simple
part of the cuticle, including some parts hairs); (b) plumose hairs on abdomen of
Paracolletes (Anthoglossa) sp.; (c) very short,
that are invaginated (such as the male highly plumose hairs forming a felt-like
genitalia). In their simplest form they are covering (tomentum) on thorax of Megachile
single filaments or bristles. However, it is a (Austrochile) sp.
characteristic of bees that at least some,
and often most, setae are branched or plu- bear a few to dozens of lateral filaments
mose. In the latter case, the main stem may along its length.

15
A G U I D E T O N AT I V E B E E S O F A U S T R A L I A
Glands
Bees of all kinds depend heavily on glan-
dular products in living their lives. Some
of these products are discussed below
under Bees’ glandular products. Here, I
shall mention only those known as epider-
mal glands, so named because they open to
the surface of the ‘skin’ and have chitinous
ductules or sacs. The gland openings are
often quite visible and can be useful taxo-
nomically. In other cases, they are micro-
scopic and scattered or concealed.
Mandibular glands
A gland and gland sac lie on each side of
the head above each mandible and secre-
tion is discharged via a pore in the soft
membrane between the inner base of each
mandible and the head capsule.
Antennal glands
Males of several bees in the colletid sub-
family Hylaeinae have swollen antennal
scapes that accommodate glands and
gland sacs opening onto the posterior sur-
faces (Houston 1975a). The most highly
developed antennal glands are those of the
two species of Hylaeus (Sphaerhylaeus)
(see p. 153). Each scape contains a reservoir
to hold the gland secretion which is dis-
16
charged via a single pore on the posterior
surface. Lying in the frons behind each
scape is a large, hair-filled cavity which
evidently absorbs the secretion.
Facial and abdominal foveae
These specialised areas of cuticle (see
under Head and Abdomen above) mark the
locations of wax glands lying on the inner
side of the integument. The waxy secretion
is probably groomed over the head and
abdomen with the tarsi.
Abdominal glands
Large glands commonly occur in the abdo-
men and open by numerous microscopic
pores in the intersegmental membrane or in
the terga themselves. These glands tend to
be more common in males, so their secre-
tions may be involved in mate attraction. In
certain species of Hylaeus and Leioproctus,
the hidden basal portions of the abdominal
terga are distinctly different from the
exposed apical portions in lacking pigment
and being covered with dense, short hairs.
Hundreds of gland pores occur among the
hairs, the latter probably serving to main-
tain an air space between the overlapping
abdominal plates so that the gland secretion
is able to evaporate and disperse.
Origin and evolution of bees
It has long been understood that bees are
an offshoot of the hunting wasps formerly
placed in the family Sphecidae and now
placed in the family Crabronidae (Melo
1999; Debevec et al. 2012). Bees and cra-
bronid wasps share a very similar body
plan and there are strong similarities in
nesting habits between at least some mem-
bers of the two groups. How the transfor-
mation occurred from predatory habits to
pollen and nectar collecting has never
been fully explained and it is difficult to
imagine intermediate stages in that trans-
formation. Yet a similar transformation
occurred in a second family of wasps – the
Vespidae – leading to the pollen wasps
(Masarinae). Like bees, masarine wasps
collect and store pollen and nectar as food
for their larvae but, unlike bees, they have
no branched hairs or scopae. Their females
swallow pollen along with nectar and
transport it to their nests in their crops (or
honey stomachs). This is known as alimen-
tary transport of pollen. Females regurgi-
tate the pollen and nectar mixture as a
paste in the brood cells.
Alimentary transport of pollen and
nectar also occurs in bees of the colletid
subfamilies Hylaeinae and Euryglossinae,
females of which lack scopae. Bees in these
two subfamilies are relatively hairless and
often possess white, cream or yellow mark-
ings so that they are very wasp-like in
appearance. Colletids generally exhibit
several wasp-like characteristics (short,
broad tongues, simple palpi, solitary habits
and most construct simple ground-nests)
and were long viewed as the most primitive
of bees. Among the colletids, the Hylaeinae
and Euryglossinae were once considered to
be closest to the ancestral bee. More recent
studies, however, have challenged that
view (see more below).
The fossil record so far has proved to be
of limited help in understanding the origin
of bees. The oldest known bee fossil is a
tiny (<3 mm) male embedded in amber
from Myanmar (Burma) that has been
dated at ~100 million years old. That puts
it in the Middle Cretaceous when dino-
saurs were at the height of their evolution.
Named Melittosphex burmensis, the fossil
forms the basis of a new family Melitto-
sphecidae. It combines both wasp-like and
bee-like features, the latter including
branched hairs (Poinar and Danforth
2006; Danforth and Poinar 2011). Because
the specimen is a male, we lack informa-
tion about whether females of the species
had a scopa and it is not possible to say that
M. burmensis collected nectar and pollen.
Previously, the oldest known bee fossil
was Cretotrigona prisca, found in Creta-
ceous amber from New Jersey dated at ~80
million years old. It is clearly a member of
the highly social, long-tongued bees in the
family Apidae, tribe Meliponini, a group
generally considered to be one of the most
highly evolved of the bees.
The Cretaceous period was notable for
the rise of another group of organisms –
the flowering plants (angiosperms). Co-
17
A G U I D E T O N AT I V E B E E S O F A U S T R A L I A
evolution is understood to have occurred
