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Moult and Ageing of European Passerines

Second Edition
Moult and Ageing of European Passerines

Second Edition

Lukas Jenni
Schweizerische Vogelwarte Sempach
Swiss Ornithological Institute

Raffael Winkler
Naturhistorisches Museum, Basel
Natural History Museum, Basel

Photographs by
Thomas Degen and Raffael Winkler
with additional material by Paul Mosimann, Christian Berger and Lukas Jenni
 HELM
Bloomsbury Publishing Plc
50 Bedford Square, London, WC1B 3DP, UK

BLOOMSBURY, HELM and the Helm logo are trademarks of Bloomsbury Publishing Plc

Second edition published in the UK in 2020

First published by Academic Press in 1994

Copyright © Lukas Jenni & Raffael Winkler, 2020

Lukas Jenni & Raffael Winkler have asserted their right under the Copyright, Designs and Patents Act, 1988, to be
identified as Authors of this work

All rights reserved. No part of this publication may be reproduced or transmitted in any form or by any means,
electronic or mechanical, including photocopying, recording, or any information storage or retrieval system, without
prior permission in writing from the publishers

A catalogue record for this book is available from the British Library

Library of Congress Cataloguing-in-Publication data has been applied for

ISBN: HB: 978-1-4729-4151-0;

ePDF: 978-1-4729-4150-3

Design by Marcel Burkhardt

To find out more about our authors and books visit www.bloomsbury.com and sign up for our newsletters

Cover drawings by Angelo Boog. They show a first-year male European Goldfinch Carduelis carduelis after the post-
juvenile moult which included the body-feathers, marginal and medium coverts, and greater coverts 4 – 10. The single
feather is the central alula feather of an adult Eurasian Jay Garrulus glandarius.
Contents

Preface XI

PART I

Chapter 1 The function and consequences of moult 1

1.1 Functions of the plumage 1

1.2 Plumage maintenance and the need for plumage renewal 2

1.2.1 Feather maintenance and wear 2
1.2.2 Adjustments to the plumage 3
1.3 The processes and costs of moult 5
1.3.1 The products and processes of moult 6
1.3.2 Costs of moult 6
1.3.3 Conclusions 7
1.4 Environmental effects on new feather quality 7

1.5 Fitting moult into the annual cycle 7

Chapter 2 Terminology and methods in moult research 9

2.1 Arrangement of the feathers 9
2.1.1 Flight-feathers 9
2.1.2 Wing-coverts 11
2.2 Terminology of plumages, feather generations and moults 11
2.2.1 Concepts of moult and plumage terminologies 11
2.2.2 General terms 13
2.2.3 Moult terms 13
2.2.4 Terms for plumages, feathers and feather generations 14
2.2.5 Age classes 14
2.3 Recording moult, plumage and feather properties 15
2.3.1 What to record: moult progress or moult intensity and of
which feather tract? 15
2.3.2 Scoring flight-feather moult 16
2.3.3 Scoring body-feather moult 17
2.3.4 Mass of feathers 17
2.3.5 Quantification of feather wear 19
2.3.6 Feather growth rates 19
2.3.7 Physical properties of feathers and fault bars 20
2.4 Analysing moult data 21
2.4.1 Seasonal timing and duration (progress) of moult 21
2.4.2 Moult intensity and moult speed 23
2.4.3 Sequence of moult 24
2.4.4 Extent of moult 24

Chapter 3 The moult of adults 25

3.1 Introduction to the moult strategies 25

3.2 Sequence of moult 26

3.2.1 Basic sequence of the complete moult 26
Flight-feathers 26
Body-feathers and wing-coverts 27
VI Contents

3.2.2 Functional aspects of the basic sequence of moult 27

3.2.3 Variations and exceptions to the basic moult sequence 29
Primaries 29
Secondaries 29
Tertials 31
Rectrices 31
Wing-coverts 32
Variation in the relationships between flight-feather tracts 32
3.3 Moult strategies 32
3.3.1 Complete post-breeding moult in the breeding area: Moult strategies 1 and 2 32
Complete post-breeding moult in the breeding area 32
Partial pre-breeding moult 33
3.3.2 Complete moult in the non-breeding area: Moult strategy 3 34
Partial moult before autumn migration 34
Suspension of the complete moult within the non-breeding area 34
Additional partial pre-breeding moult 34
Conclusions 35
3.3.3 Seasonally divided moult of remiges: Moult strategy 4 35
Seasonally divided primary moult (moult strategy 4a) 35
Seasonally divided secondary moult (moult strategies 4b and 4c) 36
3.3.4 Partial and complete biannual moult of remiges: Moult strategy 5 37
3.3.5 Summary and concluding remarks 38
3.4 Timing and duration of the complete moult 39
3.4.1 Timing and duration of the complete post-breeding moult in the breeding area 39
Timing of moult 39
Moult duration 40
3.4.2 Moult under time constraints 41
Reduction of moult duration 41
Overlap between breeding and moult 42
Overlap between moult and autumnal activities 43
Arrested moult 43
Transfer of moult to the non-breeding season 44
3.4.3 Timing of moult in trans-Saharan migrants 44
3.5 Summary and concluding remarks 47

3.6 Summary graphs and table of the moult strategies of European passerines 49

Chapter 4 The moult during the first year of life 61

4.1 Introduction 61

4.2 The juvenile plumage 61

4.2.1 Development of the juvenile plumage and completion after fledging 61
4.2.2 Structure and quality of the juvenile plumage 62
4.2.3 Coloration of the juvenile plumage 63
4.3 Sequence of post-juvenile moult 64
4.3.1 Sequence of the complete post-juvenile moult 64
4.3.2 Sequence of the partial post-juvenile moult 64
General sequence 64
Wing-coverts and alula 65
Tertials 66
Secondaries 66
Rectrices 67
4.3.3 Eccentric and other sequences of partial primary moults (post-juvenile and first pre-breeding) 67
4.3.4 Sequence and priorities of moult 69
4.4 How juveniles assume the adult moult strategies 71
4.4.1 Moult strategies 1 and 2: partial (complete) post-juvenile and complete
post-breeding moult in the breeding area 71
Partial pre-breeding moult 72
4.4.2 Moult strategies 3 and 5: complete moult in the non-breeding area 72
Incomplete first pre-breeding moult 73
  Contents VII

