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The Black-capped Chickadee
 The Black-capped
CHICKADEE
Behavioral Ecology and Natural History

Susan M. Smith
Department of Biological Sciences
Mount Holyoke College

Comstock Publishing Associates

a division of
Cornell University Press
Ithaca and London
 For May Annetts Smith

Copyright © 1991 by Cornell University

All rights reserved. Except for brief quotations in a review, this book,
or parts thereof, must not be reproduced in any form without
permission in writing from the publisher. For information, address
Cornell University Press, 124 Roberts Place, Ithaca, New York
14850.

First published 1991 by Cornell University Press.

International Standard Book Number 0-8014-2382-1 (cloth)

International Standard Book Number 0-8014-9793-0 (paper)
Library of Congress Catalog Card Number 91-55072

Printed in the United States of America

Librarians: Library of Congress cataloging information appears

on the last page of the book.

© The paper in this book meets the minimum requirements of the

American National Standard for Information Sciences—Permanence
of Paper for Printed Library materials, ANSI Z39.48-1984.

Epigraph: Henry David Thoreau, The Journal of Henry D. Thoreau, ed.

Bradford Torrey and Francis H. Allen (1906; reprint [14 vols. in 2], New
York: Dover Publications, 1962), 2: 1386-1387.
 Contents

Preface
The Black-capped Chickadee and Related Species x
1. General Natural History 1
2. Some Study Techniques 1?
3. Food and Feeding 28
4. Communication 55
5. The Early Breeding Season 90
6. The Later Part of the Breeding Season 125
7. The Nonbreeding Season 168
8. Surviving the Winter 223
9. Survivorship and Population Dynamics 253
10. Future Research 313
Glossary 321
References 326
Index 355
The fields are bleak, and they are, as it were, vacated.
The very earth is like a house shut up for the winter, and
I go knocking about it in vain. But just then I heard a
chickadee on a hemlock, and was inexpressably cheered
to find that an old acquaintance was yet stirring about
the premises, and was, I was assured, to be there all
winter. All that is evergreen in me revived at once.
—Henry David Thoreau
November 7, 1858
 Preface

Familiar to many and immensely popular, the Black-capped

Chickadee is one of the best-studied species in North America.
When I set out to write this book, I simply wanted to produce a
compilation of everything I could Find published about these re¬
markably interesting birds. As my work progressed, however, I
realized I had underestimated my task.
First, two other species—the Willow Tit of the Old World and
the North American Carolina Chickadee—have been considered
by some to be the same species as the Black-capped Chickadee. I
have therefore included as close to complete coverage as I can of
the literature on Carolina Chickadees and much of the recent
literature on Willow Tits. This in turn has forced me to attempt to
summarize the recent (and enormous) literature on European tits,
at least as it pertains to our knowledge of chickadees: a rather
daunting task.
One of my goals in writing this book was to encourage people to
write more about chickadees. Although I have tried to present all
the information published so far, as fairly and clearly as possible, I
am well aware that not all my interpretations or conclusions are
necessarily correct—just that they do fit the data currently avail¬
able to me. Science constantly strives to get closer to the truth, and
I hope this book will encourage people with unpublished informa¬
tion about chickadees to write it up. From the review of the rele-
viii Preface

vant literature provided here, they can easily see how their data fit
in with what else is known about the subject.
The book opens with an introductory chapter that brings to¬
gether some basic information on the natural history of Black-
capped Chickadees. Next, Chapter 2 discusses some of the more
common study techniques used in work with chickadees. Chapters
3 and 4 deal with two important daily activities of chickadee life:
food and feeding (Chapter 3) and communication (Chapter 4).
The next three chapters follow Black-capped Chickadees through
their annual cycle, paying particular attention to behavior and
social organization throughout the year. Chapter 8 deals with vari¬
ous aspects of chickadee physiology, focusing on an exploration of
how such tiny birds can withstand the long winter nights without
freezing to death. Chapter 9 explores survivorship and population
dynamics. Finally, Chapter 10 discusses some of the more exciting
areas for future research.
I have tried to make each chapter able to stand alone, since some
readers may wish to go straight to the chapters that interest them
most. To this end, I have given the scientific names of other chick¬
adees and titmice the first time I mention them in each chapter. To
minimize overlap of information, I refer readers to the place
where a topic is covered most fully.
Throughout the book I have been careful to identify the locality
of each study. Some readers may wonder why I bothered to in¬
clude this information. The Black-capped Chickadee is a non-
migratory bird that occupies an unusually large geographic range.
Population differences in morphological characteristics such as
wing and tail length or plumage coloration have been well docu¬
mented. It is likely that other consistent differences, including
some in behavioral ecology, also occur; indeed, a few have already
been discovered. I find such differences particularly exciting be¬
cause comparative studies between two differing populations of a
single species seem to have enormous potential for allowing us to
discover how and why such differences arise and are maintained.
I recently read a review by Charles Blem (1989) in which he
mentioned a complaint he had heard about today’s “chart- and
table-studded articles” on bird behavior, and it occurred to me that
some readers might be dismayed by the profusion of charts and
tables in this book. With that in mind, I tried to make all the
general points in the text itself; the tables and graphs merely
provide illustrations of the data on which those points are based.
 Preface ix

The tables and graphs are there for readers who wish to use them,
but my hope is that readers who choose not to will be able to read
right on with comprehension but without distraction.
Studying chickadees can be addictive—I’ve been hooked for
years. I can’t imagine ever getting tired of working with them.
They are endlessly fascinating, endlessly intriguing. Each new bit
of information discovered simply brings up important new ques¬
tions to pursue. I hope this book not only conveys but also explains
my excitement in working with these truly marvelous birds.

During the writing of this book, much of my thinking has been

shaped and clarified by conversations with other biologists. Even
those who did not necessarily agree with my interpretations (and
still don’t!) were extraordinarily generous in taking time to talk
over ideas and data with me. I am particularly grateful for discus¬
sions with Christopher P. L. Barkan, Andre Desrochers, Andre A.
Dhondt, Jan Ekman, Millicent Sigler Ficken, Thomas C. Grubb,
Jr., Svein Haftorn, Jack P. Hailman, Christine L. Hitchcock, Olav
Hogstad, James L. Howitz, Donald E. Kroodsma, D. Archibald
McCallum, Alison M. Mostrom, Stephen Nowicki, Craig D. Orr,
Christopher M. Perrins, Laurene Ratcliffe, David F. Sherry, and
Charles M. Weise.
Many people shared unpublished data and manuscripts with
me; I especially thank Christopher Barkan, Andre Dhondt, Jan
Ekman, Thomas C. Grubb, Jr., Christine Hitchcock, Olav Hog¬
stad, Lloyd Kiff, Donald Kroodsma, Archibald McCallum, Alison
Mostrom, and Stephen Nowicki.
It is a pleasure to thank Christopher and Mary Perrins for their
hospitality in Oxford; I am grateful for their generosity and kind¬
ness while I was there. The discussions we had during that short
time were most helpful.
Special thanks go to my friends Muriel W. Harris and Janice A.
Gifford for lending me negatives of their photographs to use in
this book, to Marjorie Kaufman for tracking down the particulars
on the quotation from Thoreau’s journal, and to Doris S. Atkinson
for help with many of the figures as well as for continued support
throughout the preparation of the manuscript.

Susan M. Smith

South Hadley, Massachusetts

 The Black-capped Chickadee and Related Species
These sketches of the main species discussed appear on figures
throughout the book to identify the species on which the study was
done.

Black-capped Chickadee
(.Parus atricapillus)

Carolina Chickadee
(Parus carolinensis)
 Willow Tit
(.Parus montanus)

Crested Tit
{Parus cristatus)

Great Tit
{Parus major)

Blue Tit
{Parus caeruleus)
The Black-capped Chickadee
 1
General Natural History

The Black-capped Chickadee (Parus atricapillus) is one of the

most familiar and popular of North American birds. Found from
coast to coast, including much of Canada and approximately the
northern two-thirds of the United States, chickadees readily ap¬
proach people’s houses and show little fear of humans. Flocks of
chickadees crowd to feeders in areas where most other songbirds,
especially those as small as chickadees, have left for the winter.
Who has not gladly stopped to watch the antics of these birds on a
cold winter day?
Another factor contributing to their popularity is that chick¬
adees, unlike many other common birds, do not cause any serious
nuisance to people. In fact, their summer diet is primarily insec¬
tivorous, making their presence all the more welcome.
Chickadees make ideal research subjects. They are resident
year-round, so field studies can be done in any season. They come
readily to feeders, making certain kinds of observations easier to
get. They are easily caught for banding. Relatively unafraid of
humans, they can be approached closely for behavioral observa¬
tions. And even wild-caught adults adapt readily to captivity. For
such reasons, there is a vast and growing literature on the biology
of these birds. But, as Gill and Flicken (1989) pointed out, re¬
search opportunities on chickadees are nowhere close to being
exhausted—more information simply generates new and fresh
questions about chickadee biology.
2 The Black-capped Chickadee

The more than 40 species of chickadees and titmice (genus

Parus), found in most habitats in the Northern Hemisphere, pro¬
vide ideal subjects for comparative studies. Moreover, many, in¬
cluding the Black-capped Chickadee, inhabit enormous and di¬
verse geographical ranges, permitting within-species comparisons
as well (something that is essential to any understanding of
between-species data). Some of the most exciting work on avian
learning and memory is being conducted today on Black-capped
Chickadees (see Food Storage and Memory, Chapter 3); this work,
along with certain other chickadee studies (e.g., Nowicki and Ca-
pranica 1986a, 1986b), helps to increase our understanding of
how the avian brain actually operates. Research on how chickadees
respond to cold temperatures has helped further our understand¬
ing of the winter physiology of warm-blooded vertebrates. Long¬
term studies on chickadees and titmice continue to provide the
basis for many of our insights into vertebrate population ecology.
Chickadees have already taught us a great deal and will surely
continue to do so in the foreseeable future.

Appearance and Behavior

All North American chickadees have the same basic pattern of

dark cap and bib; white cheeks; unstreaked dark back, wings, and
tail; and pale underparts with a darker wash along the flanks and
under the tail (the crissum). The cap and bib of Black-capped
Chickadees are solid black, the back is usually some shade of
greenish gray, there is at least some white edging on the gray wing
feathers (including the wing coverts), and the wash on the flanks
and crissum is often a warm buff color (Fig. 1-1). Occasionally,
chickadees with some form of aberrant plumage have been re¬
ported (Fig. 1-2; also see, for example, Wetherbee 1933, Tanner
1934, Low 1969, and Tallman 1987). Often such birds have either
too much or too little dark pigment (melanin) in their feathers.
Some have been known to grow normally colored feathers after
going through a molt (Tallman 1987; see below).
Chickadees are small. Most Black-capped Chickadees weigh be¬
tween 10 and 14 grams, with northern individuals being perhaps
somewhat heavier than their southern counterparts. By contrast,
House Sparrows {Passer domesticus) weigh about 30 grams, and
 General Natural History 3

Fig. 1-1. The Black-capped Chickadee. Note the prominent pale edgings on
both the upper wing feathers (secondaries) and the wing coverts. (© Alan
Cruickshank / VIREO.)