between the angiosperms and the bees,
each group enabling more rapid evolution
and diversification of the other.
Studies of the evolutionary pathways
and relationships (phylogeny) of the major
groups of bees, based on both morphology
and genetics, have resulted in numerous
hypotheses. Sadly, consensus among the
world’s bee specialists is slow in coming.
The hypotheses are too numerous and
complex to discuss in detail here but com-
prehensive reviews have been provided by
Michener (2007) and Plant and Paulus
(2016). There does now seem to be some
consensus arising out of the many phylo-
18
genetic studies: (1) the melittids, not
colletids, are considered to occupy the
most basal position in the family tree
(melittids are short-tongued, burrowing
bees in which the glossa is short and
pointed or occasionally slender); (2) all
other bees are believed to have evolved
from the melittids, forming two major lin-
eages – the short-tongued lineage and the
long-tongued lineage; (3) in the short-
tongued lineage, the Colletidae is consid-
ered to be the most derived (because of the
short, obtuse or bifid glossa) and, within
the Colletidae, the Euryglossinae and
Hylaeinae are considered derived rather
than basal.
Australian bee fauna
Australia’s native bee fauna is large and
diverse, and differs in major respects from
the bee faunae of other continents. Though
the total number of bee species inhabiting
Australia is yet to be established, it could
be as high as 2000. At the time of writing,
1546 named species were listed on the
Australian Faunal Directory (ABRS 2009).
Many unnamed species are known in col-
lections, especially in the genera Hylaeus,
Euhesma, Leioproctus and Megachile, and
it is anyone’s guess how many species
remain to be discovered.
Australia’s bee species are classified
into 58 genera in five families (using
Michener’s 2007 system). Four of the fami-
lies represented in Australia occur world-
wide, while one, the Stenotritidae, is
exclusively Australian. Two families nota-
bly absent from Australia, but otherwise
with world-wide distributions, are the
Andrenidae and Melittidae. Also, for some
of the world-wide families represented in
Australia, certain subfamilies or tribes are
conspicuously absent; these include Osmi-
ini (Megachilidae), Augochlorini (Halicti-
dae), the honeybee genus Apis and the
bumblebee genus Bombus. The tribe
Anthidiini (Megachilidae) is barely repre-
sented in Australia by two putative species
in north-eastern Queensland. Compared
with other continents, too, Australia has
relatively few cuckoo bees, with represent-
atives of just four genera.
The most notable feature of the Aus-
tralian bee fauna is the predominance of
the family Colletidae which accounts for
55% of the species and 60% of the generic-
level names (i.e. genera and subgenera).
One of its subfamilies, the exclusively Aus-
tralian Euryglossinae, is represented
across the continent by 15 genera and 385
species, many of which occur in great
abundance. A second colletid subfamily,
the Hylaeinae, is much more diverse in
Australia than elsewhere. Its largest genus,
Hylaeus, is believed to have originated in
Australia and subsequently spread to other
regions (Kayaalp et al. 2013). The colletid
tribe Paracolletini (as interpreted by
Michener 2007) is also very diverse and
species-rich in Australia, as it is in South
America (Almeida et al. 2012).
The evolutionary history and biogeog-
raphy of Australia’s bee fauna is a subject
still relatively poorly understood but one
attracting increasing attention, often
assisted by modern genetics. Bees are
believed to have evolved during the Early
Cretaceous before the break-up of Gond-
wana and one bee family in particular, the
Colletidae, has long been viewed as a
primitive group that was present in Gond-
wana before the separation of Africa and
then South America (Almeida et al. 2012).
The colletid tribe Paracolletini and one of
its largest genera, Leioproctus, is shared
between Australia and South America.
Comprising mainly ground-nesting bees,
paracolletines are unlikely to have dis-
persed across expanses of ocean. By
contrast, some wood-nesting bees,
­
19
A G U I D E T O N AT I V E B E E S O F A U S T R A L I A
­particularly Megachilidae and Xylocopi-
nae, could potentially have reached Aus-
tralia more recently from elsewhere by
‘rafting’ in driftwood. The xylocopine
tribe Allodapini is most diverse in Africa
and genetic studies suggest it originated
on that continent, some species later dis-
persing to Madagascar, Asia and Aus-
tralia. This would necessarily involve
crossing expanses of ocean. Chenoweth
and Schwarz (2011) suggested that the
ancestors of Australia’s exoneurine allo-
dapines (Exoneura and close relatives)
could have arrived from the south-east via
a then temperate Antarctica. A separate
study (Fuller et al. 2005) postulates that
the allodapine genus Braunsapis probably
spread from Africa through Asia and then
to Australia from the north. Rainforest-
20
associated bees (such as Palaeorhiza,
Reepenia and Ctenoplectra) that are shared
with Papua New Guinea are believed to
have arrived from that region via a land
bridge during the last glacial maximum
when the sea level was significantly lower
than today.
Within Australia, bees occupy almost
all terrestrial habitats, from coastal dunes
to the central deserts. The greatest diver-
sity of species is found in the semi-arid,
temperate regions while diversity is least in
the coolest and wettest regions and is
diminished in the tropical north. Distinc-
tive bee communities are to be found in
different regions, such as the arid interior,
the rainforest habitats of eastern Australia
and the floristically rich south-west of
Western Australia.
Another random document with
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 Figure 51. William Sellers