4.4.3 Moult strategy 4: seasonally divided moult 73

First partial pre-breeding moult including primaries 73
First partial pre-breeding moult including secondaries 73
4.4.4 Summary and concluding remarks 74
4.5 Variation in extent, timing and duration of the post-juvenile moult in the breeding area 75
4.5.1 Experimental evidence of control of post-juvenile moult 75
4.5.2 Intraspecific variation in timing and extent of the partial post-juvenile moult 76
Variation with hatching date 76
Differences between populations 77
Differences between the sexes and effects of energetic and nutrient stress 77
Intraspecific variation in the extent of post-juvenile moult during the course of the non-breeding
season and between wintering sites 78
4.5.3 Interspecific variation in timing and extent of the partial post-juvenile moult 79
Timing and duration 79
Extent 80
4.5.4 Partial post-juvenile primary moult in the breeding area 82
4.5.5 Complete post-juvenile moult in the breeding area 83
4.6 Summary and concluding remarks 84

PART II

Chapter 5 Ageing European passerines 87

5.1 Ageing criteria in live birds 87

5.2 Ageing using plumage characters 88

5.2.1 Recognition of juvenile feathers 88
Structure and shape 88
Coloration 88
Wear 88
Growth bars and fault bars 89
5.2.2 Differences in extent of moult 90
5.2.3 Differences between post-juvenile and subsequent feather generations 90
5.3 General ageing criteria in European passerines based on moult 90
5.3.1 Species with a complete post-juvenile moult in the first summer/autumn: Moult Type A 91
5.3.2 Species with a partial post-juvenile/complete post-breeding moult in the breeding area: Moult Type B 92
5.3.3 Species with a partial post-juvenile/complete post-breeding moult in the breeding area and a partial
pre-breeding moult in winter/spring: Moult Type C 95
5.3.4 Species with a complete moult in the non-breeding area: Moult Type D 96

Chapter 6 Species accounts 99

6.1 Presentation of the data and how to use it 99
6.1.1 Material 99
6.1.2 Relevance of the data 99
6.1.3 Presentation and analysis of the data 99

6.2 Procedure of ageing 100

Species accounts
Lanius collurio Red-backed Shrike 102
Lanius senator Woodchat Shrike 105
Lanius nubicus Masked Shrike 108
Oriolus oriolus Eurasian Golden Oriole 110
Garrulus glandarius Eurasian Jay 112
Pica pica Eurasian Magpie 114
Nucifraga caryocatactes Spotted Nutcracker 116
Corvus corone Carrion Crow 117
Bombycilla garrulus Bohemian Waxwing 120
VIII Contents

Periparus ater Coal Tit 122

Cyanistes caeruleus Eurasian Blue Tit 124
Parus major Great Tit 126
Riparia riparia Sand Martin 129
Hirundo rustica Barn Swallow 130
Delichon urbicum Common House Martin 131
Phylloscopus trochilus Willow Warbler 133
Phylloscopus collybita Common Chiffchaff 136
Acrocephalus schoenobaenus Sedge Warbler 140
Acrocephalus scirpaceus Eurasian Reed Warbler 142
Acrocephalus palustris Marsh Warbler 144
Hippolais icterina Icterine Warbler 146
Locustella naevia Common Grasshopper Warbler 147
Locustella fluviatilis River Warbler 150
Sylvia atricapilla Eurasian Blackcap 152
Sylvia borin Garden Warbler 156
Sylvia nisoria Barred Warbler 159
Sylvia curruca Lesser Whitethroat 162
Sylvia communis Common Whitethroat 165
Troglodytes troglodytes Eurasian Wren 172
Sitta europaea Eurasian Nuthatch 174
Sturnus vulgaris Common Starling 176
Turdus torquatus Ring Ouzel 177
Turdus merula Common Blackbird 179
Turdus pilaris Fieldfare 182
Turdus iliacus Redwing 184
Turdus philomelos Song Thrush 185
Turdus viscivorus Mistle Thrush 188
Muscicapa striata Spotted Flycatcher 190
Erithacus rubecula European Robin 194
Luscinia svecica Bluethroat 197
Luscinia luscinia Thrush Nightingale 199
Luscinia megarhynchos Common Nightingale 200
Ficedula hypoleuca European Pied Flycatcher 202
Phoenicurus ochruros Black Redstart 206
Phoenicurus phoenicurus Common Redstart 209
Saxicola rubetra Whinchat 212
Oenanthe oenanthe Northern Wheatear 215
Cinclus cinclus White-throated Dipper 218
Passer domesticus House Sparrow 219
Passer montanus Eurasian Tree Sparrow 221
Prunella modularis Dunnock 222
Motacilla flava Western Yellow Wagtail 224
Motacilla cinerea Grey Wagtail 229
Motacilla alba alba White Wagtail 232
Anthus campestris Tawny Pipit 236
Anthus pratensis Meadow Pipit 239
Anthus trivialis Tree Pipit 242
Anthus spinoletta Water Pipit 245
Fringilla coelebs Common Chaffinch 248
Fringilla montifringilla Brambling 250
Coccothraustes coccothraustes Hawfinch 253
Pyrrhula pyrrhula Eurasian Bullfinch 255
Chloris chloris European Greenfinch 257
Linaria cannabina Common Linnet 260
Acanthis cabaret Lesser Redpoll 263
Loxia curvirostra Red Crossbill 265
Carduelis carduelis European Goldfinch 271
Carduelis citrinella Citril Finch 274
Serinus serinus European Serin 276
Spinus spinus Eurasian Siskin 279
Emberiza citrinella Yellowhammer 283
Emberiza cia Rock Bunting 285
Emberiza hortulana Ortolan Bunting 287
Emberiza schoeniclus Common Reed Bunting 290
  Contents IX

APPENDIX: The use of skull pneumatization for ageing 293

The process of skull pneumatization 293
Recognition of skull pneumatization 293
Skull pneumatization scores 294
Age determination using skull pneumatization 295
Explanations of the graphs 295

References 301

Scientific names with their English, German, French, Italian and Spanish translations 317