American Robins (Turdus migratorius) often weigh well over 80

grams.
Chickadees are most definitely not known for their fast flight.
Evidently, their ordinary flight speed is approximately 11 miles
4 The Black-capped Chickadee

Fig. 1-2. This partial albino Black-capped Chickadee had much white on the
top of its head. Curiously, although the rest of the feathers had normal
coloration, its beak and feet were pink, rather than dark as in normal chick¬
adees. (Photograph courtesy of Janice Gifford.)

per hour (just under 20 km per hour), a speed they typically

achieve after only four or five wingbeats (Greenewalt 1955). By
contrast, the White-breasted Nuthatch (Sitta carolinensis) has slower
acceleration but a faster terminal velocity.
Like all birds, chickadees must preen frequently to maintain
their feathers in good condition. Oil from the uropygial, or preen,
gland, located at the base of the tail, is spread over each feather,
increasing its flexibility and insulative properties. Where condi¬
tions permit, chickadees also bathe frequently. When taking a
bath, a chickadee typically jumps into the water, rolls its body and
flicks its wings, then retires to the bank or to a low perch to vibrate
its wet feathers and preen before returning for another plunge
 General Natural History 5

(Slessers 1970). In times of drought, chickadees may use dew for

bathing, fluttering repeatedly among wet leaves (Verbeek 1962);
they have also been known to take baths in new-fallen snow
(Mitchell 1950).
Some of the most interesting and readily observed chickadee
behavior relates to their feeding. Chickadees are seldom still; they
seem to be in almost constant motion in their never-ending search
for food. They and their relatives are well known for being partic¬
ularly acrobatic, often hanging upside down from the tips of twigs
and larger branches as they forage. The fact that chickadees’ legs
are both somewhat longer and stronger than those of most other
birds of similar weight (Perrins 1979, Moreno 1990) may account
for their impressive acrobatic abilities.
If a chickadee catches something that needs manipulation be¬
fore it can be eaten (such as a large caterpillar or even a sunflower
seed), the bird typically holds the food down with one or both feet
and pounds away with its short but stout bill. This use of the feet is
fairly unusual among birds, but is characteristic of virtually all
members of the genus Parus.
Chickadees’ appearance changes little as they age; that is, they
have no distinct immature plumage. Newly hatched Black-capped
Chickadees are sparsely covered with grayish natal down, but this
is soon pushed out by the growing contour feathers of their juve-
nal plumage, which is virtually identical to that of adults, although
the feathers are softer and fluffier than those of older birds.
Young chickadees next undergo a partial postjuvenal molt about
midsummer, in which essentially every feather except the larger
wing and tail feathers is replaced. Once past the postjuvenal molt,
every chickadee undergoes a single annual molt, typically begin¬
ning in July and August and continuing into autumn, in which
each feather is replaced (Bent 1946; see Postbreeding Molt, Chap¬
ter 6).
When you stop to think about it, the consequences of this simple
fact are quite remarkable. Since chickadees molt only once a year,
the feathers that keep them warm through the cold winter are the
same ones in which they must survive the summer’s heat. Of
course, some wear occurs over time, so the feathers are not quite
so thick by the time summer arrives (see Physical Adjustments
Chapter 8); nevertheless, the ability of such small birds to maintain
a fairly constant body temperature throughout the year with one
set of feathers is indeed amazing.
6 The Black-capped Chickadee

Life History

Black-capped Chickadees live in nonbreeding flocks in the fall

and winter, then shift to monogamous, territorial breeding pairs
during the spring and summer. Chickadees in winter flocks are
hard to miss, but during the breeding season chickadees are rela¬
tively silent and inconspicuous, as they go quietly about the busi¬
ness of nesting and rearing offspring.
As do most other members of the genus, Black-capped Chick¬
adees nest in holes, although they are far less likely to use nest
boxes than are many of their European relatives (Perrins 1979),
preferring instead to excavate their own cavities in stumps or rot¬
ting branches. Both sexes excavate nest cavities, but the female
alone builds the nest and incubates the eggs. Once the eggs hatch,
however, both parents share the duties of feeding the young. Such
feeding typically continues for up to four weeks after the young
have left the nest. Each young chickadee then disperses indepen¬
dently from its natal area, often moving several kilometers away.
Chickadees rarely have a second brood; by the time the fledglings
disperse, the adults will generally have begun their postbreeding
molt.
The dispersing young soon settle and join newly forming flocks.
These flocks typically consist of local breeding pairs together with
unrelated young birds that hatched some distance away. The
flocks, which persist through the nonbreeding season, typically
exhibit stable, linear dominance hierarchies (peck orders) in which
every member has its own particular position, or rank. A chick¬
adee’s rank within the flock can profoundly affect both its behavior
and its chances of survival (see Chapters 3 [Foraging Niche], 7
[Dominance Hierarchies], and 9 [Predators and Antipredator Be¬
havior]).
Throughout most of its range, the Black-capped Chickadee is
usually resident; that is, individuals inhabit the same area year-
round. Many of the most northerly chickadee populations don’t
migrate, even in the absence of feeders (Taverner 1949, Kron
1975). Nevertheless movements can and do occur. Some are sim¬
ply vertical: chickadees inhabiting mountainous areas may de¬
scend to lower, warmer localities for the winter, then return to
higher altitudes the following spring (Smith and Van Buskirk
1988). These migrants may well be mostly young birds; Dixon and
Gilbert (1964) found that altitudinal migration by Mountain
 General Natural History 7

Chickadees (Parus gambeli) is done almost exclusively by immature

birds. Other movements, such as the more familiar pattern of
temperate zone migration, are apparently latitudinal—several
banding studies contain reports of birds captured in fall and (to a
somewhat lesser extent) in spring that were not seen again (e.g.,
Loery and Nichols 1985). Moreover, in many winters, Black-
capped Chickadees occur farther south than they do during the
breeding season. Little is known about these apparent migrants;
however, two important features emerge. First, again, the vast
majority are young birds. Second, the number that moves in this
manner varies markedly from year to year; in some years appar¬
ently few to none move, while in others, called irruption years,
very large numbers may be involved (Bagg 1969; see Migration
and Irruptions, Chapter 9).

Longevity and Natural Enemies

The oldest known Black-capped Chickadee to date was at least

12 years and 5 months old (Kennard 1975). A bird of such age,
while not unique, is quite unusual. The average life span of a
chickadee is far shorter—approximately 2.5 years or less accord¬
ing to most studies (Smith 1967c, Elder and Zimmerman 1983).
Throughout the year, chickadees must withstand the attacks of
several natural predators. Among avian predators, hawks of the
genus Accipiter, such as Sharp-shinned (A. striatus) and Cooper’s (A.
cooperii) Hawks can be particularly efficient at capturing chick¬
adees. Other predators, such as American Kestrels (Falco spar-
verius) and Merlins (F. columbarius), and more locally, Northern
Pygmy Owls (Glaucidium gnoma) and Northern Shrikes (Lanius ex-
cubitor) can also take their toll (Bent 1946). Often, however, a
healthy chickadee is agile enough to escape such predators; the
individuals that are caught often turn out to have been weakened
by age, disease, or some other factor (Geer 1982).
The house cat, an introduced species, is most dangerous to
chickadees around that human-constructed item, the feeder. Nat¬
ural nonavian predators are more likely to catch chickadees at nest
or roost sites. Nest predators must be able to fit, either partially or
entirely, into the entrance hole of the nest cavity. Many such holes
are small enough to exclude all but the smallest squirrel (e.g.,
Tamiasciurus sp.) or chipmunk (Tamias or Eutamias sp.), although
8 The Black-capped Chickadee

certain weasels (Mustela sp.) and climbing snakes still pose a danger
(Bent 1946, Perrins 1979; see the section on predators and anti¬
predator behavior in Chapter 9 for a discussion of these and other
nest predators).
Brown-headed Cowbirds (.Molothrus ater) occasionally manage to
lay eggs in Black-capped Chickadee nests (Friedmann et al. 1977;
Lowther 1983). However, the cavity entrances of most chickadee
nests are too small for female cowbirds to enter, so such instances
are rare.
As cavity nesters go, chickadees support a relatively low level
and diversity of external parasites (see Parasites and Diseases,
Chapter 9)—perhaps because they seldom reuse a nest site, pre¬
ferring to excavate a new cavity each time they nest.

Relationships

All Northern American chickadees and crested titmice are cur¬

rently placed in the genus Parus, within the family Paridae. The
Paridae is a family of the suborder Oscines (songbirds) within the
order Passeriformes, or perching birds. This huge order contains
over half of all living bird species. Within this order, parids are
generally thought to be fairly closely related to the families Sittidae
(nuthatches) and Certhiidae (creepers and their allies) (Sibley et al.
1988). Members of the family Paridae inhabit more or less wooded
areas over most of North America, Eurasia, and much of Africa,
but are absent from both South America and Australia. Evidently
the North American parids are all descended from Eurasian lin¬
eages that crossed the Bering land bridge during the Pleistocene
(Mayr 1946, Parkes 1958).
The Black-capped Chickadee’s scientific name, Parus atricapillus,
was coined in 1766 by Linnaeus; roughly translated, it means “tit¬
mouse with a black crown.” The type specimen was collected in
Quebec City, Canada. At one time, the Black-capped Chickadee
was considered to be conspecific with the Eurasian Willow Tit.
However, they are now considered separate species, because of,
among other things, considerable differences in voice and, to some
extent, in plumage (Snow 1956). Although some biologists still
question this split (e.g., Desfayes 1964), recent work has provided
solid evidence that the two are separate species (Gill et al. 1989; see
below). The Willow Tit is now known as Parus montanus.
 General Natural History 9

Among North American chickadees (all of which are in the

subgenus Poecile), two groups have traditionally been recognized
(e.g., Mayr and Short 1970): the “brown-capped” chickadees, in¬
cluding the Boreal Chickadee (Parus hudsonicus), the Chestnut-
backed Chickadee (P. rufescens), and the Siberian Tit (P. cinctus);
and the “black-capped” group, consisting of the Black-capped
Chickadee, the Carolina Chickadee (P. carolinensis), the Mountain
Chickadee (P. gambeli), and the Mexican Chickadee (P. sclateri).
Both Mountain Chickadees, of western Canada and the United
States, and Carolina Chickadees, of the southeastern United
States, are known to hybridize with Black-capped Chickadees.
While Mountain/Black-capped hybrids are relatively scare, Caroli-
na/Black-capped hybrids are fairly common in certain areas, and
the whole problem of interaction at the interface of these two
chickadees’ ranges has received much attention (e.g., Brewer
1963, Rising 1968, Johnston 1971, Braun and Robbins 1986, Gill
et al. 1989). Everyone recognizes that there are substantial dif¬
ferences between Mountain and Black-capped Chickadees; no¬
body has seriously suggested lumping them. However, there is
definite disagreement over whether Carolina and Black-capped
Chickadees are really sufficiently different to be regarded as sepa¬
rate species or just races of a single species. This question is not
easily resolved.
Many of the existing data have been interpreted as supporting
each side of the debate. The breeding ranges of Carolina and
Black-capped Chickadees are for the most part contiguous but
nonoverlapping; indeed, actual narrow gaps often exist between
the two ranges where no chickadee breeds. Those favoring two
separate species suggest that these gaps may be the result of in¬
terspecific competitive exclusion (Tanner 1952). Others, who dis¬
pute the two-species theory, see the gaps as merely habitat of such
poor quality that no chickadee would settle and breed there (Rob¬
bins et al. 1986).
Whatever the causes of these narrow gaps, contact is made in
several places within the midwestern states (from Kansas east to
Illinois) and in the southern Appalachians, where interbreeding
apparently is fairly regular. In such contact zones, it is not unusual
to find chickadees that can sing the two-note fee-bee of Black-
capped Chickadees, the four-note fee-bee, fee-bay of Carolina
Chickadees, and several intermediate whistled vocalizations as well
(Johnston 1971, Merritt 1981; see Fig. 1-3). Ward and Ward (1974)
10 The Black-capped Chickadee

Seconds

Fig. 1-3. Tracings of spectrograms of Missouri chick¬

adee vocalizations, showing a normal Black-capped
Chickadee song (a); a normal Carolina Chickadee song
(e); and two-, three-, and four-note songs, all sung by a
single individual, of unknown affinities, from the contact
zone (b—d). Kilohertz scale is logarithmic. (From Robbins
et al. 1986, courtesy of Auk.)

interpreted this apparent convergence in song pattern as evidence

of interspecific competition between the two kinds of chickadees,
suggesting it represents vocal mimicry between members of two
different species. There is much to support the Wards’ views.
Whistled songs can be involved in repelling rivals (Ficken 1981b).
Moreover, chickadees do imitate; in fact, each young Black-
capped Chickadee probably must learn (by imitation) what to
whistle (Ficken 1981b). When Donald E. Kroodsma played taped
whistled songs to mixed groups of hand-reared nestling Black-
capped and Carolina Chickadees (all from nests well outside the
contact zone), he found that each kind readily learned to imitate
elements of the other’s whistled songs (D. E. Kroodsma, pers.
 General Natural History 11

comm.). Furthermore, a chickadee may be able to continue to add

new notes it hears to its repertoire throughout its life. A chickadee
in the contact zone would hear both songs, and could well experi¬
ment with various combinations of the notes, using whatever com¬
bination is most effective in driving off any given rival. With such
imitative abilities, a chickadee’s song should not, as Robbins et al.
(1986) seem to claim, necessarily indicate the genetic makeup of
the singer.
Black-capped and Carolina Chickadees differ behaviorally in
many ways. For example, their precopulatory vocalizations, by
both males and females, are very different (see Copulation, Chap¬
ter 5). Another area of divergence is their winter social organiza¬
tion: Carolina Chickadee flocks can have markedly less stable
dominance hierarchies than those of typical Black-capped Chick¬
adee flocks, and not infrequently have unbalanced sex ratios, even
when first formed (Mostrom 1988), a decidedly unusual phe¬
nomenon in Black-capped Chickadee flocks over most of their
range (see Flock Formation and Flock Composition, Chapter 7). In
addition, the mechanisms underlying whistled song production in
the two chickadees may be quite different. In the whistled fee-bee
of Black-capped Chickadees, relative pitch seems to be far more
important than the absolute pitch of the song; that is, a bird may
show considerable variability in how high the first note of a song is,
but far less variability in the pitch interval between adjacent notes
(Weisman et al. 1990). By contrast, Carolina Chickadees seem to
rely much more on absolute pitch of their songs (Lohr et al. 1991).
Such behavioral differences, while not constituting proof, none¬
theless support the idea that the two chickadees are different spe¬
cies.
Recently, an entirely new approach, involving biochemical analy¬
sis, has been used. One such method entails the analysis of tissue
samples for protein (and, more specifically, enzyme) similarity.
The enzymes studied are referred to variously as allozymes, iso¬
zymes, or electromorphs. The idea behind this analysis is that the
structure of these enzymes can give information concerning the
genetic material that codes for the enzymes’ manufacture within
the cells. The greater the difference in the allozymes, the greater
the genetic difference, and hence the more likely the samples are
from two different species. Conversely, if few or no differences are
found, the samples may represent only one species.
While the reasoning seems clear, there are evidently many pit-
 12 The Black-capped Chickadee

falls associated with interpretation and analysis of the data. Results

can vary depending on which enzymes were examined and even
on which computer program was used (e.g., Gill et al. 1989,
Springer and Krajewski 1989). Nevertheless, although an early
study (Braun and Robbins 1986) indicated that there was virtually
no difference between Black-capped and Carolina Chickadees, a
somewhat more extensive analysis by Gill et al. (1989), involving
most of the same allozymes and analyzed by the same computer
program, suggested fairly substantial differences between the two.
Indeed, this later study seems to indicate that the Black-capped
Chickadee is genetically closer to the Mountain Chickadee and
even the Mexican Chickadee than to the Carolina Chickadee, and
that it is closer to the Chestnut-backed and Boreal Chickadees
than it is to the Eurasian Willow and Marsh Tits (Fig. 1-4).
Using a somewhat different approach, Mack et al. (1986) exam¬
ined nucleotide sequences in the mitochondrial DNA of Black-
capped and Carolina Chickadees and found considerable dif-