This paper had as great influence in America as Whitworth’s paper

of 1841 had in England. A committee was appointed to investigate
the question and recommend a standard. On this committee, among
others, were William B. Bement, C. T. Parry of the Baldwin
Locomotive Works, S. V. Merrick, J. H. Towne, and Coleman Sellers.
Early in the next year the committee reported in favor of the Sellers
standard, the Franklin Institute communicated their findings to other
societies, and recommended the general adoption of the system
throughout the country. The Sellers’ thread was adopted by the
United States Government for all government work in 1868, by the
Pennsylvania Railroad in 1869, the Master Car Builders’ Association
in 1872, and soon became practically universal. After exhaustive
investigation the Sellers’ form of thread was adopted in 1898 by the
International Congress for the standardization of screw threads, at
Zurich, and is now in general use on the continent of Europe.[209]
[209] For the discussion of the Sellers’ screw thread and the
circumstances surrounding its adoption, see: Journal of the Franklin
Institute, Vol. LXXVII, p. 344; Vol. LXXIX, pp. 53, 111; Vol. CXXIII, p. 261;
Vol. CXXV, p. 185.

In 1868 William Sellers organized the Edgemoor Iron Company

which furnished the iron work for the principal Centennial buildings
and all the structural work of the Brooklyn Bridge. In the
development of this business, he led the way in the distinctly
American methods and machinery by which the building of bridges
has been, to a great extent, put upon a manufacturing basis. This
involved the design and introduction of hydraulic machinery, large
multiple punches, riveters, cranes, boring machines, etc.
The excellence of his machinery soon brought him into contact
with government engineers and throughout his life his influence in
the War and Navy Departments was great. In 1890 the Navy
Department called for bids on an eight-foot lathe, with a total length
of over 128 feet, to bore and turn sixteen-inch cannon for the Naval
Gun Factory at Washington. Sellers disapproved of the design and
refused to bid on it. He proposed an alternative one of his own,
argued its merits in person before the Board of Engineers, and
secured its adoption and a contract for it. This great lathe, weighing
over 500,000 pounds, has attracted the attention of engineers from
all parts of the world. In 1873 Mr. Sellers reorganized the William
Butcher Steel Works as the Midvale Steel Company and became its
president. Under his management the company grew rapidly, and
later became a leader in production of heavy ordnance.
It was here that Frederick W. Taylor began in 1880 his work on the
art of cutting metals, which resulted in modern high-speed tool steels
and a general re-design of machine tools. These experiments,
covering a period of twenty-six years, cost upwards of $200,000. Mr.
Taylor has frequently acknowledged his indebtedness in this work to
the patience and courage of Mr. Sellers, who was then an old man
and might have been expected to oppose radical change. It was he
who made the work possible, however, and he supported Taylor
unwaveringly in the face of constant protests.[210] Mr. Sellers was a
man of commanding presence, direct but gracious in manner, who
won and held the respect and loyalty of all about him. His judgment
was almost unerring and he dominated each of the great
establishments he built up.
[210] F. W. Taylor: Paper on the “Art of Cutting Metals,” Trans. A. S. M. E.,
Vol. XXVIII, p. 34.

The firm of William Sellers & Company had another master mind
in that of Dr. Coleman Sellers, a second cousin of William
Sellers.[211] He was born in Philadelphia in 1827, his father, Coleman
Sellers, being also an inventor and mechanic. Like Nasmyth he
spent his school holidays in his father’s shop, which was at
Cardington. In 1846, when he was nineteen years old, he went to
Cincinnati and worked in the Globe Rolling Mill, operated by his elder
brothers, where the first locomotives for the Panama Railroad were
built; and in two years he became superintendent. In 1851 he
became foreman of the works of James and Jonathan Niles, who
were then in Cincinnati and building locomotives. Six years later he
returned to Philadelphia, became chief engineer of William Sellers &
Company, and remained with them for over thirty years, becoming a
partner in 1873. During these years he designed a wide range of
machinery, which naturally covered much the same field as that of
William Sellers, but his familiarity with locomotive work especially
fitted him for the design of railway tools. His designs were original,
correct and refined. The Sellers coupling was his invention and he
did much to introduce the modern systems of power transmission.
[211] See Trans. A. S. M. E., Vol. XXIX, p. 1163; Cassier’s Magazine,
August, 1903, p. 352; Journal of the Franklin Institute, Vol. CXLIX, p. 5.