Index 320

Quick reference key 322

  Preface
Preface
The two complementary aims of this book have remained unchanged
since its first edition in 1994. First, it presents an up to date summary
of the moult patterns of European passerines with due emphasis
on the extent of partial moults (Part I). This information is used to
describe, with the aid of photographs, detailed ageing criteria for 74
species (Part II). Although not an exhaustive reference to all aspects
of moult and ageing in European passerines, this book provides a
valuable complement to other guides, both through its extensive
collection of photographs and its detailed discussion of the extent,
sequence and strategies of moult.
Part I gives an overview of the various moult strategies of
European passerines based on our own observations and on data
from the literature. In chapter 1, we provide a short summary of the
basic aspects of moult: the functions of the plumage, the need for
plumage renewal, the costs and consequences of moult, and broadly
how moult is fitted in the annual cycle. Our understanding of all
these topics has increased tremendously since the first edition. In
consequence, we have prepared a separate, companion book (Jenni
& Winkler 2020) which treats these aspects of moult for birds of all
kinds, and make frequent reference to it here. Chapter 2 regarding
terms and methods in moult study has been expanded and now
not only explains the terminology, but covers the various ways of
recording moult, plumage and feather properties, and discusses
the various statistical methods of moult data analysis, which have
developed significantly over the last 25 years. Chapters 3 and 4,
which describe the moult of adults and young birds, respectively,
have been thoroughly re-written and updated. We describe the
moult strategies of European passerines based on the seasonal
occurrence and extent of moults of the entire plumage, which
may provide a more realistic insight into the inter- and intraspecific
variations in how moult fits into the annual cycle. We also highlight
how incomplete our knowledge of the moult strategies of trans-
Saharan migrants remains.
In section 3.6, we have added a graphical presentation of the
various moult strategies of European passerines, and a substantial
table which summarizes our current understanding of the moults of
all European passerine species. We hope that this will help to explain
the sometimes complex moult strategies of long-distance migrants.
Part II is devoted to ageing. We start with a general introduction to
ageing using plumage characters, followed by the species accounts.
Knowing the moult strategy of a species, and the variation in the
extent of its moult, is a prerequisite for ageing using plumage
characters. We have therefore deduced ageing criteria explicitly from
the observed moult patterns. This has led to our recognition of four
main moult types among the passerines breeding in Europe and a
description of their corresponding general ageing criteria (chapter 5).
We hope that this procedure, together with the information given
in the first part of the book (particularly in the summary table), will
enable readers to age birds in active moult more easily, to recognize
unusual and undescribed moult patterns and their implications for
ageing, and to develop plumage ageing criteria for species not
previously studied in detail.
We have added 16 species to the species accounts, bringing
the total to 74. They provide detailed information on the range in
extent of moult and explain how this can be used for ageing. We
avoid simplistic rules based on the most frequent moult patterns and
consider anomalous moult patterns as well. We are well aware that
XI
the data provided on the extent of moult relate mainly to central
Europe and that the restricted number of species treated prohibits
the use of this book as a general ageing guide to European passerines,
such are those by Svensson (1992) and Demongin (2013, 2016) (see
also the many websites and blogs mentioned in Demongin 2016, p.
386, which we could not deal with comprehensively here). Ageing
criteria for some more species can be found in the small ageing
and sexing guide by Winkler & Jenni (2007, 2009; in German and
Czech; see Conings et al. 1999 for a Dutch version). The main aim
of this book is to explain, with the examples given and the general
ageing criteria provided in chapter 5, how feather generations can be
recognized and how plumage-based ageing criteria can be deduced.
The colour photographs illustrate the variation in moult patterns
and the recognition of feather generations leading to ageing
criteria. They replace tedious descriptions of feather wear and colour
differences and should also speed up the process of acquiring a
feel for the clues to ageing any particular species. However, many
ageing criteria rely on the recognition of very slight differences in
feather colour and wear, and some experience in recognizing them
is certainly required.
The colour photographs concentrate on the wing, since most
plumage-based ageing criteria appear there. Almost the whole range
of moult extent is reflected on the wing since the marginal coverts
are among the first, and the secondaries among the last tracts to
be moulted.
The sections on ‘best criteria’ also include useful ageing criteria
other than plumage characters, and we systematically provide
information on the period of validity of skull pneumatization
(ossification) for ageing, based on our own data (and more fully
presented in the Appendix). Skull pneumatization is a very reliable
criterion for many species in summer and autumn, when it offers
the possibility of checking the reliability of newly discovered or
ambiguous plumage-based ageing characters. Many of the ageing
criteria and unusual moult patterns described here could only be
verified by ‘skulling’; which has also enabled us to identify certain
ageing criteria mentioned in the literature as unreliable.
The moult data and colour photographs presented here were
collected at several bird ringing stations in Switzerland (principally
Col de Bretolet) and elsewhere (see 6.1.1), and from material held
at the Natural History Museum, Basel. Our interest in moult began in
1972, but our systematic data collection only started in 1980. In total,
moult data have been collected from about 180,000 birds by the
authors and by Paul Mosimann, Christian Berger, Markus Leuenberger
and Thomas Degen. Skull pneumatization was recorded for all birds
observed in the summer/autumn.
All the photographs were taken from live birds, except about 45
which were from wing preparations in the Natural History Museum,
Basel. The feathers of the right wing were carefully arranged with a
pair of fine tweezers. Then, the wing was placed on an oblique board
covered with a standard grey paper. A small piece of double-sided
adhesive tape, 1.5 mm thick, held the wing just distally of the wrist.
Photographs were taken on 35 mm slide film, 64 ASA, with a reflex
camera mounted on a tripod and equipped with a ring flash. This
relatively simple technical equipment had the advantage that it was
easy to transport and install, even in the field. A disadvantage was the
inability to ensure exact equivalence of the colours between the slides.
For our purposes, however, absolute colours are not as important as
XII Preface
relative differences in colour between feathers on the same wing.
For the second edition all of the slides were photographed on a light
box by Marcel Burkhardt using a digital camera, which gave results
at least as good as conventional scanning of the slides. The digital
photographs were then treated by Jörg Brunner and the background
replaced by a uniform picture of a grey paper.
Throughout, we followed Gill & Donsker (2017) for names and
the order of species.
This work would have been impossible without the tremendous
help of many people and we are greatly indebted to them all. For
the first edition these were: Christian Berger, Thomas Degen, Markus
Leuenberger and Paul Mosimann who all showed great enthusiasm
and perseverance in the collection of moult data and photographs
over a number of years. Others too numerous to name assisted at the
bird ringing stations in Switzerland, although Hildegard Messer­knecht
and Roland Ammann deserve our special thanks. Ernst Sutter laid
the foundations for the carefully documented skin collection in the
Natural History Museum, Basel, and provided the data for Garrulus
glandarius. Fernando Spina, head of the Italian Ringing Scheme,
enabled Raffael Winkler and Paul Mosimann to work on Ventotene
Island. Peter Keusch, John Attard Montalto and Frank Neuschulz
made it possible to catch and take photographs of Emberiza
hortulana, Cisticola juncidis and Sylvia nisoria in their study areas.
Markus Leuenberger and Thomas Degen helped with the analysis of
the data. Les Underhill carried out the analyses of the dependence
of moult extent on skull pneumatization scores and feather-length,
and the analysis of the timing and duration of moult of some species
from our data. Unpublished data were provided by Bruno Bruderer,
Carlos de la Cruz, André A. Dhondt, Thord Fransson, Charles Gauci,
Hans Leuzinger, Åke Lindström, Gerhard Nikolaus, David Pearson,
Werner Prünte, Fernando Spina, Joe Sultana and Michael Widmer.
Tomasz Wesołowski translated important literature from Russian.
Eberhard Gwinner, Dominique G. Homberger, Åke Lindström and
David Pearson provided valuable comments on earlier drafts of parts
of the manuscript.
For the second edition, we are indebted to: the late David Pearson
who enabled the authors and Matthias Kestenholz to examine birds
at Ngulia, Tsavo West National Park in Kenya; Reuven Yosef and
John Morgan who made data collection for RW possible at the
International Ornithological Center Eilat, Israel; Tatiana Rymkevich
and George Noskov who invited RW to Gumbaritsy, NW Russia;
Nicola Baccetti allowed RW to scrutinize a confiscated collection
of illegally shot birds at the Istituto Superiore per la Protezione e
la Ricerca Ambientale ISPRA, Italy; the Fritz Sarasin Foundation
of the Freiwillige Akademische Gesellschaft, Hilfsfonds für die
Schweizerische Vogelwarte, Rudolf Geigy Fonds of the Natural History
Museum, Basel and Swiss Academy of Sciences for financial help for
travel; Sandro Gröflin, Natural History Museum, Bern, David Norman,
Daniel Peyer, the late Werner Prünte (through his son Falko Prünte),
Florian Steiner, Daniel Tschanz, Lorenzo Vinciguerra, the ringers of
the Sahara project of the Swiss Ornithological Institute in Mauritania,
and ringers participating in the European-African Songbird Migration
Network for providing valuable data and material; Birgit Erni, Fränzi
Korner-Nievergelt, Ian Newton, Michael Schaub, Les Underhill and
Jan von Rönn for commenting on chapter 2; Volker Salewski for
commenting on chapter sections 3.4.3 and 3.5; Barbara Helm for
helping with chapter section 4.5; Gabriel Gargallo, Natalia Iovchenko
and Hadoram Shirihai for fruitful discussions; Patricia Dürring and
Anne Tampe for literature searches and checks; Shannon Luepold
for help with the English language; Susanne Jenni-Eiermann for
constant support; Daniel Scherl, Simon Hohl and Tabea Kölliker for
reconstructing and modifying the graphs from the first edition and
for drawing the new ones; Marc Kéry for statistical advice and for
producing Fig. 60; Isabelle Kaiser for help with the layout; Bernard
Volet for the multilingual list of bird names.
We owe a great deal to Marcel Burkhardt, Swiss Ornithological
Institute, who digitized all the slides, made the photographs for
chapter 2, created the layout, and produced the book up to the
final stage for the printers; and to the Swiss Ornithological Institute
for financing the preparation of this book. We were fortunate that
Andrew Richford, editor and ornithologist at Academic Press for the
first edition, was able to improve the English language usage and
advise on this second edition. We thank Jim Martin of Bloomsbury
Publishing for his patience and help in the production of this book
and its companion volume.
  The Function and Consequences of Moult 1