Black-capped Chickadee
t - Mountain Chickadee
- Mexican Chickadee
- Chestnut-backed Chickadee
Boreal Chickadee
—I- Marsh 1
II - Willow Tit
I - Carolina Chickadee
Crested Tit

- Tufted Titmouse
+--//-+ +—+—+—+—+—+—+—+—+—+—+—+—+—+—+—+
0 . 10 .15 .20 .25 .30 .35 .40 .45 .50

Distance from root

Fig. 1-4. Genetic relationships among chickadees and titmice of the subgenus Poecil
portrayed as a distance Wagner tree. Black rectangles indicate shared, derived alleh
(suggesting closeness); pepper rectangles indicate alleles unique to species (suggestin
distance). According to these data, the Black-capped Chickadee is more closely related t
Chestnut-backed and Boreal Chickadees than to either Willow Tits or Carolina Chic!
adees. This tree is a subset of a fuller set of species rooted with the nuthatch genus Sitti
Total length of the full tree = 2.86; percent SD = 10.99. (From Gill et al. 1989, courtesy (
Wilson Bulletin.)
 General Natural History 13

ferences between the two. In light of this study and the one by
Grill et al., and with a note of caution based on Braun and Rob¬
bins’s data and the fact that hybridization undoubtedly occurs, I
will follow the lead of the latest (1983) AOU checklist, and treat
these two chickadees as closely related but distinct species.

Subspecies

The Black-capped Chickadee is nonmigratory, and its enormous

geographic range encompasses a wide variety of climates and hab¬
itats. For these reasons, Black-capped Chickadees show marked
local variation in size and, to some extent, plumage. Where recog¬
nizably different populations occur, especially if they seem to show
little geographical overlap, they are often described as subspecies.
There is some disagreement as to how many valid subspecies of
Black-capped Chickadees there actually are. For example, Snow
(1967) recognized only seven subspecies, although both Duvall
(1945) and the 1957 AOU checklist (the last one to include sub¬
species) recognized more. The nine subspecies of Black-capped
Chickadees listed by the 1957 AOU checklist (see Fig. 1-5) are
described below.
The eastern black-capped chickadee, Parus atricapillus atricapillus
Linnaeus, ranges from west of the Great Lakes to the Atlantic, and
from James Bay in the north down to central Missouri and north¬
ern New Jersey in the south. The back is olive-gray, and the wings
have broad white edgings on the secondaries and wing coverts.
The flanks are strongly tinged with buff, especially in early fall.
Measurements (from Duvall 1945): adult males (36 specimens),
wing 60-67.5 mm (average, 65.1 mm), tail 58.5-66 mm (average,
62.1 mm); adult females (31 specimens), wing 60-67 mm (63.4
mm), tail 57-64 mm (60.5 mm). See also Glase 1973 and Gochfeld
(1977 for further measurement data.
The Appalachian black-capped chickadee, P. a. practicus
Oberholser, lives from Ohio and southwestern Pennsylvania south
along the Appalachian Mountains to the Great Smokies. It is
somewhat smaller than the eastern black-capped chickadee and
has a shorter tail; the upper parts are darker and less green, and
the white edgings on the wing coverts and tail feathers are less
broad. Measurements (from Duvall 1945): adult males (12 spec¬
imens), wing 61.5-67 mm (64 mm), tail 57-61.5 mm (59.9 mm);
14 The Black-capped Chickadee

J Columbian Eastern

Rocky Mountain Newfoundland

Pallid Appalachian

Fig. 1-5. Breeding range of the Black-capped Chickadee, showing the dis¬
tribution of the nine subspecies recognized by the 1957 AOU checklist. Of
these, Snow (1967) recognized all but the Columbian and Rocky Mountain
forms, which he lumped with the long-tailed chickadee (Columbian) and
pallid chickadee (Rocky Mountain). The boundaries between the subspecies’
ranges are probably far less clear-cut than they appear here.
 General Natural History 15

adult females (12 specimens), wing 59.5-65.5 mm (63 mm), tail

57—60.5 mm (59.3 mm).
The Newfoundland black-capped chickadee, P. a. bartletti Alrich
and Nutt, is restricted to the islands of Newfoundland and Mi¬
quelon. It is darker and slightly larger than the eastern black-
capped chickadee, browner above, and has darker buff on the
sides and flanks. Measurements (from Duvall 1945): adult males
(7 specimens), wing 65-68 mm (66.4 mm); tail 59-64 mm (62.4
mm); adult females (6 specimens), wing 61-64.5 mm (62.3 mm),
tail 57.7—61.5 mm (60 mm).
The long-tailed chickadee, P. a. septentrionalis Harris, inhabits
west central Canada and central United States. It has markedly
longer wings and tail than the eastern black-capped chickadee,
and still broader white edgings on the wing and tail feathers.
Measurements (from Duvall 1945): adult males (62 specimens),
wing 64.5—73 mm (68.7 mm), tail 63—72.5 mm (67.4 mm); adult
females (30 specimens), wing 64-71 mm (67.2 mm), tail 64-71
mm (64.2 mm).
The Oregon chickadee, P. a. occidentalis Baird, is the West Coast
form of the Black-capped Chickadee, ranging from southwestern
British Columbia (excluding Vancouver Island) south to north¬
western California. It is a particularly distinctive form, being de¬
cidedly smaller than the eastern black-capped chickadee and con¬
siderably darker. The back tends to be dark gray-brown, and the
sides and flanks are much richer buff than in most other races,
especially in autumn and early winter. The white edgings on the
wings and tail are narrower than in many other races, also contrib¬
uting to the overall dark appearance of these western birds. Mea¬
surements (from Duvall 1945): adult males (41 specimens), wing
57.5-64 mm (61.2 mm), tail 53.5-60.5 mm (57.1 mm); adult
females (35 specimens), wing 57.5—63. 5 mm (60 mm), tail 53-59.5
mm (56 mm).
The pallid black-capped chickadee, P. a. nevadensis (Linsdale),
inhabits the Great Basin area, ranging south to northern Arizona
and Utah. The birds are about the same size as the long-tailed
chickadee, but are even paler and have broad white edgings on the
wing and tail feathers. Measurements (from Duvall 1945): adult
males (22 specimens), wing 64-72.2 mm (68 mm), tail 64-71.6
mm (67.5 mm); adult females (13 specimens), wing 63.5-70.5 mm
(66.9 mm), tail 63.5—70.5 mm (67 mm).
The Rocky Mountain black-capped chickadee, P. a. garrinus
16 The Black-capped Chickadee
Behle, inhabits much of the Rocky Mountain area of the United
States. It is a fairly large, buffy chickadee, about the same size as
the long-tailed chickadee, but with a brown back and strongly buff
rump. Both the wing and tail feathers have quite broad pale edg¬
ings. It was lumped by Snow (1967) with the Pallid black-capped
chickadee. Measurements (from Behle 1951): males (10 spec¬
imens), wing 64.3—70.6 mm (68.6 mm), tail 61.5—69.8 mm (66.7
mm); females (10 specimens), wing 63.4—76.7 mm (67.6 mm), tail
63.1-70.0 mm (65.6 mm).
The Columbian black-capped chickadee, P. a. fortuitus (Dawson
and Bowles), lives in southern British Columbia (except on the
coast) and in Washington and Oregon east of the Cascades. A
decidedly buffy race, it is very similar to the Rocky Mountain form,
but is somewhat smaller, and the back and rump are essentially the
same shade. Snow (1967) lumped it with the long-tailed chickadee.
Measurements (from Duvall 1945): adult males (27 specimens),
wing 60-67 mm (64.1 mm), tail 57.5-65 mm (62.1 mm); adult
females (23 specimens), wing 59—67 mm (63.2 mm), tail 57—65
mm (61.4 mm).
The Yukon chickadee, P. a. turneri Ridgway, inhabits Alaska,
extreme northwestern British Columbia, the Yukon and western
Northwest Territories north to the tree line. It is a fairly large, pale
form with a pale gray back, sides and flanks very pale buff, and
with very broad white edgings on the wing and tail feathers. Mea¬
surements (from Duvall 1945): adult males (20 specimens), wing
61-68 mm (65.4 mm), tail 62-70.5 mm (65.7 mm); adult females
(15 specimens), wing 61—66.5 mm (63.6 mm), tail 61—66 mm (63.7
mm).
 Some Study Techniques

This chapter discusses a few of the more common techniques

applied in the study of chickadees and titmice along with a few
that seem to have especially exciting potential for use in future
research.

Banding

Banding is essential for many areas of chickadee research. The

most common ways to catch chickadees are with mist nets (Fig.
2-1), often using playbacks of vocalizations to help attract chick¬
adees into the nets, and with wire traps (McCamey 1961). Very
young chickadees can be banded before they leave the nest, pref¬
erably when they are about ten days old; that is, old enough to
have legs the right size but too young for handling to cause early
fledging.
Much important information can be obtained through the use
of aluminum bands alone (see, for example, Loery and Nichols
1985, Loery et al. 1987). For many kinds of research questions,
however, it is necessary to be able to identify individuals from a
distance. Sight identification is typically achieved by the use of
auxiliary markers such as color bands. When each individual is
18 The Black-capped Chickadee

Fig. 2-1. A Black-capped Chickadee caught in a mist net. Mist nets are partic¬
ularly effective for catching every flock member, including those that are
relatively shy.

given its own set of colors, it can easily be distinguished by sight,

making further handling unnecessary.
Investigators have been using color bands on chickadees for
many years (e.g., Butts 1931; Odum 1941a, 1941b). Few ques¬
tioned the effect of this technique until Burley (1981) reported
that mate choice of Zebra Finches (Poephila guttata) can be influ¬
enced to a measurable extent by color bands: both males and
females apparently preferred to spend time with birds wearing
red bands rather than with birds wearing bands of any other color,
or even with unbanded Finches. Not too surprisingly, Burley’s data
caused quite a stir among researchers. Although Burley’s in¬
terpretation of her original data was challenged (Immelmann et
al. 1982), she and her colleagues later published additional data
that support her claims (Burley et al. 1982; but see also Ratcliffe
and Boag 1987). The fact that the bills of juvenile Zebra Finches
 Some Study Techniques 19

are black and those of adults are orange-red may explain the
apparent preference for red bands, red being a signal of sexual
maturity in this species. Similarly, Metz and Weatherhead (1991)
discovered that both red and black color bands, when placed on
male Red-winged Blackbirds (Agelaius phoeniceus), can act as social
signals.
Will a chickadee’s chances of obtaining a mate be influenced by
the colors it happens to be given when it is banded? For several
reasons, I suspect the answer is no. Unlike Zebra Finches, chick¬
adees do not have any natural color changes that signal sexual
maturity. Moreover, since much of pair formation occurs during
flock formation (see the section on pair formation in Chapter 5),
most chickadees will already be paired before they are banded.
Nevertheless, some pair formation does occur at other times of the
year. Because it remains to be proven that a chickadee’s color
bands have no effect whatsoever on its relations with other chick¬
adees, banders should randomize colors among birds.
When handling wild birds, banders should take precautions to
guard against ticks that might carry the spirochete that causes
Lyme disease, because several avian species are now known to
harbor carrier ticks (Anderson et al. 1990, Burgess 1990; see Para¬
sites and Diseases, Chapter 9).