Doctor Sellers was a good physicist, an expert photographer,

telegrapher, microscopist, and a professor in the Franklin Institute,
his lectures always drawing large audiences. Like William Sellers, he
was a member of most of the great engineering and scientific
societies, here and abroad; and he was president of the American
Society of Mechanical Engineers, of which he was a charter
member. He was received with the greatest distinction in his visits to
Europe. In 1886 impaired health compelled his relinquishing regular
work and he resigned his position of engineer for William Sellers &
Company, being succeeded by his son, the present president of the
company. His last great work was in connection with the power
development of Niagara Falls. He was engineer for the Cataract
Construction Company and served on the commission which
determined the types of turbines and generators and the methods of
power transmission finally adopted. Among the others on this
commission were Lord Kelvin, Colonel Turretini, the great Swiss
engineer, and Professor Unwin, and its report forms the foundation
of modern large hydro-electric work. William Sellers & Company has
a unique distinction among the builders of machine tools in having
had the leadership of two such men as William and Coleman Sellers.
William B. Bement, the son of a Connecticut farmer and
blacksmith, was born at Bradford, N. H., in 1817. His education was
obtained in the district schools and in his father’s blacksmith shop.
His mechanical aptitude was so clear that he was apprenticed to
Moore & Colby, manufacturers of woolen and cotton machinery at
Peterboro, N. H. His progress at first was rapid. Within two years he
became foreman, and on the withdrawal of one of the partners, was
admitted into the firm. He continued there three years, already giving
much thought to machine tools, for which he saw the rising need. In
1840 he went to Manchester and entered the Amoskeag shop when
it was just finished, remaining there two years as a foreman and
contractor under William A. Burke, to whom we have referred
elsewhere. From there Bement went to take charge of a shop for
manufacturing woolen machinery at Mishawaka, Ind. Unfortunately it
was burned to the ground while Bement had gone back to New
Hampshire for his family, so that when he returned with them he
found himself without employment and with only ten dollars in hand.
For the time being he worked as a blacksmith and gunsmith, and
made an engine lathe for himself in the shop of the St. Joseph Iron
Company, which gave him permission to use their tools in return for
the use of his patterns to make a similar machine for themselves.
Much of the work in making this lathe was done by hand as there
was no planer within many hundred miles. The St. Joseph Iron
Company, seeing his work, offered him the charge of their shop, to
which he agreed, provided the plant were enlarged and equipped
with proper tools. This was done, but just as everything was
completed this plant also was burned down. Bement had plans for
another shop ready the following day, went into the woods with
others, cut the necessary timber, and a new shop was soon
completed. He remained there for three years, constructing a variety
of machine tools, one of which was a gear cutter said to have been
the first one built in the West, or used beyond Cleveland.
Figure 52. Coleman Sellers
 Figure 53. William B. Bement