PART I
Chapter 1

The Function and Consequences of Moult

1.1 Functions of the plumage

The process of moult serves to keep the plumage in good condition

and adjust its characteristics to the changing needs of a bird during its
lifetime. Understanding moult therefore requires a knowledge of the
various functions of the plumage (see Jenni & Winkler 2020: chapter 1
for a more detailed treatment and supporting literature).
Feathers form a flexible, lightweight protective barrier against
mechanical impact, water, and solar and terrestrial ionizing radiation.
In addition, the plumage forms the principal layer of thermal insulation
and plays an important role in thermoregulation (Wolf & Walsberg
2000). Short-term adjustments to the degree of insulation provided can
be effected by changes in the position of the feathers (e.g. fluffing).
Long-term adjustments can be made by changing the number, mass or
length of the feathers, through feather loss or the growth of additional
or differently structured feathers during moult (see 1.2.2). The density
and structural properties of body-feathers and down differ between
species, depending on phylogeny, climate and habitat characteristics
(Pap et al. 2017, Osváth et al. 2018).
Fig. 1. European Goldfinch Carduelis carduelis ad , 22 October. After the post-breeding
The unique function of feathers is that they enable birds to fly by moult in autumn, the entire plumage is fresh, the tertials, secondaries and primaries
forming the major part of the aerofoil-like wings and by clothing the having unworn, white tips.
body in an aerodynamically efficient shape (Homberger & de Silva
2000, Videler 2005).
The plumage is also largely responsible for the appearance of a bird,
giving it its colour and shape, and is therefore important for camouflage
and visual communication (e.g. interspecific signalling, intraspecific
signalling of e.g. individual dominance and quality, and facilitation of
foraging; Bortolotti 2006, Dale 2006, Hill 2006). Specially coloured
parts or adornments have often evolved which may be apparent only
in one sex or only during certain times of the year. Every plumage
trait is produced only during the moult period(s) (although they may
change over time with wear). Therefore, the environmental conditions
experienced during moult, and how a bird copes with these, play an
important role in determining the appearance of a bird (see also 1.4).
Specialized feathers may act as tactile sensors, such as facial bristles
(which can have various functions), and filoplumes (hair-like structures
with a few short barbs at the end), which are associated with contour
and flight-feathers and provide information about the position and
vibration of flight-feathers (Lucas & Stettenheim 1972, Necker 1985,
Cunningham et al. 2011, Persons & Currie 2015).
Many other functions of the plumage, or individual feathers, only
Fig. 2. European Goldfinch Carduelis carduelis ad , 25 April. After about 7 months,
appear in certain taxa or species (mainly non-passerines) including: the plumage is worn. Note how abrasion has especially affected the less resistant white
regulation of buoyancy (waterfowl); production of sound (snipe); tips of the exposed tertials and outer primaries. However, the white tips remain on the
protected inner primaries and secondaries.
reduction of the sound produced by the wings in flight (owls and nightjars);
 2 The Function and Consequences of Moult
enhanced hearing (owls); transport of water (sandgrouse); mechanical
support (woodpeckers and treecreepers); chemical defence and
chemical communication (e.g. pitohuis); maintenance of the plumage
through the production of powder-down (pigeons); provision of nesting
material (certain waterfowl); and assisting digestion (grebes) (reviewed
in Jenni & Winkler 2020).
Many functions of the plumage exist in potential conflict with each
other, so that the ‘best’ plumage is always a well-balanced compromise
between opposing requirements which may differ between species,
sexes, age-classes, seasons and environmental conditions. For example,
the remiges are under strong selection pressure for optimum flight
performance, which generally militates against the evolution of
elaborate structural ornaments, while ornamental colour patches on
wings are frequent. The structure of the wing-feathers themselves is
always a finely tuned compromise between different flight requirements
(e.g. long primaries efficient for migration versus shorter ones to enable
foraging in dense vegetation; e.g. Leisler & Winkler 1985, Vágási et
al. 2016).
There is a general conflict between the need for concealment and
conspicuous signal coloration. Almost all birds are under predation
pressure or are predators themselves, both in need of camouflage,
but at the same time using plumage patches or patterns for signalling.
Compromise solutions to this conflict include: distinctive colour
patterns which are unusual or easily distinguished close to, but are
cryptic at long range, as opposed to conspicuous coloration which
is easy to see at a distance; reduction of conspicuous coloration to
certain feathers which are normally hidden and are presented only
on certain occasions; changing the colour of the plumage seasonally
through the abrasion of cryptic feather fringes or by moulting twice
a year (see 1.2); and confining conspicuous coloration to only one
sex or to certain age groups. Usually, plumage patterns intimately
mix patches selected for camouflage and patches selected for
conspicuousness (e.g. Butcher & Rohwer 1989, Gomez & Théry 2007).
The general rule whereby a bird appears cryptic at long range and
conspicuous at close range is variously modified from very cryptic
to very conspicuous, according to the relative importance of the
selection pressures for camouflage and conspicuousness, very often
depending on sex, age and season. The compromise struck between
conspicuousness and crypsis, or between adornments and flight
capability, is ‘used’ by sexual selection so that males can honestly
advertise their own quality to potential mates or rivals by displaying
a genuine handicap which they have clearly overcome (Zahavi 1975,
1977). Some cryptic plumage patterns may also serve a dual function
as signals of quality, such as quality related variations in the regularity
of camouflage barring patterns (e.g. Hasson 1991).
A further conflict may arise because a high melanin content (which
produces black, brown, red-brown and yellow feather colours) increases
feather durability so that feathers exposed to intense wear should not
be light in colour (Burtt 1986, Bonser 1995; Fig. 1 and 2, see 1.2).
Resistance to wear may therefore conflict with the colour requirements
of camouflage or display. In contrast, plumage coloration and heat gain
from solar radiation are more or less independent, because colour is
only one of several factors determining the penetration of radiation into
the plumage (Walsberg 1983, Wolf & Walsberg 2000).
Very few studies have tried to use an integrated approach to
understanding how the various requirements determine the properties