Aging and Sexing Techniques

Chickadees that are less than a year old can be distinguished

from older birds relatively easily. Two methods can be used. One is
to check for skull pneumatization. As a chickadee develops, the
growing bones (in this case, those of the skull) become thicker and
typically come to include many very small pockets of gas; hence
the term pneumatization. Young chickadees’ darker, less pneu¬
matized skulls can easily be told apart from the paler, more pneu¬
matized skulls of the older birds. However, this technique is effec¬
tive only in late summer and early fall because most young
chickadees’ skulls are fully pneumatized by mid-October (Yunick
1980, 1981; Meigs et al. 1983). The other technique involves ex¬
amining the tail feathers. In autumn, when most banding is done,
juvenile chickadees’ tail feathers, which start to grow in the nest,
20 The Black-capped Chickadee

are actually older and thus more worn than those of adults, which
are grown during the postbreeding molt of late summer and early
fall. Owing in part to this differential wear, first-year chickadee tail
tips are more pointed and have less pale edging than those of
adults (Fig. 2-2). This difference, described for Black-capped
Chickadees by Meigs et al. (1983), is easily seen after a bit of
practice and can be used to age chickadees with well over 90 per¬
cent accuracy. Moreover, it remains useful throughout the fall and
early winter.
Sexing chickadees accurately presents more of a problem. A few
years ago Mosher and Lane (1972) suggested that males and
females could be told apart by the shape of their caps and bibs:
males, they thought, had bibs that were fairly broad just below the
bill, with poorly defined lower margins, while females’ bibs were

Fig. 2-2. The age of Black-capped Chickadees can be determined by examin¬

ing the tail feather (rectrix) tips, (a) Representative immature (first basic)
outer rectrix. The feather is pointed and worn, showing no white on the tip of
the inner vein, (b) Representative adult (basic) outer rectrix. The feather is
less worn than that of an immature and usually shows some white on both
edges of the feather tip. (Modified from Meigs et al. 1983.)
 Some Study Techniques 21
Table 2-1. Plumage categories of Black-capped Chickadees

Males Females

Bib shape Bib margin Cap margin Adult Juvenile Adult Juvenile

Truncate 6 1 5 2
Rough
Pointed 2 0 1 0
Wide
Truncate 11 2 2 3
Smooth
Pointed 8 3 6 0
Rough Truncate 4 1 5 1
Pointed 3 1 1 1
Pinched
Truncate 4 3 4 3
Smooth
Pointed 2 1 4 0

Statistical comparison (adult and juveniles combined)

Males Females Males Females
Wide 33 19 Rough 18 16
Pinched 19 19 Smooth 34 22
FEP2 > .30 FEP2 > .50
Males Females Males Females
Truncate 32 25 Wide-pointed 12 7
Pointed 20 13 Pinched-truncate 13 13
FEP2 > .30 FEP2 > .30

Source: Gochfeld 1977.

Note: Mid-April to mid-September plumages. FEP2 = Fisher exact probability test
(two-tailed, designated).

supposed to be narrower right under the bill and to have sharper,

more clear-cut lower margins. As for caps, they claimed that the
hind end, where the cap joins the back, was neatly squared in
females and more pointed in males (Mosher and Lane 1972). Gos-
ler and King (1989) found similar sex-related differences in bib
shape in a British population of Coal Tits (Parus ater). Unfortu¬
nately, this method, even if accurate in a small population of
Black-capped Chickadees, seems not to be of general use. The
careful work of Gochfeld (1977) showed that, while variability in
these characters does exist, it is not broadly correlated with sex
differences in this species (Table 2-1). No consistent differences
were found either between males and females or between adult
immature birds.
Several morphological measures can help determine the sex of a
22 The Black-capped Chickadee

chickadee in the hand once the subspecies is known; as shown at

the end of Chapter 1, the nine subspecies of the Black-capped
Chickadee exhibit markedly different measurements. One impor¬
tant measure is wing length. In general, males have longer wings
than do females (Glase 1973, Weise 1979). This indicator must be
used with caution, however, because chickadee wings get longer
with age (Blake 1956, Stewart 1963, Glase 1973). It is therefore ne¬
cessary to know a chickadee’s age before using wing length to de¬
termine its sex. Furthermore, in most populations there is at least
some overlap in wing length, so individuals with intermediate-
length wings must be sexed by some other means. Another mea¬
sure is tail length. Males typically have longer tails than do fe¬
males, although here again there is considerable overlap. Most
males also weigh more than most females—yet chickadee weights
(or mass) vary from season to season, and also with time of day
(Lawrence 1958), so both season and time must be standardized
before such data will be useful. Perhaps the best way to sex chick¬
adees is to use a combination of wing and tail length and weight.
Indeed, Desrochers (1990) found that by using a discriminant
analysis of these three measures, he could accurately sex over 90
percent of both males and females in his population.
Besides such morphological clues, various behavioral indices,
such as how often a bird gives gargle vocalizations, can also give
some indication of sex. Males are far more likely to give this call,
particularly in the nonbreeding season (Ficken, Ficken, and
Witkin 1978; Weise 1979).
Laparotomy, a surgical procedure in which a bird is anesthe¬
tized, cut open, and the gonads examined, can give unequivocal
data on the sex of birds in the hand (Risser 1971). Ketterson and
Nolan (1986) showed that laparotomized individuals of larger spe¬
cies, such as sparrows and juncos, had just as good long-term
survival as did unlaparotomized controls. Nevertheless, even these
experienced investigators reported that a few birds died while
they were improving their technique in giving the anesthetic and
in performing the operation itself. Chickadees, being quite a bit
smaller, would be still harder to laparotomize. Recently a new and
related technique, known as laparoscopy, was used on passerines
(Richner 1989). This process uses fiber optics, specifically a diag¬
nostic arthroscope, to observe the gonads of the birds. Richner
(1989) claimed that this technique is faster and less harmful than
laparotomy; however, it has yet to be tested on birds as small as
 Some Study Techniques 23
chickadees. Perhaps someday researchers (who can afford it) may
sex chickadees using some of the new high-technology methods of
DNA analysis. These include sex-specific DNA probes from blood
samples, a method recently developed for geese (Quinn et al.
1987, 1990), and flow cytometry (fluorescence-activated cell sort¬
ing), in which the DNA content of blood or other nucleated tissue
samples is compared with that of reference cells (Nakamura et al.
1990, Tiersch et al. 1991). Meanwhile, sexing chickadees on the
basis of behavioral observations, while taking longer to confirm,
seems preferable to subjecting them to the risks of tissue sampling,
laparotomy, or even laparoscopy, especially in studies where sur¬
vivorship is being measured.

Nest Boxes

Black-capped Chickadees do not readily accept nest boxes, nor

do the closely related Willow Tits (Parus montanus) of Europe. In
contrast, several other European titmice, such as Great (P. major)
and Blue (P. caeruleus) Tits, actually seem to prefer such nest sites
(Perrins 1979).
Nevertheless, a rather simple trick can markedly increase the
chances of a box’s being used by chickadees. Kluyver (1961) found
that nest boxes can be made far more acceptable by filling them
half-full of sawdust. The sawdust provides the chickadees with
something they can excavate. Kluyver used boxes that had en¬
trance holes of 3.0—3.5 cm (about llA inches) in diameter.
Because so many European tits use nest boxes readily, most of
the really interesting techniques involving boxes have been de¬
vised for these species; only a few of these techniques have been
applied to Black-capped Chickadees so far. Mechanical devices can
be attached that can record each time a bird enters or leaves the
nest (Fig. 2-3). Boxes with one or more glass sides have been
constructed, allowing inside activities to be monitored, either by a
video camera or by an observer in a blind close-by. The perch at
the entrance of a box can be rigged to a scale so that the weight of
the entering bird (plus any food it might be carrying) can be
measured (Perrins 1979). Nest boxes have even been converted
into metabolic chambers so that precise physiological measure¬
ments can be made of the occupants (Haftorn and Reinertsen
1985).
24 The Black-capped Chickadee

Fig. 2-3. A Black-capped Chickadee with food for its nestlings at a nest
box that has contact apparatus. The entering parent moves a hanging
strip, which makes the chronograph operate, yielding a record of each
time an adult enters or leaves the nest. (From Kluyver 1961, courtesy of
Auk.)
 Some Study Techniques 25

Grundel (1987, 1990) applied some of these techniques very

effectively to nesting Mountain Chickadees (Parus gambeli). He got
his chickadees to nest in cement nest boxes (see also Dahlsten and
Copper 1979) and then constructed three automated camera sys¬
tems around cement nesting boxes identical to those used by the
chickadees. Each was mounted with a movie camera and an elec¬
tronic flash. When a brood was ready to be recorded, he could
transfer it from the regular nest box to the camera box. A pho¬
tocell activated by a bird passing through the nest entrance trig¬
gered the camera shutter and flash. The camera then pho¬
tographed each bird as it entered (showing kind, number, and size
of food being carried) and again as it left. A watch mounted near
the entrance allowed precise timing data to be taken. Dots painted
on each parent’s forehead allowed each bird to be distinguished on
the film.
Several of the nest box techniques might profitably be applied to
nesting Black-capped Chickadees. Areas where nest boxes are
most likely to be accepted by chickadees include suburban settings:
places that have plenty of canopy to provide food for the offspring
but relatively few unpruned older trees, where natural cavities
might occur.

Ptilochronology

Ptilochronology, a technique recently pioneered by Thomas C.

Grubb, Jr., is based on the fact that as each feather grows, it
produces marks, termed growth bars, that reflect daily growth.
These bars are most visible in the larger feathers of the wings and
tail; they run more or less at right angles to the length of the
feather. There is somewhat of a knack to seeing these bars. It is
best to tilt the feather back and forth under a bright light to pick
them out; they will appear as a series of alternating darker and
paler bands. Evidently the darker bands are derived from material
laid down during the day, and the paler bands, from material laid
down during the night (Grubb 1989). Of greatest importance to
researchers is that the breadth of the growth bars apparently re¬
flects the bird’s nutritional status when the feather was being
grown.
Grubb (1989) suggested that researchers experimentally induce
the growth of new feathers to learn more about the nutritional
26 The Black-capped Chickadee

ecology of a species. One can capture a bird, mark it for future

identification, then pluck, label, and save one standard feather,
such as the right outer tail feather. After the bird has grown a
replacement feather (in about six weeks or so), recapture the bird,
collect the induced feather, and compare it with the one removed
earlier.
Ptilochronology has already been applied to parid ecology (see,
for example, Grubb and Cimprich 1990a, 1990b and Hogstad
1990b). It seems to be a particularly sensitive method for measur¬
ing the effect of various events or conditions on the nutritional
ecology of free-living birds.

Telemetry

One of the most exciting new techniques to be used on parids is

radio telemetry. Various kinds have been developed, some of
which have been used in studies of European tits.
Reinertsen and Haftorn (1983) implanted small (0.75-gram)
transmitters into the bodies (intraperitoneal cavities, to be precise)
of Willow Tits. This technique allowed them to obtain continuous
recordings of deep body temperatures of undisturbed birds. In¬
deed, their setup permitted continuous recordings for almost 500
hours. However, because the strength of the signal is not very
great, this kind of recording is best done on captive birds.
East and Hofer (1986) did a pilot study on radio-tracking free-
living Great Tits in England. They considered various methods of
attachment for the transmitters, including direct adhesion, har¬
nesses, and tail clips. Direct adhesion has the advantage of being
lighter than the other two, since the glue is lighter than either a
harness or a tail clip; nevertheless, East and Hofer rejected this
method because removal of the transmitter could result in dan¬
gerous reduction of the insulative properties of the feathers it had
been attached to. They avoided harnesses for fear of disrupting
the birds’ balance and because the harnesses might have become
tangled in vegetation. In the end they decided that the least dis¬
ruptive attachment was tail clips, placed close to the body on the
two or three central tail feathers.
East and Hofer, using special receiving equipment, found that
the signal from their birds was detectable at distances of up to 50
meters (almost 55 yards) in woodland habitat. The technique al-
 Some Study Techniques 27

lowed them to follow the movement of relatively undisturbed

birds, giving information on home range size and movement pat¬
terns. In fact, their data led them to speculate that standard obser¬
vations on free-living Great Tits may be somewhat biased by ob¬
server proximity. Although this is potentially an exciting and
important technique, some problems remain to be solved before it
can easily be applied to Black-capped Chickadees. One is weight.
The combination of East and Hofer’s radio transmitter plus the
tail dip weighed 1.8 grams. For a 29-30-gram bird such as the
Great Tit, this is well within the recommended limit of 10 percent
of a bird’s body mass (Brander and Cochran 1969, Amlaner and
MacDonald 1980). For a 10—14-gram bird such as a Black-capped
Chickadee, however, this weight is unacceptably high. Another
problem is that the batteries East and Hofer used in their transmit¬
ters had a mean life of only seven days. Still, as radio miniaturiza¬
tion and the production of superior batteries improves, the day
may not be far off when a transmitting device small enough to fit
without harm on a Black-capped Chickadee may be developed.
Indeed, researchers in Europe recently developed a 0.9-gram
transmitter that is suitable for use on Blue Tits (Arie J. van Noord-
wijk, pers. comm.). Moreover, the life of this transmitter is said to
be considerably longer than just one week. Transmitters this small
would seem to be perfectly suitable for use on Black-capped
Chickadees. If so, it may even now be possible to obtain accurate
data on the movements and behavior of undisturbed chickadees in
the field.
 3
Food and Feeding