He returned to New England as a contractor in the Lowell Machine

Shop under Burke, who had gone there from the Amoskeag Mills in
1845. On account of Bement’s resourcefulness and skill in
designing, Burke induced him to relinquish his contracts and take
charge of their designing, which he did for three years, his residence
at Lowell covering in all about six years.
In 1851 Elijah D. Marshall, who had established a business of
engraving rolls for printing calicos in 1848 and had a small shop at
Twentieth and Callowhill Streets in Philadelphia, offered Bement a
partnership. He moved to Philadelphia in September of that year,
and with Marshall and Gilbert A. Colby, a nephew, he began the
manufacture of machine tools under the name of Marshall, Bement &
Colby, thus starting only a year or so after Sellers. Marshall was a
large man, dignified and deliberate in speech. Bement was strong,
vigorous, a born designer, a remarkably rapid draftsman, and had a
capacity for work rarely equalled. Colby was also a man of
considerable mechanical ability, with advanced business ideas. Their
shop consisted of a single three-storied, stone, whitewashed
building, 40 by 90 feet. Their entire machine shop was on the first
floor, with a 10- by 12-foot room for an office. The engine, boiler and
blacksmith shop were in small outbuildings. Part of the second floor
was rented to another factory and the rest was sometimes used for
religious meetings, while the third floor was used for engraving
printing rolls. Their tools were few and crude; among them were a
36-inch lathe with a wooden bed and iron straps for ways, and a 48-
inch by 14-foot planer with ornate Doric uprights. Marshall and Colby
soon retired, the latter going to Niles, Mich., where he was very
successful. James Dougherty, an expert foundryman, and George C.
Thomas entered the firm, which became Bement & Dougherty, the
plant being known as the “Industrial Works.” Mr. Thomas contributed
considerable capital, and a new shop and a foundry were built. At
the same time they installed a planer 10 feet wide by 8 feet high, to
plane work 45 feet long, a notable tool for that day.
After a few years of struggle, the plant began to grow rapidly and
at one time was the largest of its kind in the country. Bement and
Sellers were among the first to concentrate wholly on tool building.
They confined themselves to work of the highest quality. Both made
much heavier tools, as we have said, than the New England
builders, their only competitors, and in a short time had established
great reputations. Bement relied little on patent protection, trusting to
quality and constant improvement. Thomas retired from the
partnership in 1856 and Dougherty in 1870; and Clarence S. Bement
joined the firm, which became William B. Bement & Son. John M.
Shrigley became a partner in 1875, William P. Bement in 1879, and
Frank Bement in 1888.
Frederick B. Miles was an employee of Bement & Dougherty who
established a tool business under the name of Ferris & Miles, which
afterward became the Machine Tool Works. While head of these
works, Miles greatly improved the steam hammer, particularly its
valve mechanism, and many details of what is known as the Bement
hammer were invented by Miles. In 1885 the Machine Tool Works
consolidated with William Bement & Son, forming Bement, Miles &
Company. Mr. Miles was an accomplished engineer and designer,
with the unusual equipment of six languages at his command, an
asset of value in the firm’s foreign business. William Bement, Senior,
died in 1897, and in 1900 the business became a part of the Niles-
Bement-Pond Company. Mr. Miles retired at that time and has not
since been active in the tool business.[212]
[212] Most of the foregoing details in regard to the Bement & Miles Works
have been obtained from Mr. Clarence S. Bement and Mr. W. T. Hagman,
their present general manager.

Although Bement and Sellers contributed more to the art of tool

building than any of the other Philadelphia mechanics, some of these
others ought to be mentioned. Matthias W. Baldwin, a native of New
Jersey, began as a jeweler’s apprentice. In partnership with David H.
Mason he began making bookbinders’ tools, to which he added in
1822 the engraving of rolls for printing cotton goods and later of bank
notes. From the invention and manufacture of a variety of tools used
in that business they were led gradually into the machine tool
business, the building of hydraulic presses, calender rolls, steam
engines, and finally locomotives. In 1830 Baldwin built a model
locomotive for the Peale Museum which led to an order from the
Philadelphia & Germantown Railroad for an engine which was
completed in 1832 and placed on the road in January, 1833. An
advertisement of that time says: “The locomotive engine built by Mr.
M. W. Baldwin of this city will depart daily, when the weather is fair,
with a train of passenger cars. On rainy days horses will be attached
in the place of the locomotive.”
From this beginning has sprung the Baldwin Locomotive Works,
which employs approximately 20,000 men. In 1834 they built five
locomotives; in 1835, fourteen; in 1836, forty. Their one thousandth
locomotive was built in 1861; the five thousandth in 1880 and the
forty thousandth in 1913. These works have naturally greatly
influenced the neighboring tool makers. From the beginning, both
Bement and Sellers specialized on railway machinery and they have
always built a class of tools larger than those manufactured in New
England.
The Southwark Foundry was established in 1836, first as a
foundry only, but a large machine shop was soon added. The owners
were S. V. Merrick, who became the first president of the
Pennsylvania Railroad Company, and John Henry Towne, who was
the engineering partner. The firm designed and built steam engines
and other heavy machinery and introduced the steam hammer into
the United States under arrangement with James Nasmyth. From the
designs of Capt. John Ericsson they built the engines for the
“Princeton,” the first American man-of-war propelled by a screw, and
later were identified with the Porter-Allen steam engine. Mr. Towne
withdrew from the firm about 1848, and the firm name became
successively Merrick & Son, Merrick & Sons, Henry G. Morris, and
finally the Southwark Foundry & Machine Company.
I. P. Morris & Company came from Levi Morris & Company,
founded in 1828, and for many years were engaged in a similar
work. In 1862 Mr. J. H. Towne, above referred to, was admitted to
the firm as the engineering partner, and the firm name then became
I. P. Morris, Towne & Company, until about 1869 when Mr. Towne
withdrew. At his withdrawal the firm name was restored to its original
form, I. P. Morris & Company. It is now a department of the Cramp
Ship Building Company. During the Civil War the works were
occupied largely in building engines and boilers for government
vessels, and blast furnace and sugar mill machinery. During this
period Henry R. Towne, son of J. H. Towne, entered the works as an
apprentice, served in the drawing room and shops, and finally was
placed in charge of the erection at the navy yards of Boston and
Kittery of the engines, boilers, etc., built for two of the double-
turreted monitors. Returning to Philadelphia, he was made assistant
superintendent of the works.
J. H. Towne was a mechanical engineer of eminence in his day,
whose work as a designer showed unusual thoroughness and finish.
He was a warm friend and admirer of both William and Coleman
Sellers, and through his influence, Henry R. Towne was at one time
a student apprentice in the shops of William Sellers & Company,
acquiring there an experience which had a marked influence on his
future work. Both of the firms with which J. H. Towne was connected
built machine tools for themselves and for others, especially of the
heavier and larger kinds, and thus were among the early tool
builders. I. P. Morris & Company, about 1860, designed and built for
their own use what was then the largest vertical boring mill in this
country.[213]
[213] From correspondence with Mr. Henry R. Towne.