of the plumage (e.g. Burtt 1986). Our incomplete knowledge of
the significance of plumage properties makes it difficult to always
understand the timing, extent and sequence of moult of a particular
species or individual.
1.2 Plumage maintenance and
the need for plumage
renewal
Full-grown feathers are dead structures consisting mainly of avian
keratin. Keratin is one of the most durable biological materials,
with great strength, flexibility and resistance to hydrolytic, protein-
digesting enzymes and bacteria. Nevertheless, feathers can be lost
or become irreparably damaged.
However, unlike other keratin structures like hair and claws, they
cannot be renewed continuously from the base. Consequently,
unsuitable feathers must be replaced totally. Furthermore, feathers
can only be replaced by pushing out the old feather long before the
new one is fully grown and functional. This is a major disadvantage of
feathers since it results in a significant reduction in plumage function
when many feathers must be replaced simultaneously.
The regular replacement of all or part of the plumage is called
moult (see Jenni & Winkler 2020: 3.1.1 for definition). There are two
main reasons for plumage renewal: (a) damaged or lost feathers need
to be replaced in order to maintain the function of the plumage; and
(b) any feathers unsuitable for their current or future purpose must
be replaced in order to adjust the plumage to changing requirements.
For a more detailed treatment and supporting literature see Jenni &
Winkler (2020: chapter 2).
1.2.1 Feather maintenance and wear
Feathers are regularly maintained by a variety of comfort behaviours
such as preening (with the bill), scratching (with the foot), shaking,
bathing and drying, oiling with the secretion from the uropygial or
preen gland, dusting, sunning, and possibly anting (reviewed by
Simmons in Campbell & Lack 1985, Clayton et al. 2010, Bush &
Clayton 2018). Inevitably, however, feathers are lost or irreparably
damaged due to mechanical abrasion, photochemical processes,
ectoparasites, bacteria, and fungi. We group all of these factors
(except pathological damage) under the more general term ‘wear’
(other authors use the term ‘abrasion’ in the same general sense; e.g.
Ginn & Melville 1983, Rogers 1990).
Entire feathers may be lost through traumatic mechanical
interaction with vegetation, rocks, the nest, conspecifics, or predators.
Feathers may also be lost when a bird is stressed (‘fright moult’;
Dathe 1955, Höglund 1964, Stiefel in Bub 1985, Lindström & Nilsson
1988, Diesing 2001, Møller et al. 2006) and in forming the brood
patch (Schifferli 1981).
Feathers are damaged by sunlight. Ultraviolet light alters the
physical structure of keratin and pigments leading to bleaching.
Flight-feathers suffer from mechanical fatigue, i.e. the loss of bending
stiffness caused by their repeated bending upwards and downwards
during every wing beat (e.g. Weber et al. 2005).
Abrasion, caused by rubbing against other feathers, objects in
the environment, and particles in the air, or by the tearing apart of
feathers frozen together, damages the structure of the feathers by
breaking off barbicels, pennulae and barbules, and by cracking and
splitting the barbs and the rachis (Burtt 1986, Willoughby 1991). This
impairs, or may completely prevent, the cohesion of the vane. Abrasion
of the exposed, and often more loosely textured, feather fringes
  The Function and Consequences of Moult
reduces the feather’s surface area and may substantially change
its shape (Fig. 5). Abrasion also affects plumage colour. The
feathers of many birds have dark centres and light fringes. As
the plumage wears, it increasingly takes on the colour of the
feather centres and may become more or less bright or cryptic.
Ectoparasites such as feather lice (Insecta, Phthiraptera: Amblycera
and Ischnocera; formerly collectively termed Mallophaga) actually eat
feather material. The effect of feather mites (Acari, Prostigmata and
Astigmata) on feathers differs between species. A few mite species
(the syringicoles) may damage the feather quill by feeding on the
medulla of the rachis. However, most feather mites live on the feather
vanes (the plumicoles) and seem to be commensals feeding on fungi,
and possibly bacteria, admixed with the uropygial gland secretions,
as well as algae, pollen, debris and other micro-organisms (Proctor
& Owens 2000, Blanco et al. 2001, Proctor 2003, Doña et al. 2018).
Although it is widely believed that moult may reduce the ectoparasite
burden (e.g. Baum 1968, Post & Enders 1970), more recent studies
have shown that feather lice are as abundant after moult as before,
but more concealed, or even more abundant (Moyer et al. 2002,
Hamstra & Badyaev 2009; reviewed in Clayton et al. 2010), and
plumicolous mites can apparently sense which wing-feathers are
about to be dropped and change feathers to avoid being dropped with
the feather (Jovani & Serrano 2001, Jovani et al. 2006, Pap et al. 2006).
More recently, a whole suite of bacteria capable of decomposing
β-keratin was discovered on the feathers of birds, but their effects
on feathers or fitness in live birds have not been studied in detail
(Clayton 1999, Burtt 2009). Nevertheless, feather parts coloured
by melanins are more resistant to feather-degrading bacteria than
white and carotenoid-coloured feathers in vitro; an effect which may
not only be caused by the pigments, but also by concomitant more
thickly keratinized feather elements (Goldstein et al. 2004, Gunderson
et al. 2008, Burtt et al. 2011, Ruiz-de-Castañeda et al. 2012, but
not Grande et al. 2004). Indeed Bacillus licheniformis preferentially
colonizes the unmelanized parts of a feather (Justyn et al. 2017).
Interactions between different species or strains of feather-
degrading bacteria, or interactions between feather-degrading
bacteria and other bacteria or fungi, may result in a complex
‘ecosystem’ on a bird’s feathers and body and have not yet been
investigated. For example, the bacteriocins produced by enterococci
bacteria in the uropygial gland of female and nestling Eurasian
Hoopoes Upupa epops inhibit the keratinolytic activity of feather-
degrading bacteria (Ruiz-Rodriguez et al. 2009).
Wear affects different feathers differently depending on the
exposure, shape, structure and pigmentation of the feather in
question and also on the behaviour of the bird (Fig. 3 – 6). Generally,
the more loosely textured feathers of the juvenile plumage are more
prone to abrasion than the feathers of subsequent generations (Fig.
7). Exposed feathers abrade and bleach more readily than concealed
feathers (Fig. 4 – 7). In many species, the tertials, rectrices, inner (but
not the innermost tenth) greater coverts, and the tips of the primaries
are especially exposed. The secondaries, inner primaries and primary
coverts, outer greater coverts, and the carpal covert are normally
well protected (Fig. 4 – 7). These differences in exposure may lead to
situations in which feathers generated more than six months apart