One thing a chickadee must do every day of its independent life

is find and eat food. Some people assume that in particularly bad
weather chickadees do not feed at all. While it is true that they
might curtail their activity somewhat and stay in sheltered areas as
much as possible, they do have to eat—and the worse the weather,
the more they have to eat. Birds are warm-blooded animals, and
they need the energy provided by food to keep warm. Addi¬
tionally, small animals spend more energy in staying warm than do
large animals, because the smaller the individual, the greater its
ratio of surface area to volume. Put in practical terms, very small
creatures such as chickadees have a proportionally greater amount
of surface area (over which body heat can be lost) with respect to
the volume of warm core where heat can be stored. Various studies
on European titmice have shown that birds the size of Black-
capped Chickadees need approximately 10 kcal of energy each
day (Perrins 1979). One gram of stored fat yields about 9 kcal. But,
especially in bad winter weather, all fat reserves are typically de¬
pleted by the end of the long night, so the bird must get out the
next morning and forage for that day’s fuel.
Not only food but also water is essential for Black-capped Chick¬
adees (Fig. 3-1). Although chickadees must have water in some
 Food and Feeding 29

Fig. 3-1. Chickadees frequendy visit water to drink and to bathe. (© Alan
Cruickshank / VIREO.)

form throughout the year, probably their greatest need for water
comes with the summer’s heat. Simple metabolism in the summer
can produce about 0.24 kcal in just one hour, even in a resting
bird; flight adds to the amount of heat produced. Much of this
heat must be dissipated through evaporative cooling (Rising and
Hudson 1974). Clearly, the need for evaporative cooling is far
lower in cooler seasons, but water must still be obtained. During
the winter, Black-capped Chickadees can occasionally be seen eat¬
ing snow (Bent 1946).
30 The Black-capped Chickadee

Diet

Throughout the year, about 70 percent of Black-capped Chick¬

adees’ food is animal and 30 percent is vegetable matter (Bent
1946). These proportions vary with the season: most plant mate¬
rial is eaten in the fall and winter, and the chickadees’ typical
summer diet is almost entirely animal material. Martin et al. (1951)
reported that the winter diet of Black-capped Chickadees was ap¬
proximately 50 percent animal and 50 percent plant material,
while in the summer, 80—90 percent of their food was animal.
Plant matter eaten is mostly seeds and berries. Chickadees seem
particularly fond of small wax-covered berries such as those of
poison ivy (Rhus radicans) and bayberry (.Myrica sp.). In autumn
they also eat seed of ragweed (Ambrosia sp.), goldenrod (Solidago
sp.), and staghorn sumac (Rhus typhina), as well as other things such
as blueberries (Vaccinium sp.), blackberries {Rubus sp.), wild cherries
{Prunus sp.), and, where available, fruits of the tulip tree {Lirio-
dendron tulipifera) (Bent 1946, Brewer 1963, Baird 1980). In
winter, the bulk of their plant food is seeds of coniferous trees.
Hemlock (Tsuga sp.) seeds are particular favorites (Odum 1942a).
The two animal groups eaten most by Black-capped Chickadees
are lepidopterans (moths and butterflies, especially in the form of
caterpillars) and spiders (Robinson and Holmes 1982). Chickadees
will also eat beetles, true bugs, katydids, cicadas, plant lice, and
scale insects. They are even known to eat bees. Other animals,
such as centipedes and small snails and slugs, are sometimes also
eaten (Bent 1946).
On occasion, chickadees will actually eat some of the hairy cater¬
pillars so often rejected by most other insectivorous birds. These
include both gypsy moth {Limantria dispar) and tent caterpillars
{Malacosoma americanum). The only gypsy moth caterpillars I have
seen Black-capped Chickadees eating were very tiny, having not
yet dispersed from the egg patch. A pair of chickadees fed on the
emerging caterpillars, taking turns visiting the patch to get bill-
fulls. I have also seen chickadees repeatedly tearing open “tents” to
get at tent caterpillars. Although the caterpillars I have seen chick¬
adees eating were all an inch (2.5 cm) or less in length, reports
exist of Black-capped Chickadees eating even larger tent cater¬
pillars (Terres 1940).
Perhaps the most unexpected type of animal food that chick-
 Food and Feeding 31

adees eat naturally is animal fat. If a chickadee flock finds a dead

mammal, even one as large as a white-tailed deer (Odocoileus virgi-
nianus), the birds may peck through the skin to get at the sub¬
cutaneous fat, returning again and again until the source is de¬
pleted. Black-capped Chickadees are also known to eat dead
fish—very likely for the fat stored under the fish’s skin (Southern
1966). The largest group of chickadees that Hamerstrom (1942)
ever saw away from human habitation was feeding on the fat of a
dead skunk (.Mephites mephites). Similarly, Glase (1973) reported
aggregati°ns of more than one flock feeding on the fat of a dead
deer. Small wonder that chickadees visit suet feeders so readily!
Finally, at least in eastern North America, within the range of
sugar maple trees (Acer saccharum), chickadees show strong evi¬
dence of a fondness for sweet substances. In early spring when the
sap runs, icicles of maple sap can form where branches have bro¬
ken during the winter. Black-capped Chickadees regularly visit
these icicles, both to drink if the sap is melting (Fig. 3-2) and to eat
the frozen sap. Some hover below and snatch mouthfuls; some
actually hang from the larger sap icicles. I have watched a chick¬
adee knock a small sap icicle off a branch, catch it in midair, carry
it to a branch, and eat the whole thing. European titmice also show
evidence of similar taste preferences. Warren and Vince (1963)
found that Great Tits (Parus major) prefer sweet to salty, sour, or
bitter solutions, and Perrins (1979) reported that Blue Tits (P.
caeruleus) are known to visit hummingbird-type feeders in En¬
gland.
At certain times in their lives, chickadees may need extra
amounts of particular sorts of foods. One such time is during
nestling and early fledgling periods—that is, during posthatching
growth and development. Kluyver (1961), working in Mas¬
sachusetts, recorded 65 food items brought by parent Black-
capped Chickadees to their offspring. These were 35 caterpillars,
11 spiders, 10 small insect larvae, 6 termites, 1 white moth or
butterfly, 1 pupa, and 1 fly. The points to stress here are that all
were animal foods (high in protein to meet growth requirements)
and that almost 70 percent were larvae, which are usually softer
than adult insects, and thus easier to digest.
Another time of particular nutrient requirements occurs every
spring for breeding females. Eggs are costly to produce, being
packed with nutrients for the embryos’ development during in-
32 The Black-capped Chickadee

Fig. 3-2. A Black-capped Chickadee drinking sugar maple sap (photograph

© Marie Reed).

cubation. Females must obtain food that is particularly rich in

protein just before the onset of egg laying (see Factors That Affect
the Onset of Breeding, Chapter 5). Moreover, female chickadees
must also obtain enough calcium to form strong egg shells.
Females of some European tits are known to store bits of snail shell
in their gizzards during egg laying (Perrins 1979). A few reports
exist of Black-capped Chickadees eating small snails (Bent 1946),
and Ficken (1989) reported a case of Boreal Chickadees (Parus
hudsonicus) that repeatedly ate ash high in calcium (clearly prefer¬
ring that to ash low in calcium). It is not clear, however, how the
preference was related to egg laying because Ficken’s observations
were made in October, well past the breeding season. I know of no
study that focuses directly on how any chickadee species solves the
problem of obtaining sufficient calcium for egg-shell production.
This remains an interesting area for future research.
 Food and Feeding 33

Food Selection and Optimal Foraging

Optimal foraging models are based on the idea that an animal

collects food in a manner that will maximize its rate of energy
intake (Pyke et al. 1977, Krebs et al. 1978). The animals is assumed
to do this by discriminating among alternative sources of energy
(Grubb 1979). Foraging animals are expected to “figure” into the
equation not only rates of energy intake, but also rates of energy
expenditure (usually measured by human investigators in terms of
search time, pursuit time, and handling time) (Grubb 1979).
Foraging theory typically assumes a hierarchy of feeding deci¬
sions: foragers first choose among habitats, then among patches
within habitats, and finally among food items within patches (Ste¬
phens et al. 1986). One of the predictions of optimal patch choice
theory has to do with a forager’s giving up-time. The foraging bird
should leave a patch when its rate of food intake there has

Fig. 3-3. A graphical

representation of an optimal
foraging model of patch use. The
graphs show the number of prey
(or food) that survived in a patch
plotted against the time the
predator (or forager) spent
searching in the patch. The two
curves show the change in capture
rate with time spent searching in a
patch (assuming that the capture
rate within a patch is proportional
to the number of prey present).
The predator leaves a patch when
its capture rate drops to a critical UJ
level, called the giving-up TIME IN PATCH
threshold. This model predicts
that the giving-up threshold is
equal to the average rate of food
intake for the habitat. The giving-
up threshold is thus higher in the
rich habitat (upper graph) than in
the poor one. D, prey surviving
after predator has left; gp, good
patch; pp, poor patch. (From
Krebs et al. 1975, courtesy of
Animal Behaviour.)
34 The Black-capped Chickadee

dropped to the average rate for that habitat; therefore, the giving-
up time should be shorter in rich patches and relatively longer in
poorer ones (Fig. 3-3). The behavior of Black-capped Chickadees
foraging in aviaries fitted this prediction very well (Krebs et al.
1975): the birds’ giving-up time was inversely related to the aver¬
age capture rate of the environment (Table 3-1). In a study on
wild-caught Great Tits, Krebs et al. (1978) found that when given
two different patch types of differing profitability, the Great Tits
would first sample each one, then exploit the more profitable
patch. Although Plowright and Plowright (1987) criticized the in¬
terpretation of these data by Krebs et al., Kamil et al. (1988) later
demonstrated remarkably similar foraging behavior by Blue Jays
(Cyanocitta cristata).
Early versions of optimal foraging models predicted that forag¬
ing birds would never exhibit partial preferences; rather, for any
encounter with food, the response should be either always attack
or always ignore (Pyke et al. 1977). Later, however, several studies
(all on Great Tits) showed that at least this species does indeed
exhibit partial preferences, occasionally rejecting profitable food
and taking less profitable items. Why would these birds spend time
and energy capturing so-called nonoptimal kinds of food? One
reason is that, especially when dealing with cryptic prey, Great Tits
show some lag in their response to newly available food types
(Getty and Krebs 1985). Moreover, the actual value of a given sort
of food might change over time, in which case the “nonoptimal”
food types should be sampled periodically in case they have
changed in relative value (Rechten et al. 1983).

Table 3-1. The average giving-up times (for all patch types) of
four Black-capped Chickadees

Giving-up time (s)

Bird Rich environment Poor environment

YY 7.73 11.69
OW 8.11 15.57
BW 7.45 9.94
GW 6.98 8.30
Average 7.56* ± 0.24 (SD) 11.38* ± 1.56 (SD)

Source: Krebs et al. 1975, courtesy of Animal Behaviour.

*t = 2.4 (6 df); P < 0.025 (one-tailed).
 Food and Feeding 35

A theory related to optimal foraging is the optimal diet theory

(e.g., Stephens et al. 1986). Barkan and Withiam (1989) per¬
formed some interesting experiments on prey choice by captive
Black-capped Chickadees. They were concerned with what would
happen if two sorts of food were available—one more profitable
but smaller than the other. One version of the optimal diet model
predicts that a predator would maximize its long-term rate of food
intake by specializing on the larger but less profitable prey if the
encounter rate was sufficiently low. However, Barkan and With-
iam’s chickadees did not behave as this model predicts; they always
preferred the higher profitability food type, regardless of encoun¬
ter rate. Barkan and Withiam reasoned that their birds could have
been choosing food so as to either minimize pursuit time or max¬
imize profitability; evidently, from the results of their second ex¬
periment (Fig. 3-4) the chickadees chose food so as to maximize
profitability.
Apparently, foraging chickadees tend to prefer a sure thing
over a more uncertain one. Barkan (1990) demonstrated this pref¬
erence in the field with free-living Black-capped Chickadees. He
gave the birds a choice of two food types, each hidden in holes in a
feeder. The two types of food had equal expected values but dif¬
ferent variances. That is, the two food types had the same energy
content and were presented in equal total amounts (equal ex¬
pected values), but one kind (the low-variance type) was placed in
the feeder’s holes such that every hole contained the same amount
of food, while the other kind (the high-variance type) was dis¬
tributed such that some holes contained a large amount of food
and other holes contained nothing. The chickadees showed a
clear, significant preference for the low-variance food (that is, the
sure thing). In this regard, the chickadees’ behavior fits the predic¬
tions of still another variant on optimal foraging, known as the
risk-sensitive foraging theory (see, for example, Stephens and
Krebs 1986).
Lucas (1987) and others (e.g., Grundel 1990) have pointed out
that actual (as opposed to theoretical) foraging by birds in nature
is regularly influenced by a wide variety of constraints, each of
which may influence different birds in different ways. Some of
these constraints are predation (Lima 1985), rank and the pres¬
ence of more dominant individuals (Pierce and Grubb 1981,
Schneider 1984), and the individual’s need at that moment for the
expression of other sorts of behavior, such as defense of territory
36 The Black-capped Chickadee

Experiment

Profitability

Low High

io r

c
CD
CO
O
JZ
O
c
o
tr 0-5
o
CL
o

00

Pursuit Time

Fig. 3-4. A comparison of the proportion of each food type chosen by

four Black-capped Chickadees under two sets of conditions. The chick¬
adees consistently chose high-profitability food, regardless of pursuit time.
Error bars indicate ± 2 SE. (From Barkan and Withiam 1989, courtesy of
American Naturalist, © 1989 by The University of Chicago.)