It may surprise some to learn that the well-known New England

firm, the Yale & Towne Manufacturing Company in Stamford, Conn.,
is a descendant of these Philadelphia companies. It was organized
in October, 1868, by Linus Yale, Jr., and Henry R. Towne, who were
brought together by William Sellers. Mr. Yale died in the following
December. This company, under the direction and control of Mr.
Towne, has had a wide influence on the lock and hardware industry
in this country. While the products of the Yale & Towne
Manufacturing Company have always consisted chiefly of locks and
related articles, they have added since 1876 the manufacture of
chain blocks, electric hoists, and, during a considerable period, two
lines allied to tool building, namely, cranes and testing machines.
This company was the pioneer crane builder of this country,
organizing a department for this purpose as early as 1878, and
developing a large business in this field, which was sold in 1894 to
the Brown Hoisting Machine Company of Cleveland, Ohio. The
building of testing machines was undertaken in 1882, to utilize the
inventions of Mr. A. H. Emery, and was continued until 1887, when
this business was sold to William Sellers & Company, for the same
reason that the crane business was sold; namely, that both were
incongruous with the other and principal products of the company.
In recent years the Bilgram Machine Works, under the leadership
of Hugo Bilgram, an expert Philadelphia mechanic, has made
valuable contributions to the art of accurate gear cutting.
In the cities between New York and Philadelphia, and here and
there in the smaller towns of Pennsylvania, are several tool builders
of influence. Gould & Eberhardt in Newark is one of the oldest firms
in the business, having been established in 1833. Ezra Gould, its
founder, learned his trade at Paterson, and started in for himself at
Newark in a single room, 16 feet square. Within a few years the
Gould Machine Company was organized, the business moved to its
present location, and a line of lathes, planers and drill presses was
manufactured. To these they added fire engines. Ulrich Eberhardt
started as an apprentice in 1858 and became a partner in 1877, the
firm name becoming E. Gould & Eberhardt, and later Gould &
Eberhardt. Mr. Gould retired in 1891, and died in 1901. Mr. Eberhardt
also died in 1901; the business has since been incorporated and is
now under the management of his three sons. They employ about
400 men in the manufacture of gear and rack cutting machinery and
shapers.
The Pond Machine Tool Company, which moved from Worcester
to Plainfield, N. J., in 1888, was founded by Lucius W. Pond.[214] It is
a large and influential shop and one of the four plants of the Niles-
Bement-Pond Company. Their output is chiefly planers, boring mills
and large lathes.
[214] See p. 222.

The Landis Tool Company, of Waynesboro, Pa., builders of

grinding machinery, springs from the firm of Landis Brothers,
established in 1890 by F. F. and A. B. Landis. One was
superintendent and the other a tool maker in a small plant building
portable engines and agricultural machinery. A small Brown &
Sharpe grinding machine was purchased for use in these works. Mr.
A. B. Landis became interested in the design of a machine more
suited to their particular work, and from this has developed the
Landis grinder.
 CHAPTER XX
THE WESTERN TOOL BUILDERS
Prior to 1880 practically all of the tool building in the United States
was done east of the Alleghenies. The few tools built here and there
in Ohio and Indiana were mostly copies of eastern ones and their
quality was not high. In fact, there were few shops in the West
equipped to do accurate work. “Chordal’s Letters,” published first in
the American Machinist and later in book form,[215] give an excellent
picture of the western machine shop in the transition stage from
pioneer conditions to those of the present day.
[215] Henry W. See: “Extracts from Chordal’s Letters”; McGraw-Hill Book
Co., N. Y. 12th Edition. 1909.