can no longer be distinguished by the degree of wear (see e.g. spring
individuals of the Tawny Pipit Anthus campestris, Western Yellow
Wagtail Motacilla flava and Common Whitethroat Sylvia communis
in the species accounts).
It has long been known that melanic feathers are more resistant to
ultraviolet radiation and mechanical wear than non-melanic feathers,
3
while carotenoids do not increase wear resistance (e.g. Barrowclough
& Sibley 1980, Bergman 1982, Burtt 1986, Ellis 2006; Fig. 1 and
2). The inclusion of melanin granules increases the hardness of
feather keratin by 39 % (Bonser 1995). This may be partly due to
the resulting increase in the thickness of keratinized feather elements
since certain mechanical properties of feather barbs are determined
by morphology (e.g. increased cross-section) rather than melanin
content per se (Voitkevich 1966, Butler & Johnson 2004).
Wear depends on habitat, weather, climate and season. Birds
sliding through dense, hard vegetation such as reeds, grass, sedges
or thorny bushes, or exposed to sand and wind suffer more abrasion
than aerial and perching birds. Birds exposed to intense sunlight suffer
more bleaching than those living in shady forests (cf. Fig. 3 with Fig.
4 – 6). Bleaching may be more pronounced in tropical winter quarters
than in the temperate breeding area. Changes in wing-length indicate
that abrasion is stronger during the breeding season (due to activities
in the nest and intensive foraging) than outside the breeding season
(see Jenni & Winkler 2020: 2.1).
The fact that the extent of wear differs with habitat, climate,
weather, exposure, and pigmentation of a feather must always be
kept in mind when assigning feathers to different feather generations
and in reconstructing the extent of moult in birds outside the
moulting season.
Wear results in the loss of feather material and a change in the
shape and colour of feathers. Therefore, wear affects all functions
of the plumage, in particular thermal insulation, flight function and
appearance. In particular, feather ornaments may break or fade.
During the time over which body-feathers become worn, their mass
and number decreases substantially (e.g. Newton 1968b, Schifferli
1981). Wear also results in the abrasion of flight-feather tips, especially
the exposed feathers and those forming the tips of the wing and tail,
and this leads to a change in wing shape (e.g. in the Eurasian Reed
Warbler Acrocephalus scirpaceus; Peiró 2003) and a reduction of
wing-length and tail-length with feather age (for details see Jenni
& Winkler 2020: 2.1.5). Although wear is clearly associated with a
deterioration of plumage ornaments, abrasion can also change the
coloration of a bird in an adaptive way (see 1.2.2).
The plumage of a bird after moult, although composed of dead
material, is therefore by no means a constant trait, but one which
changes over time dependent on wear and plumage maintenance.
1.2.2 Adjustments to the plumage
Birds may need to adjust their plumage in response to seasonal
changes, age and changing environmental conditions. For some
time, studies have focussed on the description of changes in plumage
coloration according to age and season, but only recently has it been
realized that the actual plumage structure may also change.
Many passerine species have a distinct juvenile plumage (see
4.2) which is usually replaced with a more colourful adult plumage
within the first year. In some sexually dimorphic passerines, first-year
males acquire a more female-like plumage rather than the adult-male
plumage, although they may in fact breed. This delayed plumage
maturation has been explained in various ways, but most evidence
suggests that it is part of a strategy to reduce competition with older
individuals by honestly signalling subordinate status (Berggren et al.
2004, Hawkins et al. 2012). Actually, in most European passerines
the juvenile plumage is only partly replaced during the post-juvenile
moult (see 4.4). Therefore, the remaining juvenile feathers can still be
4 The Function and Consequences of Moult
Fig. 3. Eurasian Blackcap Sylvia atricapilla ad , 25 April. This species shows hardly any
signs of wear in spring, although the plumage was acquired about nine months previously,
during the postbr moult.
Fig. 4. Subalpine Warbler Sylvia cantillans ad , 21 April. The entire plumage is postbr,
except for some MaC and MeC, GC 4 and probably T which are prebr. The postbr feathers,
acquired about nine months before, are noticeably abraded and bleached. This species
winters in the tropics and flying through thorny scrub has focussed wear on the exposed
tertials, the inner greater coverts (except the protected GC 10), the tips of the primaries
and the outer primary coverts. The tips of the protected secondaries are hardly worn,
although the more exposed fringes of their outer webs are slightly abraded.
recognized by conspecifics and may constitute a special case of
delayed plumage maturation (Senar et al. 1998, Hawkins et al. 2012).
In European passerines, classical delayed plumage maturation
through a distinct subadult plumage is rare. In Black Redstart
Phoenicurus ochruros about 90 % of first-year males have a female-
like plumage while the remainder have an almost adult-male-like
plumage (see p. 206). Lesser Redpoll Acanthis cabaret males do not
have a red breast in their first year (Winkler & Jenni 2007). In the Red-
backed Shrike Lanius collurio and the Barred Warbler Sylvia nisoria,
the feathers acquired during the post-juvenile moult are markedly
different from those of adults, especially in male Red-backed Shrikes
(hence termed second juvenile plumage by Stresemann 1963a),
but are moulted into the adult plumage only a few months later,
during the subsequent moult in the winter quarters. Male Common
Fig. 5. Greater Short-toed Lark Calandrella brachydactyla, 23 April. The entire plumage is
postbr or postjuv. The feathers, acquired about nine months previously, are considerably
abraded. This species lives in a sunny, dusty environment and running through grassy
vegetation has primarily worn the tertials, the innermost greater coverts and the outer
webs of the inner secondaries, which are most exposed when the wing is closed. The
primaries and outer secondaries are protected by the long tertials, and the central and
outer greater coverts and the primary coverts by the inner greater coverts, and so show
very little wear.
Fig. 6. European Stonechat Saxicola rubicola 2y , 1 May. The MaC, MeC, GC, CC, Al
1, T and S 6 are postjuv, the remainder of wing juv. The feathers acquired 8 – 12 months
previously are considerably worn. This sit-and-wait predator hunts mainly by flying into
low vegetation and wear has mainly affected the tips of the remiges, including the
secondaries, these feathers being those exposed when the wing is open. The relatively
short tertials and the greater coverts are less worn, both because they are less exposed
on the open wing and because they were renewed in summer/autumn.
Rosefinches Carpodacus erythrinus acquire a female-like plumage