(Ydenberg 1987). Clearly, chickadee food selection, at least that of

captive birds, does fit well with some of the predictions of optimal
foraging theory and some of its later derivatives. Nonetheless,
Lucas (1987) may well have been right when he concluded that
foraging by any wild bird is an extremely complex, dynamic pro¬
cess that may never fit very well with sweeping general models.

Foraging Niche

Exactly where Black-capped Chickadees feed can vary with the

individual, the weather, the season, and what is available in the
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probably arises from their having first written upon papyrus, after
the use of stone was laid aside, and only adopted skins upon their
embracing the Jewish religion. The Ethiopians, indeed, write upon
parchment, yet use the same form of books as we do. The outer
boards are made of wood and covered with leather. It was the law
only they say they were in use to preserve in one long roll of
parchment, upon the foreside of which it was written; it being
indecent and improper to write any part of it on the back, or a less
honourable place of the skin: And such was the roll we have just
mentioned as presented to Ptolemy, where such pains were taken in
joining the several skins together, for this very reason.

The manner paper was made has been controverted; but whoever
will read Pliny12 attentively, cannot, as I imagine, be long in doubt.
The thick part of the stalk being cut in half, the pellicle between the
pith and the bark, or perhaps the two pellicles, were stript off, and
divided by an iron instrument, which probably was sharp-pointed,
but did not cut at the edges. This was squared at the sides so as to
be like a ribband, then laid upon a smooth table or dresser, after
being cut into the length that it was required the leaf should be.
These stripes, or ribbands of papyrus, were lapped over each other
by a very thin border, and then pieces of the same kind were laid
transversely, the length of these answering to the breadth of the
first. The book which I have is eleven inches and a half long, and
seven inches broad, and there is not one leaf in it that has a ribband
of papyrus of two inches and a half broad, from which I imagine the
size of this plant, formerly being fifteen feet long, was pretty near
the truth. No such plant, however, appears now; I do not remember
to have ever seen one more than ten feet high. This is probably
owing to their being allowed to grow wild, and too thick together,
without being weeded; we know from Herodotus13, that the
Egyptians cut theirs down yearly as they did their harvest.

These ribbands, or stripes of papyrus, have twelve different names

in Pliny14, which is to be copious with a vengeance. They are,
philura, ramentum; scheda, cutis, plagula, corium, tænia,
subtegmen, statumen, pagina, tabula, and papyrus. After these, by
whatever name you call them, were arranged at right angles to each
other, a weight was placed upon them while moist, which
compressed them, and so they were suffered to dry in the sun.

It was supposed that the water of the Nile15 had a gummy quality
necessary to glue these stripes together. This we may be assured is
without foundation, no such quality being found in the water of the
Nile. On the contrary, I found it of all others the most improper, till it
had settled, and was absolutely divested of all the earth gathered in
its turbid state. I made several pieces of this paper, both in Abyssinia
and Egypt, and it appears to me, that the sugar or sweetness with
which the whole juice of this plant is impregnated, is the matter that
causes the adhesion of these stripes together, and that the use of
the water is no more than to dissolve this, and put it perfectly and
equally in fusion.

There seemed to be an advantage in putting the inside of the pellicle

in the situation that it was before divided, that is, the interior parts
face to face, one long-ways, and one cross-ways, after which a thin
board of the cover of a book was laid first over it, and a heap of
stones piled upon it. I do not think it succeeded with boiled water,
and it was always coarse and gritty with the water of the Nile. Some
pieces were excellent, made with water that had settled, that is, in
the state in which we drink it; but even the best of it was always
thick and heavy, drying very soon, then turning firm and rigid, and
never white; nor did I ever find one piece that would bear the
strokes of a mallet16, but in its greenest state the blow shivered and
divided the fibres length-ways; nor did I see the marks of any stroke
of a hammer or mallet in the book in my custody, which is certainly
on Saitic or Hieratic paper. I apprehend by a passage in Pliny17, that
the mallet was used only when artificial glue or gum was made use
of, which must have been as often as they let these stripes of the
ribband or pellicle dry before arranging them.
Pliny18 says, the books of Numa were 830 years old when they were
found, and he wonders, from the brittleness of the inside of the
paper, it could have lasted so long. The manuscript in my
possession, which was dug up at Thebes, I conjecture is near three
times the age that Pliny mentions; and, though it is certainly fragil,
has substance and preservation of letter enough, with good care, to
last as much longer, and be legible.

If the Saitic paper was, as we imagine, the first invented, it should

follow, contrary to what Isidore advances, that it was not first
invented in Memphis, but in Upper Egypt in Seide, whose language
and writing obtained in the earliest age, though Lucan seems to
think with Isidore,

Nondum flumineas Memphis contexere biblos

Noverat.——
Lucan, lib. iii.

After the hieroglyphics were lost, perhaps some time before, we

know nothing the Egyptians adopted so generally as paper, and
there were probably19 religious reasons that impeded in those early
days the people from falling upon the most natural, the skins of
beasts. However this be, it is certain under the Egyptians, naturally
averse to novelty and improvement, paper arrived to no great
perfection till taken in hands by the Romans. The Charta Claudia was
thirteen inches wide, the Hieratica, or Saitica, eleven, and such is
the length of the leaf of my book in the Saitic dialect, that is, the old
Coptic, or Egyptian of Upper Egypt. I have no idea what the
Emporetic paper was, which obtained that degree of coarseness and
toughness, as to serve for shopkeepers’ uses to tie up goods, unless
it was like our brown paper employed to the same purposes.

If the date of the invention of this useful art of making paper is

doubtful, the time when it was lost, or superseded by one more
convenient, is as uncertain. Eustathius says it was disused in his
time in the 1170. Mabillon endeavours to prove it existed in the 9th,
and even that there existed some Popish bulls wrote upon it as late
as the 11th century. He gives, as instances, a part of St Mark’s
Gospel preserved at Venice as being upon papyrus, and the
fragment of Josephus at Milan to be cotton paper, while Maffei
proves this to be just the reverse, that of St Mark being cotton, and
the other indisputably he thinks to be Egyptian papyrus, so that
Mabillon’s authority as to the bulls of the pope may be fairly
questioned.

The several times I have been at these places mentioned, I have

never succeeded in seeing any of these pieces; that of St Mark at
Venice I was assured had been recognized to be cotton paper; it was
rendered not legible by the warm saliva of zealots kissing it from
devotion, which I can easily comprehend must contain a very
corrosive quality, and the Venetians now refuse to shew it more. I
have seen two detached leaves of papyrus, but do not believe there
is another book existing at the present time but that in my
possession, which is very perfect. I gave Dr Woide leave to translate
it at Lord North’s desire; it is a gnostic book, full of their dreams.

The general figure of this plant Pliny has rightly said to resemble a
Thyrsus; the head is composed of a number of small grassy
filaments, each about a foot long. About the middle, each of these
filaments parts into four, and in the point, or partition, are four
branches of flowers; the head of this is not unlike an ear of wheat in
form, but which in fact is but a chaffy, silky, soft husk. These heads,
or flowers, grow upon the stalk alternately, and are not opposite to,
or on the same line with each other at the bottom.

Pliny20 says it has no seed; but this we may be assured is an

absurdity. The form of the flower sufficiently indicates that it was
made to resolve itself into the covering of one, which is certainly
very small, and by its exalted situation, and thickness of the head of
the flower, seems to have needed the extraordinary covering it has
had to protect it from the violent hold the wind must have had upon
it. For the same reason, the bottom of the filaments composing the
head are sheathed in four concave leaves, which keep them close
together, and prevent injury from the wind getting in between them.

The stalk is of a vivid green, thickest at the bottom, and tapering up

to the top21; it is of a triangular form. In the Jordan, the single side,
or apex of the triangle, stood opposed to the stream as the cut-
water of a boat or ship, or the sharp angle of a buttress of a bridge,
by which the pressure of the stream upon the stalk would be greatly
diminished. I do not precisely remember how it stood in the lakes in
Ethiopia and Egypt, and only have this remark in the notes I made
at the Jordan.

This construction of the stalk of the papyrus seems to reproach

Aristotle with want of observation. He says that no plant had either
triangular or quadrangular stalks. Here we see an instance of the
contrary in the papyrus, whose stalk is certainly and universally
triangular; and we learn from Dioscorides that many more have
quadrangular stalks, or stems of four angles.

It has but one root, which is large and strong22, Pliny says, as thick
as a man’s arm: So it was, probably, when the plant was fifteen feet
high, but it is now diminished in proportion, the whole length of the
stalk, comprehending the head, being a little above ten, but the root
is still hard and solid near the heart, and works with the turning
loom tolerably well, as it did formerly when they made cups of it. In
the middle of this long root arises the stalk at right angles, so when
inverted it has the figure of a T, and on each side of the large root
there are smaller elastic ones, which are of a direction perpendicular
to it, and which, like the strings of a tent, steady it and fix it to the
earth at the bottom. About two feet, or little more, of the lower part
of the stalk is cloathed with long, hollow, sword-shaped leaves,
which cover each other like scales, and fortify the foot of the plant.
They are of a dusky brown, or yellow colour. I suppose the stalk was
cut off below, at about where these leaves end.
The drawing represents the papyrus as growing. The head is not
upright, but is inclined, as from its size it always must be in hot
countries, in which alone it grows. In all such climates, there is some
particular wind that reigns longer than others, and this being always
the most violent, as well as the most constant, gives to heavy-
headed trees, or plants, an inclination contrary to that from which it
blows.

This plant is called el Berdi in Egypt, which signifies nothing in

Arabic, and I suppose is old Egyptian. I have been told by a learned
gentleman23, that in Syria it is known by the name of Babeer, which
approaches more to the sound of papyrus, and paper; this I never
heard myself, but leave it entirely upon his authority.
 BALESSAN, BALM, or BALSAM.

T HE great value set upon this drug in the east remounts to very
early ages; it is coeval with the India trade for pepper, and the
beginning of it consequently lost in the darkness of the first ages.
We know from scripture, the oldest history extant, as well as most
infallible, that the Ishmaelites, or Arabian carriers and merchants,
trafficking with the India commodities into Egypt, brought with them
balm as part of the cargo with pepper; but the price that they paid
for Joseph was silver, and not a barter with any of their articles of
merchandise.

Strabo alone, of all the ancients, hath given us the true account of
the place of its origin, “Near to this, that historian says, is the most
happy land of the Sabeans, and they are a very great people.
Among these, frankincense, myrrh, and cinnamon grow, and in the
coast that is about Saba the balsam also.” Among the myrrh-trees
behind Azab all along the coast to the Straits of Babelmandeb is its
native country. It grows to a tree above fourteen feet high,
spontaneously and without culture, like the myrrh, the coffee, and
frankincense tree; they are all equally the wood of the country, and
are occasionally cut down and used for fuel. We need not doubt but
that it was early transplanted into Arabia, that is, into the south part
of Arabia Felix, immediately fronting Azab, the place of its nativity.
The high country of Arabia was too cold to receive it, being all
mountainous; water freezes there.
 Balessan

London Publish’d Decr. 1.st 1789 by G. Robinson & Co.

 Balessan.

There is an anecdote relating to Sir William Middleton, who was

surprised and taken prisoner by the Turks in the first attempt to
open the trade of the Red Sea, that when about to set24 out for
Sanaa, corruptly called Zenan, the residence of the Imam, or prince
of Arabia Felix, he was by the people desired25 to take his fur cloak
along with him to keep him from the cold; he thought they were
ridiculing him upon what he had to suffer from the approaching
heat, which he was convinced in the middle of Arabia must be
excessive.
The first plantation that succeeded seems to have been at Petra, the
ancient metropolis of Arabia, now called Beder, or Beder Hunein,
whence I got one of the specimens from which the present drawing
is made.

Josephus26, in the history of the antiquities of his country, says, that

a tree of this balsam was brought to Jerusalem by the queen of
Saba, and given, among other presents, to Solomon, who, as we
know from scripture, was very studious of all sort of plants, and
skilful in the description and distinction of them. Here it seems to
have been cultivated and to have thriven, so that the place of its
origin came to be forgotten.