Good tool building appeared in Ohio in the early eighties, and

within ten years its competition was felt by the eastern tool builders.
The first western centers were Cleveland, Cincinnati and Hamilton.
Of these, Cleveland seems to have been the first to build tools of the
highest grade.
We have already noted that the Pratt & Whitney shop in Hartford
furnished Cleveland with a number of its foremost tool builders. The
oldest of these and perhaps the best known is the Warner & Swasey
Company. This company has the distinction, shared with only one
other, of having furnished two presidents of the American Society of
Mechanical Engineers. Oddly enough the other company is also a
Cleveland firm, the Wellman, Seaver, Morgan Company, builders of
coal- and ore-handling machinery, and of steel mill equipment.
Worcester E. Warner, of the Warner & Swasey Company, was
born at Cummington, Mass., in 1846. Although a farmer’s son and
denied a college education, he had access in his own home to an
admirable library, which he used to great advantage. When nineteen
years old he went to Boston and learned mechanical drawing in the
office of George B. Brayton. Shortly afterwards he was transferred to
the shop at Exeter, N. H., where he first met Ambrose Swasey. Mr.
Swasey was born at Exeter, also in 1846, went to the traditional “little
red schoolhouse,” and learned his trade as a machinist in the shop
to which Warner came. In 1870 they went together to Hartford,
entered the Pratt & Whitney shop as journeymen mechanics, and in
a short time had become foremen and contractors. Mr. Swasey soon
gained a reputation for accurate workmanship and rare ability in the
solution of complex mechanical problems. He had charge of the gear
department, and invented and developed a new process of
generating spur gear teeth, which was given in a paper before the
American Society of Mechanical Engineers.[216] Mr. Warner, also,
became one of the company’s most trusted mechanics, was head of
the planing department, and had charge of the Pratt & Whitney
exhibit at the Centennial Exposition in Philadelphia.
[216] Trans. A. S. M. E., Vol. XII, p. 265.

In 1881 they left Hartford and went first to Chicago, intending to

build engine lathes, each putting $5000 into the venture; but finding
difficulty in obtaining good workmen there, they moved in about a
year to Cleveland, where they have remained. Their first order was
for twelve turret lathes, and they have built this type of machine ever
since. At various times they have built speed lathes, die-sinking
machines, horizontal boring mills, and hand gear-cutters, but they
now confine their tool building to hand-operated turret lathes. They
have never built automatics.
Figure 54. Worcester R. Warner
Figure 55. Ambrose Swasey
 The building of astronomical instruments was not in their original
scheme, but Mr. Warner’s taste for astronomy and Mr. Swasey’s skill
in intricate and delicate mechanical problems, led them to take up
this work. These instruments, usually designed by astronomers and
instrument makers, were in general much too light; at least the large
ones were. From their long experience as tool builders, Warner and
Swasey realized that strength and rigidity are quite as essential as
accuracy of workmanship where great precision is required. The
design of a large telescope carrying a lens weighing over 500
pounds at the end of a steel tube forty or sixty feet long, and
weighing five or six tons, which must be practically free from flexure
and vibration and under intricate and accurate control, becomes
distinctly an engineering problem. To this problem both Mr. Warner
and Mr. Swasey brought engineering skill and experience of the
highest order.
When the trustees of the Lick Observatory called in 1886 for
designs for the great 36-inch telescope, Warner & Swasey submitted
one which provided for much heavier mountings than had ever been
used before, and heavier construction throughout. They were
awarded the contract and the instrument was built and installed
under Mr. Swasey’s personal supervision. It is located on the very
top of Mount Hamilton in California, 4200 feet above sea-level; and
to give room for the observatory 42,000 tons of rock had to be
removed. The great instrument, weighing with its mountings more
than forty tons, “was transported in sections, over a newly made
mountain road, sometimes in a driving snowstorm, with the wind
blowing from sixty to eighty miles an hour.”[217]
[217] Cassier’s Magazine, March, 1897, p. 403.