in their first winter during an extensive post-juvenile moult and may
breed in this plumage during their first breeding season, the red
breeding plumage only being acquired during the second winter
(Cramp & Perrins 1994a, Glutz von Blotzheim & Bauer 1997).
In a few European passerines, the adult plumage may become
slightly more colourful with increasing age (e.g. European Pied
Flycatcher Ficedula hypoleuca, Winkel et al. 1970, Potti & Merino
1995, Potti et al. 2014; Whinchat Saxicola rubetra, Schmidt & Hantge
1954; European Robin Erithacus rubecula, Jovani et al. 2012).
As mentioned (see 1.1), the requirements of camouflage and
display may be in conflict. In many species, their relative importance
differs depending on season or on the age of the bird. Therefore, it
may be advantageous for birds to change their appearance during
  The Function and Consequences of Moult
Fig. 7. European Serin Serinus serinus 2y , 5 May. The MaC are postjuv, the MeC
1 – 4 + 6 + 8 juv, and MeC 5 + 7 postjuv. The remainder of the wing is juv. The loosely
textured juvenile median coverts were acquired about ten months previously and are
more worn than the more firmly textured postjuv median coverts acquired about nine
months earlier. Within the feather tracts, wear has primarily affected the inner greater
coverts (except the protected GC 10) which lie over the outer ones and cover them when
the wing is closed. Within the median coverts, however, the outer ones lie over the inner
ones and are therefore more heavily worn.
the year, or as they age. Apart from temporary exposure of normally
concealed colourful feather patches or conspicuous structures, there
are two ways in which appearance may be altered.
First, abrasion is a limited, but nevertheless striking method of
changing plumage colour. The feathers of many species have drab,
light-coloured fringes. When these fringes wear off, the plumage
increasingly assumes the colour of the feather centres and may
become more or less bright or less cryptic. Well known examples are
the light fringes on the body-feathers of species such as Common
Chaffinch Fringilla coelebs, Brambling F. montifringilla, Common
Linnet Linaria cannabina, Common Starling Sturnus vulgaris, etc.,
which are worn away during winter to reveal the colourful or shiny
breeding plumage in spring. Abrasion of these fringes is often
facilitated by their light colours, which contain little or no melanin.
In addition to the loss of light fringes, the carotenoid-based colours
of the feather centres may brighten over winter in time for the
breeding season, rather than fading (see above), possibly either
because barbules break away or irradiation by sunlight denatures
feather proteins that might conceal carotenoid pigments (Willoughby
et al. 2002, Blanco et al. 2005).
The second way in which a bird may change its appearance is by
moulting its body-feathers twice a year and this is the only way in
which appearance can change profoundly with age or season and
special feather structures can be introduced. Species having more
than one moult per year usually change between a colourful ‘display’
plumage worn during the mating season and a more cryptic, dull non-
breeding plumage worn during the rest of the year (see 3.3.1). Many
species change from a cryptic juvenile plumage to a more colourful
adult plumage (see 4.2.3).
Less well studied are changes of plumage structures which are
important in behaviour and display with age and environmental
conditions. Many plumage characters which play a role in sexual
selection, aggressive encounters and establishing dominance in non-
breeding flocks are less prominent in first-year birds than in older ones,
and may change with environmental conditions. Therefore, moult
provides a means to adjust an individual’s status and quality signals
5
to its age and to the prevailing environmental conditions (see Jenni
& Winkler 2020 for details).
The plumage performs functions other than governing appearance,
which may be adjusted according to season and age. It is well
known that the juvenile remiges of passerines are usually shorter
than their successors, with the exception of the outermost primary
which is usually longer. The shorter length of the juvenile remiges,
which are often also of lower mass, is usually ascribed to nutritional
constraints during the nestling period. A more likely explanation is
that it minimises the time spent in the nest waiting for the feathers to
grow fully, and therefore reduces predation risk (Nilsson 1992, Nielsen
1993). Finally, the more rounded and slotted wings of juveniles may
be an adaptation to increase flight manoeuvrability and take-off
performance as an antipredator adaptation during the first phase
of independence, at the cost of lower flight efficiency during long-
distance flights (Alatalo et al. 1984, Hedenström & Rosén 2001). In
many species, the shape of the individual juvenile feathers may also
differ from that of the following feather generations. The juvenile
rectrices of many species are more pointed than the more rounded
rectrices of adults (see e.g. Svensson 1992, Winkler & Jenni 2007).
By having two moults per year, the amount of insulation provided
by the plumage can be adjusted to suit the seasons. In the American
Goldfinch Spinus tristis, the number and length of pre-breeding and
post-breeding body-feathers are similar, but the post-breeding ones
have more plumulaceous barbs resulting in a considerably heavier,
‘winter’ plumage (Middleton 1986). In contrast, in birds with only
one moult per year, plumage insulation may be reduced through
feather loss and wear.
Because body-feathers are lost throughout the year, it would
be advantageous to restore the plumage before the winter in cold
and temperate climates and so increase insulation. Indeed, total
insulation has been found to be better after moult than before it
(reviewed in Jenni & Winkler 2020: 4.3.2). The lighter, less dense,
more loosely structured juvenile plumage might need replacement
before wintering in cold areas. Thus, the post-juvenile moult increases
the insulation effect for the oncoming winter. Likewise, feather loss
at the approach of the warmer season may also be adaptive.
The two main functions of moult, replacing worn feathers and
adjusting plumage characteristics, will often be cited in chapters 3 and
4 to explain particular moult strategies or the extent of a partial moult.
However, in practice it is often impossible to distinguish between
them, because our knowledge of the degree and effects of feather
wear and the exact requirements of plumage function (especially the
behavioural significance of particular plumage colours) are still poor.
1.3 The processes and costs
of moult
Feathers are cornified and are, therefore, dead structures, so need
to be replaced periodically (see 1.2). They can be replaced only by
pushing out the old feather long before the new one is fully developed
and functional, which inevitably results in a temporary reduction
in plumage function. Therefore, birds face two major interrelated
challenges during the period of moult, both dependent on moult