Notwithstanding this positive authority of Josephus, and the great

probability that attends it, we are not to put it in competition with
what we have been told from scripture, as we have just now seen,
that the place where it grew, and was sold to merchants, was Gilead
in Judea, more than 1730 years before Christ, or 1000 before the
queen of Saba; so that reading the verse, nothing can be more plain
than that it had been transplanted into Judea, flourished, and had
become an article of commerce in Gilead long before the period
Josephus mentions: “And they sat down to eat bread, and they lifted
up their eyes and looked, and behold, a company of Ishmaelites
came from Gilead with their camels, bearing spicery, and balm, and
myrrh, going to carry it down to Egypt27.” Now, the spicery, or
pepper, was certainly purchased by the Ishmaelites at the mouth of
the Red Sea, where was the market for Indian goods, and at the
same place they must have bought the myrrh, for that neither grew
nor grows any where else than in Saba or Azabo east to Cape
Gardefan, where were the ports for India, and whence it was
dispersed all over the world.

The Ishmaelites, or Arabian carriers, loaded their camels at the

mouth of the Red Sea with pepper and myrrh. For reasons not now
known to us, they went and completed their cargo with balsam at
Gilead, so that, contrary to the authority of Josephus, nothing is
more certain, than 1730 years before Christ, and 1000 years before
the queen of Saba came to Jerusalem, the balsam-tree had been
transplanted from Abyssinia into Judea, and become an article of
commerce there, and the place from which it originally was brought,
through length of time, combined with other reasons, came to be
forgotten.

Theophrastus, Dioscorides, Pliny, Solinus,and Serapion, all say that

this balsam came only from Judea. The words of Pliny are, “But to
all other odours whatever, the balsam is preferred, produced in no
other part but the land of Judea, and even there in two gardens
only; both of them belonging to the king, one no more than twenty
acres, the other still smaller28.”

At this time I suppose it got its name of Balsamum Judaicum, or,

Balm of Gilead, and thence became an article in merchandise and
fiscal revenue, which probably occasioned the discouragement of
bringing it any more from Arabia, whence it very probably was
prohibited as contraband. We shall suppose thirty acres planted with
this tree would have produced more than all the trees in Arabia do
at this day. Nor does the plantation of Beder Hunein amount to
much more than that quantity, for we are still to observe, that even
when it had been as it were naturalised in Judea, and acquired a
name in the country, still it bore evident marks of its being a
stranger there; and its being confined to two royal gardens alone,
shews it was maintained there by force and culture, and was by no
means a native of the country. And this is confirmed by Strabo, who
speaks of it being in the king’s palace or garden at Jericho. This
place being one of the warmest in Judea, shews likewise their
apprehensions about it, so that in Judea, we may imagine it was
pretty much in the state of our myrtles in England, which, though
cultivated in green-houses in all the rest of the island, yet grow
beautifully and luxuriantly in Devonshire and Cornwall, the western
parts of it.
Diodorus Siculus says, it grew in a valley in Arabia Felix; he should
have said on a number of gentle, sloping hills in Arabia Deserta,
which have a very small degree of elevation above the plain, but by
no means resemble a valley. This place was the scene of three
bloody battles between Mahomet and his kinsmen the Beni Koreish,
who refused to be converts to his religion, or acknowledge his divine
legation. These are at large described by several of the historians of
that nation, with circumstances and anecdotes, as well interesting
and entertaining, as elegantly told. They shew plainly that
Mahomet’s tribe, the Beni Koreish, did not receive their fanatical
manners and disposition from Mahomet and his religion, but were
just as obstinate, ignorant, and sanguinary when they were Pagans,
as they were afterwards when converted and became Mahometans.
The last of these battles, which was decisive in Mahomet’s favour,
gave him the sovereignty of Mecca, and was attended with the
extirpation of some of the principal families in this tribe.

At this time the balsam is supposed, by being sold in Judea, and not
accessible by reason of the commotions in Arabia, to have become
almost forgotten in that last part, where the trade from Abyssinia, its
native country, was likewise interrupted by this innovation of
religion, and by Mahomet’s profanation of the Caaba, or temple of
the sun, the ancient resort of the Sabean merchants carrying on the
trade of India. This interval the impostor thought proper for a
pretended miracle; he said, that, from the blood of the Beni Koreish
slain, there had sprung up this grove of trees, from the juice of
which all the true believers on his side received a cure for their
wounds, however fatal they appeared, nay, some of them were
revived from even death itself. Since that time it has maintained its
reputation equal to that which it had in antiquity.

Prosper Alpinus says, that one Messoner a eunuch, governor of Cairo

in the year 1519, caused bring from Arabia forty plants, which he
placed in the garden of Mattareah, where he superintended them.
Every day he went to that garden to pay his devotions to the Virgin
Mary. It was many times renewed, and has as often perished since.
Bellonius says, that in his time there were ten plants at Mattareah,
and he is of opinion, that in all ages they grew well in Arabia, which
is not true, for those at Beder are constantly supplied with new
plants so soon as the old ones decay. There was none existing at
Mattareah the two several times I visited Cairo, but there were some
of the Christians still living there that remembered one plant in that
garden.

There were three productions from this tree very much esteemed
among the ancients. The first was called Opobalsamum, or, Juice of
the Balsam, which was the finest kind, composed of that greenish
liquor found in the kernel of the fruit: The next was Carpo-
balsamum, made by the expression of the fruit when in maturity.
The third was Xylo-balsamum, the worst of all, it was an expression
or decoction of the small new twigs of a reddish colour. These twigs
are still gathered in little faggots and sent to Venice, where I am told
they are an ingredient in the Theriac, or of some sort of compound
drug made in the laboratories there: But the principal quantity of
balsam in all times was produced by incision, as it is at this day.
Concerning this, too, many fables have been invented and
propagated.

Tacitus says, that this tree was so averse to iron that it trembled
upon a knife being laid near it, and some pretend the incision should
be made by ivory, glass, or stone. There is no doubt but the more
attention there is given to it, and the cleaner the wound is made, the
better this balsam will be. It is now, as it probably ever has been,
cut by an ax, when the juice is in its strongest circulation in July,
August, and beginning of September. It is then received into a small
earthen bottle, and every day’s produce gathered and poured into a
larger, which is kept closely corked. The Arabs Harb, a noble family
of Beni Koreish, are the proprietors of it, and of Beder, where it
grows. It is a station of the Emir Hadje, or pilgrims going to Mecca,
half way between that city and Medina.
Some books speak of a white sort brought by the caravans from
Mecca, and called Balsam of Mecca, and others a balsam called that
of Judea, but all these are counterfeits or adulterations. The balsam
of Judea, which I have already mentioned, was long ago lost, when
the troubles of that country withdrew the royal attention from it;
but, as late as Galen’s time, it not only existed, but was growing in
many places of Palestine besides Jericho, and there is no doubt but
it is now totally lost there.

When Sultan Selim made the conquest of Egypt and Arabia in the
year 1516, three pound was then the tribute ordered to be sent to
Constantinople yearly, and this proportion is kept up to this day. One
pound is due to the governor of Cairo, one pound to the Emir Hadje
who conducts the pilgrims to Mecca, half a pound to the basha of
Damascus, and several smaller quantities to other officers, after
which, the remainder is sold or farmed out to some merchants, who,
to increase the quantity, adulterate it with oil of olives and wax, and
several other mixtures, consulting only the agreement of colour,
without considering the aptitude in mixing; formerly we were told it
was done with art, but nothing is easier detected than this fraud
now.

It does not appear to me, that the ancients had ever seen this plant,
they describe it so variously; some will have it a tree, some a shrub,
and some a plant only; and Prosper Alpinus, a modern, corroborates
the errors of the ancients, by saying it is a kind of vine, (viticosus).
The figure he has given of it is a very bad one, and leaves us
entirely in doubt in what class to place it. The defect of the plant in
Judea and in Egypt, and the contradiction in the description of the
ancients as to its figure and resemblance, occasioned a doubt that
the whole plants in these two countries, and Arabia also, had been
lost in the desolation occasioned by the Mahometan conquest; and a
warm dispute arose between the Venetians and Romans, whether
the drug used by the former in the Theriac was really and truly the
old genuine opobalsamum? The matter was referred to the pope,
who directed proper inquiry to be made in Egypt, which turned out
entirely in favour of the Venetians, and the opobalsamum continuing
as formerly.

A very learned and tedious treatise was published by Veslingius, in

the year 1643, at Padua, where this affair was discussed at full
length. As both parties of the disputants seem to argue concerning
what it is from the misunderstood reports of what it was, I shall
content myself briefly with stating what the qualities of the
opobalsamum are, without taking pains to refute the opinions of
those that have reported what the opobalsamum is not.

The opobalsamum, or juice flowing from the balsam-tree, at first

when it is received into the bottle or vase from the wound from
whence it issues, is of a light, yellow colour, apparently turbid, in
which there is a whitish cast, which I apprehend are the globules of
air that pervade the whole of it in its first state of fermentation; it
then appears very light upon shaking. As it settles and cools, it turns
clear, and loses that milkiness which it first had when flowing from
the tree into the bottle. It then has the colour of honey, and appears
more fixed and heavy than at first. After being kept for years, it
grows a much deeper yellow, and of the colour of gold. I have some
of it, which, as I have already mentioned in my travels, I got from
the Cadi of Medina in the year 1768; it is now still deeper in colour,
full as much so as the yellowest honey. It is perfectly fluid, and has
lost very little either of its taste, smell, or weight. The smell at first is
violent and strongly pungent, giving a sensation to the brain like to
that of volatile salts when rashly drawn up by an incautious person.
This lasts in proportion to its freshness, for being neglected, and the
bottle uncorked, it quickly loses this quality, as it probably will at last
by age, whatever care is taken of it.

In its pure and fresh state it dissolves easily in water. If dropt on a

woollen cloth, it will wash out easily, and leaves no stain. It is of an
acrid, rough, pungent taste, is used by the Arabs in all complaints of
the stomach and bowels, is reckoned a powerful antiseptic, and of
use in preventing any infection of the plague. These qualities it now
enjoys, in all probability, in common with the various balsams we
have received from the new world, such as the balsam of Tolu, of
Peru, and the rest; but it is always used, and in particular esteemed
by the ladies, as a cosmetic: As such it has kept up its reputation in
the east to this very day. The manner of applying it is this; you first
go into the tepid bath till the pores are sufficiently opened, you then
anoint yourself with a small quantity, and, as much as the vessels
will absorb; never-fading youth and beauty are said to be the
consequences of this. The purchase is easy enough. I do not hear
that it ever has been thought restorative after the loss of either.

The figure I have here given of the balsam may be depended upon,
as being carefully drawn, after an exact examination, from two very
fine trees brought from Beder Hunein; the first by the Cadi of
Medina at Yambo; the second at Jidda, by order of Yousef Kabil, vizir
or minister to the sherriffe of Mecca. The first was so deliberately
executed, that the second seemed of no service but to confirm me in
the exactitude of the first. The tree was 5 feet 2 inches high from
where the red root begins, or which was buried in the earth, to
where it divides itself first into branches. The trunk at thickest was
about 5 inches diameter, the wood light and open, and incapable of
polishing, covered with a smooth bark of bluish-white, like to a
standard cherry-tree in good health, which has not above half that
diameter; indeed a part of the bark is a reddish brown; it flattens at
top like trees that are exposed to snow-blasts or sea-air, which gives
it a stunted appearance. It is remarkable for a penury of leaves. The
flowers are like that of the acacia-tree, white and round, only that
three hang upon three filaments, or stalks, where the acacia has but
one. Two of these flowers fall off and leave a single fruit; the
branches that bear this are the shoots of the present year; they are
of a reddish colour, and tougher than the old wood: it is these that
are cut off and put into little faggots, and sent to Venice for the
Theriac, when bruised or drawn by fire, and formerly these made
the Xylo-balsamum.
Concerning the vipers which, Pliny says, were frequent among the
balsam trees I made very particular inquiry; several were brought
me alive, both to Yambo and Jidda. Of these I shall speak in another
place, when I give the figure, and an account of that animal so
found.
 SASSA, MYRRH, and OPOCALPASUM.

A T the time when I was on the borders of the Tal-Tal, or

Troglodyte country, I sought to procure myself branches and
bark of the myrrh-tree, enough preserved to be able to describe it
and make a design; but the length and ruggedness of the way, the
heat of the weather, and the carelessness and want of resources of
naked savages always disappointed me. In those goat-skin bags into
which I had often ordered them to put small branches, I always
found the leaves mostly in powder; some few that were entire
seemed to resemble much the acacia vera, but were wider towards
the extremity, and more pointed immediately at the end. In what
order the leaves grew I never could determine. The bark was
absolutely like that of the acacia vera; and among the leaves I often
met with a small, straight, weak thorn, about two inches long.

These were all the circumstances I could combine relative to the

myrrh-tree, too vague and uncertain to risk a drawing upon, when
there still remained so many desiderata concerning it; and as the
king was obstinate not to let me go thither after what had happened
to the surgeon’s mate and boat’s crew of the Elgin Indiaman29, I was
obliged to abandon the drawing of the myrrh-tree to some more
fortunate traveller, after having in vain attempted to procure it at
Azab, as I have already mentioned.