As is well known, the instrument was a brilliant success. The

Warner & Swasey Company has since designed and built the
mountings for the United States Naval Observatory telescope, the
40-inch Yerkes telescope, the 72-inch reflecting telescope for the
Canadian Government, and the 60-inch reflecting telescope for the
National Observatory at Cordoba, Argentina, the largest in use in the
southern hemisphere. In addition to this large work, the firm has built
meridian circles, transits and other instruments for astronomical
work, range finders for the United States Government, and
introduced the prismatic binocular into this country.
In connection with this astronomical work Mr. Swasey designed
and built a dividing engine capable of dividing circles of 40 inches in
diameter with an error of less than one second of arc. A second of
arc subtends about one-third of an inch at the distance of one mile.
Although the graduations on the inlaid silver band of this machine
are so fine that they can scarcely be seen with the naked eye, the
width of each line is twelve times the maximum error in the automatic
graduations which the machine produces.
Although their reputation as telescope builders is international,
Warner & Swasey are, and always have been, primarily tool builders.
They were not the first to build tools in the Middle West, but they
were the first to turn out work comparable in quality with that of the
best shops in the East.
The Warner & Swasey shop has had the advantage of other good
mechanics besides its proprietors. Walter Allen, an expert tool
designer, did his entire work with them, rising from apprentice to
works manager. Frank Kempsmith, originally a Brown & Sharpe
man, was at one time their superintendent. Lucas, of the Lucas
Machine Tool Company, was a foreman. George Bardons, who
served his apprenticeship with Pratt & Whitney, went west with
Warner and Swasey when they started in business and was their
superintendent; and John Oliver, a graduate of Worcester
Polytechnic, was their chief draftsman. The last two left Warner &
Swasey in 1891 and established the firm of Bardons & Oliver for
building lathes.
Another old Pratt & Whitney workman is A. W. Foote of the Foote-
Burt Company, builders of drilling machines. Unlike the others,
however, Foote did not work for Warner & Swasey.
The first multi-spindle automatic screw machines were
manufactured in Cleveland. The Cleveland automatic was developed
in the plant of the White Sewing Machine Company for their own
work, and its success led to the establishment of a separate
company for its manufacture. The Acme automatic was invented by
Reinholdt Hakewessel and E. C. Henn in Hartford. Mr. Hakewessel
was a Pratt & Whitney man and Mr. Henn a New Britain boy, who
had worked first in Lorain and Cincinnati and then for twelve years in
Hartford with Pratt & Cady, the valve manufactures. In 1895 Henn
and Hakewessel began manufacturing bicycle parts in a little
Hartford attic, developing for this work a five-spindle automatic.
Seven years later the business was moved to Cleveland, where it
became the National-Acme Manufacturing Company, organized by
E. C. and A. W. Henn and W. D. B. Alexander, who came from the
Union Steel Screw Works. Their business of manufacturing
automatic screw machinery and screw machine products has grown
rapidly and is now one of the largest industries in Cleveland.
The White Sewing Machine Company and the Union Steel Screw
Works were among the first in Cleveland to use accurate methods
and to produce interchangeable work. It was at the Union Steel
Screw Works that James Hartness, of the Jones & Lamson Machine
Company, got his first training in accurate work. Their shop practice
was good and was due to Jason A. Bidwell, who came from the
American Tool Company of Providence.
The Standard Tool Company is an offspring of Bingham &
Company, Cleveland, and of the Morse Twist Drill Company of New
Bedford, Mass. From the Standard Tool Company has come the
Whitman-Barnes Company of Akron, and from that the Michigan
Twist Drill and Machine Company.
Newton & Cox was established in 1876, and built planers and
milling machines. Mr. Newton sold his share in the business to F. F.
Prentiss in 1880, went to Philadelphia, and started the Newton
Machine Tool Works. Cox & Prentiss later became the Cleveland
Twist Drill Company. They drifted into the drill business through not
being able to buy such drills as they required. They began making
drills first for themselves, then for their friends, and gradually took up
their manufacture, giving up the business in machine tools.
Cincinnati is said to have upwards of 15,000 men engaged in the
tool building industry, and to be the largest tool building center in the
world. There are approximately forty firms there engaged in this
work, many of them large and widely known.
This development, which has taken place within the past thirty-five
years, may possibly have sprung indirectly from the old river traffic.
Seventy years ago this traffic was large, and Cincinnati did the
greater part of the engine and boat building and repair work. When
the river trade vanished, the mechanics engaged in this work were
compelled to turn their attention to something else, and there may be
some significance in the coincidence of the rise of tool building with
the decline of the older industry.
There had been more or less manufacturing in Cincinnati for many
years, but little of it could be described as tool building. Miles
Greenwood established the Eagle Iron Works in 1832 on the site
now occupied by the Ohio Mechanics Institute. It comprised a
general machine shop, an iron foundry, brass foundries, and a
hardware factory which rivaled those of New England, employing in
all over 500 men. The hardware factory was important enough to
attract the special attention of the English commissioners who visited
this country in 1853.
In the fifties and early sixties, Niles & Company built steamboat
and stationary engines, locomotives and sugar machinery, and
employed from 200 to 300 men. This company was the forerunner of
the present Niles Tool Works in Hamilton. Lane & Bodley were
building woodworking machinery about the same time, and J. A. Fay
& Company, another firm building woodworking machinery, which
started in Keene, N. H., began work in Cincinnati in the early sixties.
The first builder of metal-working tools in Cincinnati was John
Steptoe; in fact, he is said to have been for many years the only tool
builder west of the Alleghenies. Steptoe came to this country from
Oldham, England, some time in the forties. It is said that he was a
foundling and that his name came from his having been left on a
doorstep. He was married before he came to Cincinnati, and had
served an apprenticeship of seven years, although he was so young
in appearance that no one would believe it. After working some time
for Greenwood, he started in business for himself, making a foot
power mortising machine and later a line of woodworking tools. The
first metal-working tool which he built was a copy of the Putnam
lathe. With Thomas McFarlan, another Englishman, he formed the
firm of Steptoe & McFarlan, and his shop, called the Western
Machine Works, employed by 1870 about 300 men. Their old
payrolls contain the names of William E. Gang of the William E.