At the same time that I was taking these pains about the myrrh, I
had desired the savages to bring me all the gums they could find,
with the branches and bark of the trees that produced them. They
brought me at different times some very fine pieces of incense, and
at another time a very small quantity of a bright colourless gum,
sweeter on burning than incense, but no branches of either tree,
though I found this latter afterwards in another part of Abyssinia.
But at all times they procured me quantities of gum of an even and
close grain, and of a dark brown colour, which was produced by a
tree called Sassa, and twice I received branches of this tree in
tolerable order, and of these I made a drawing.

Some weeks after, while walking at Emfras, a Mahometan village,

whose inhabitants are myrrh merchants, I saw a large tree with the
whole upper part of the trunk, and the large branches, so covered
with bosses and knobs of gum, as to appear monstrously deformed,
and inquiring farther about this tree, I found that it had been
brought, many years before, from the myrrh country, by merchants,
and planted there for the sake of its gum, with which these
Mahometans stiffened the blue Surat cloths they got damaged from
Mocha, to trade in with the Galla and Abyssinians. Neither the origin
of the tree which they called Sassa, nor the gum, could allow me to
doubt a moment that it was the same as what had been brought to
me from the myrrh country, but I had the additional satisfaction to
find the tree all covered over with beautiful crimson flowers of a very
extraordinary and strange construction. I began then a drawing
anew, with all that satisfaction known only to those who have been
conversant in such discoveries.
 Sassa

London Publish’d Dec.r 1.st 1789 by G. Robinson & Co.

 Sassa

London Publish’d Dec.r 1.st 1789 by G. Robinson & Co.

I took pieces of the gum with me; it is very light. Galen complains
that, in his time, the myrrh was often mixed with a drug which he
calls Opocalpasum, by a Greek name, but what the drug was is
totally unknown to us at this day, as nothing similar to the Greek
name is found in the language of the country. But as the only view
of the savage, in mixing another gum with his myrrh, must have
been to increase the quantity, and as the great plenty in which this
gum is produced, and its colour, make it very proper for this use,
and above all, as there is no reason to think there is another gum-
bearing tree of equal qualities in the country where the myrrh
grows, it seems to me next to a proof, that this must have been the
opocalpasum of Galen.

I must however confess, that Galen says the opocalpasum was so

far from being an innocent drug, that it was a mortal poison, and
had produced very fatal effects. But as those Troglodytes, though
now more ignorant than formerly, are still well acquainted with the
properties of their herbs and trees, it is not possible that the savage,
desiring to increase his sales, would mix them with a poison that
must needs diminish them. And we may therefore without scruple
suppose that Galen was mistaken in the quality ascribed to this drug,
and that he might have imagined, from tenderness to the profession,
that people died of the opocalpasum who perhaps really died of the
physician: First, Because we know of no gum or resin that is a
mortal poison: Secondly, Because, from the construction of its parts,
gum could not have the activity which violent poison has; and
considering the small quantities in which myrrh is taken, and the
opocalpasum could have been but in an inconsiderable proportion to
the myrrh, to have killed, it must have been a very active poison
indeed: Thirdly, these accidents from a known cause must have
brought myrrh into disuse, as certainly as the Spaniards mixing
arsenic with bark would banish that drug when we saw people die of
it. Now this never was the case, it maintained its character among
the Greeks and the Arabs, and so down to our days; and a modern
physician, Van Helmont, thinks it might make man immortal if it
could be rendered perfectly soluble in the human body. Galen then
was mistaken as to the poisonous quality of the opocalpasum. The
Greek physician knew little of the Natural History of Arabia, less still
of that of Abyssinia, and we who have followed them know nothing
of either.

This gum being put into water, swells and turns white, and loses all
its glue; it very much resembles gum adragant in quality, and may
be eaten safely. This specimen came from the Troglodyte country in
the year 1771. The Sassa, the tree which produces the
opocalpasum, does not grow in Arabia. Arabian myrrh is easily
known from Abyssinian by the following method: Take a handful of
the smallest pieces found at the bottom of the basket where the
myrrh was packed, and throw them into a plate, and just cover them
with water a little warm, the myrrh will remain for some time
without visible alteration, for it dissolves slowly, but the gum will
swell to five times its original size, and appear so many white spots
amidst the myrrh.

Emfras, as I have said, is a large village something more than

twenty miles south from Gondar, situated upon the face of a hill of
considerable height above the lake Tzana, of which, and all its
islands, it has a very distinct and pleasant view; it is divided from the
lake by a large plain, near which is the island of Mitraha, one of the
burying-places of the kings. The inhabitants of the lower town, close
on the banks of the small river Arno, are all Mahometans, many of
them men of substance, part of them the king’s tent-makers, who
follow the camp, and pitch his tents in the field; the others are
merchants to the myrrh and frankincense country, that is, from the
east parallel of the kingdom of Dancali to the point Cape Gardefan,
or Promontorium Aromatum; they also bring salt from the plains, on
the west of the kingdom of Dancali, where fossile salt is dug; it is on
the S. E. border of the kingdom of Tigré. These Mahometans trade
also to the Galla, to the westward of the Nile; their principal
commodity is myrrh and damaged cargoes of blue Surat cloth, which
they unfold and clean, then stiffen them with gum, and fold them in
form of a book as when they were new.

This gum, which is called Sassa, they at first brought from the myrrh
country behind Azab, till ingenious and sagacious people had carried
plants of the tree to their different villages, where they have it
growing in great perfection, and more than supply the uses of the
merchants.
This tree grows to a great height, not inferior to that of an English
elm; that from which this draught was made was about two feet
diameter; the gum grows on all sides of the trunk, in quantity
enough almost to cover it, in form of large globes, and so it does on
all the principal branches. These lumps are sometimes so large as to
weigh two pound, though naturally very light.

The bark of the tree is thin and of a bluish colour, not unlike that of
a cherry-tree when young, or rather whiter. The wood is white and
hard, only the young branches which carry the flower are red. The
leaves are joined to the sides of the small branches by a small
pedicle of considerable strength, the leaves are two and two, or
opposite to each other, and have no single leaf at the point; they are
strongly varnished both on one side and the other, the back rather
lighter than the foreside of the leaf. The branches that carry the
leaves have about an inch of the stalk bare, where it is fixed to the
larger branch. There are generally fourteen leaves, each of about
three quarters of an inch long. At the top of the branch are knots
out of which come three small stalks, bare for about an inch and a
half, then having a number of small tubes, which, when they open at
the top, put forth a long pistil from the bottom of the tube. The top
of the tube, divided into five segments, or petals, arrives about one
third up the pistil, and makes the figure of a calix or perianthium to
it. From this tube proceeds a great number of very small
capillaments of a pink colour, at the end of each of which hangs a
purple stigma. At the top of this pistil is a large bunch of still finer
fibres, or capillaments, with stigmata likewise, and at the end the
pistil is rounded as if forming a fruit; without a very distinct drawing,
it would be difficult to make a description that should be intelligible.

Nothing can be more beautiful, or more compounded, than the

formation of this flower, though it has no odour; the head is
composed of about thirty of these small branches now described,
which make a very beautiful mass, and is of a pink colour of
different shades. At sun-set, the leaves on each side of the branch
shut face to face like the sensitive tribe. I never saw any seed or
fruit that it bore, nor any thing like the rudiments of seed, unless it
be that very small rotundity that appears at the end of the pistil,
which seem to bear no proportion to so large a tree.
 ERGETT Y’DIMMO.

T HE two beautiful shrubs which I have here given to the reader

are called by the name of Ergett, which we may suppose, in
Abyssinian botany, to be the generic name of the mimosa, as both of
these have the same name, and both of the same family, of which
there are many varieties in Abyssinia.

This first is called the Bloody Ergett, as we may suppose from the
pink filaments of which this beautiful and uncommon flower is in
part composed, and which we may therefore call Mimosa Sanguinea.
The upper part of the flower is composed of curled, yellow filaments,
and the bottom a pink of the same structure. I never saw it in any
other state. Before the blossoms spread it appears in the form here
exhibited. The pink, or lower part, in its unripe state, is composed of
green tubercules, larger and more detached than where the yellow
flower is produced, whose tubercules are smaller and closer set
together. I need not say the leaves are of the double pinnated kind,
as that and every thing else material can be learned from the figure,
full as perfectly as if the flower was before them; none of the parts,
however trifling and small, being neglected in the representation,
and none of them supposed or placed there out of order, for
ornament, or any other cause whatever: a rule which I would have
the reader be persuaded is invariably observed in every article
represented in this collection, whether tree or plant, beast, bird, or
fish.
 Ergett Dimmo.

London Publish’d Dec.r 1.st 1789 by G. Robinson & Co.

 Ergett el Krone

London Published Decem.r 1.st 1789 by G. Robinson & Co.

 ERGETT EL KRONE.

T HE next of this species of Ergett or Mimosa, is called in Abyssinia

Ergett el Krone, or the Horned Ergett; I apprehend the figure of
the pods have given it that appellation. Its flower in size and form
very much resembles the acacia vera, only that it is attached to the
branch by a long and strong woody stalk, which grows out at the
bottom of the branch bearing the leaves, and is sheltered as in a
case by the lower part of it. The branches of it are all covered with
very short, strong, sharp-pointed thorns, whose point is inclined
backward towards the root. Its pods are covered with a prickly kind
of hair, which, when touched, stick in your fingers and give very
uneasy sensations. The pods are divided into thirteen divisions, in
each of which are three round seeds, hard and shining, of a dusky
brownish colour. The flower has scarcely any smell, nor do I know
that it is of any utility whatever. Both these beautiful shrubs were
found upon the banks of the river Arno, between Emfras and the
lake Tzana. The soil is black mould, with a great mixture or
composition of rotten putrified leaves, thinly covering the rock in the
temperate part of Abyssinia. What I have to observe of both these
shrubs is, that they shut their leaves upon the violent rains of winter,
and are never fully expanded till the sun and fair season again
return.
 ENSETE.

T HE Ensete is an herbaceous plant. It is said to be a native of

Narea, and to grow in the great swamps and marshes in that
country, formed by many rivers rising there, which have little level to
run to either ocean. It is said that the Galla, when transplanted into
Abyssinia, brought for their particular use the coffee-tree, and the
Ensete, the use of neither of which were before known. However,
the general opinion is, that both are naturally produced in every part
of Abyssinia, provided there is heat and moisture. It grows and
comes to great perfection at Gondar, but it most abounds in that
part of Maitsha and Goutto west of the Nile, where there are large
plantations of it, and is there almost, exclusive of any thing else, the
food of the Galla inhabiting that province; Maitsha is nearly upon a
dead level, and the rains have not slope to get off easily, but
stagnate and prevent the sowing of grain. Vegetable food would
therefore be very scarce in Maitsha, were it not for this plant.
 Ensete
Heath. Sc.

London Publish’d Dec.r 1.st 1789 by G. Robinson & Co.

 Ensete
Heath. Sc.

London Published Dec.r 1.st 1789 by G. Robinson & Co.

Some who have seen my drawing of this plant, and at the same time
found the banana in many parts of the east, have thought the
Ensete to be a species of the Musa. This however, I imagine, is
without any sort of reason. It is true, the leaf of the banana
resembles that of the Ensete, it bears figs, and has an excrescence
from its trunk, which is terminated by a conical figure, chiefly
differing from the Ensete in size and quantity of parts, but the figs of
the banana are in shape of a cucumber, and this is the part which is
eaten. This fig is sweet though mealy, and of a taste highly
agreeable. It is supposed to have no seeds, though in fact there are
four small black seeds in every fig belonging to it. But the figs of the
Ensete are not eatable; they are of a tender, soft substance; watery,
tasteless, and in colour and consistence similar to a rotten apricot;
they are of a conical form, crooked a little at the lower end, about an
inch and a half in length, and an inch in breadth where thickest. In
the inside of these is a large stone half an inch long, of the shape of
a bean or cushoo-nut, of a dark brown colour, and this contains a
small seed, which is seldom hardened into fruit, but consists only of
skin.

The long stalk that bears the figs of the Ensete springs from the
center of the plant, or rather is the body or solid part of the plant
itself. Upon this, where it begins to bend, are a parcel of loose
leaves, then grows the fig upon the body of the plant without any
stalk, after which the top of the stalk is thick-set with small leaves, in
the midst of which it terminates the flower in form of the artichoke;
whereas in the banana, the flower, in form of the artichoke, grows at
the end of that shoot, or stalk, which proceeds from the middle of
the plant, the upper part of which bears the row of figs.

The leaves of the Ensete are a web of longitudinal fibres closely set
together; the leaves grow from the bottom, and are without stalks;
whereas the banana is in shape like a tree, and has been mistaken
for such. One half of it is divided into a stem, the other is a head
formed of leaves, and, in place of the stem that grows out of the
Ensete, a number of leaves rolled together round like a truncheon,
shoots out of the heart of the banana, and renews the upper as the
under leaves fall off; but all the leaves of the banana have a long
stalk; this fixes them to the trunk, which they do not embrace by a
broad base, or involucrum, as the Ensete does.

But the greatest differences are still remaining. The banana, has, by
some, been mistaken for a tree of the palmaceous tribe, for no other
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