intensity: (a) additional energetic and nutritional requirements; and,
(b) reduced performance in all plumage functions (e.g. flight, aerobic
6 The Function and Consequences of Moult
capacity, insulation, signalling) leading to greater vulnerability (e.g. to
predation, disease, starvation). A short summary of the review given
in Jenni & Winkler (2020: chapter 3) follows.
1.3.1 The products and processes of moult 
During a complete moult, thousands of feathers are produced
including the auxilliary structures lost to the environment during
feather deployment (feather sheaths, dead supporting cells, etc.),
all consisting of 90 – 95 % protein (keratin). For small birds, the
feather-mass produced at least once a year is 6 – 12 % of total body
mass or 18 – 39 % of total lean dry body mass. Body-feathers make
up 62 – 78 % of total plumage mass, and flight-feathers 22 – 38 %
(see Jenni & Winkler 2020: 3.2.2 for details). A complete moult may
coincide with the renovation of other epidermal structures, such as
podotheca, ramphotheca, claws and the stratum corneum of the
skin. Little is known about the renovation of the skeleton during
moult (Meister 1951, Murphy et al. 1992).
Moult is a period of profound physiological change and requires
the reinstatement and operation of the entire physiology governing
feather growth. Body temperature, body water turnover, the
vascular network, blood volume, and peripheral blood flow (in
the growing feathers) all increase. Body mass at moult is generally
low, mainly because fat stores are very low at this time, but organ
sizes may also be reduced or upgraded during moult. Part of the
immune system is enhanced, possibly because the risk of injury is
increased. In addition, processes apparently not directly involved in
feather production, such as whole-body protein turnover and bone
renovation, are increased during the moult period. This results in
an increase in protein metabolism.
Various body functions appear to be downregulated during an
intensive moult, such as (other) parts of the immune system and the
physiological response to stressors. This may be a direct consequence
of moult (e.g. resources are redirected towards feather production)
or to compensate for the additional energetic and nutritional
burden of moult (see 1.3.2). Importantly, certain behaviours are
greatly reduced during moult. A reduction in behavioural activity
may protect the growing feathers from damage, reduce predation
risk and decrease energy expenditure.
1.3.2 Costs of moult
The periodic replacement of the plumage entails two types of
cost: (a) the replacement of feathers requires energy and special
nutrients as well as metabolic and physiological adaptations; and
(b) the functions of the plumage cannot be fully maintained during
moult. However, a bird has many ways in which to compensate
for these costs.
Outside the moult period, the costs of replacing experimentally
plucked feathers seem to be quite low (Schieltz & Murphy 1997,
Hoye & Buttemer 2011, Buttemer et al. 2015, Ben-Hamo et al.
2017). However, during the moult period the energetic costs of
moult during the nightly resting period at thermoneutrality generally
amount to 12 – 36 % (or exceptionally 68 and 92 %) of non-moult
resting metabolic rate and are dependent on moult intensity and
the size of the bird (Jenni & Winkler 2020: 3.4.1). Therefore, other
processes occurring during the moult period, such as increased
protein turnover, may account for a major part of the added
energy expenditure. However, it remains to be shown whether the
estimates of nocturnal moult costs are also valid during the day.
Daily energy expenditure during moult in captive birds increases
by 0 – 35 %, which is on average lower than the increase in resting
metabolic rate at night during moult. Daily energy expenditure during
moult in free-living birds increases by only 0 – 15 %, compared with
non-moulting birds not occupied with other activities (e.g. breeding),
as indicated by the small number of available studies. However,
moulting birds expend less energy per day than those engaged in all
other life-history activities (e.g. breeding, migration etc.; reviewed
in Jenni & Winkler 2020: 3.4.5), almost certainly because they can
offset a large part of the costs of moult by reducing other energy
expenditures, such as locomotion.
Feathers consist of 90 – 95 % protein with a higher proportion
of the essential sulphur amino-acid cysteine than is found in most
foods. They not only grow during the day when food is ingested,
but also at night, and some feathers are coloured with carotenoids
which cannot be synthesized by the bird, but must be obtained
from the food. This raises the questions as to whether moult is
constrained by nutritional requirements, and whether such a
constraint may be particularly pronounced during the night. The
acquisition of protein and sulphur-amino acids by birds feeding on
a ‘normal’ diet during moult does not seem to pose a problem, as
the amount ingested normally more than covers the needs of moult.
The growth of feathers during the night, when no dietary protein is
available, requires protein and glutathione to be temporarily stored
in the body during the day, when it can constitute 10 % of total
body protein (excluding feathers; Newton 1968b, Dolnik & Gavrilov
1979, Murphy 1996). Together with the increased whole-body
protein turnover, the additional protein synthesis and degradation
can be 8.5 times the amount of protein deposited as keratin each
day during peak moult in the White-crowned Sparrow Zonotrichia
leucophrys (Murphy 1996). However, the energetic cost of this
intensification of amino acid metabolism remains unknown.
The additional physiological processes during the moult period
result in a generally low efficiency of energy conversion to new
feather material. Two main processes have been proposed as
responsible for the added energetic costs of moult: (a) increased
whole-body protein metabolism; and (b) adaptation of the basal
metabolic rate in anti­cipation of reduced insulation and increased
thermoregulatory costs during moult.
During moult, the number of fully formed feathers is reduced and
this must affect plumage function. While the effects of moult on
water repellency and the effectiveness of display and status signalling
have not been assessed, the insulation provided by the plumage is
clearly compromised during feather replacement. Depending on the
number of simultaneously growing feathers, thermal conductance
in moulting birds is 30 – 60 % higher than at other times and this
may double the thermoregulatory costs and shift the lower critical
temperature during moult (Jenni & Winkler 2020: 3.4.2).
Moult clearly affects flight, and probably also aerobic capacity
(Buttemer et al. 2019), as shown by the well-known reluctance of
passerines to fly when moulting several remiges simultaneously.
In small passerines, simultaneous moult of more than about five
primaries results in serious flight impairment or almost complete
flightlessness (e.g. Newton 1966, Dolnik & Blyumental 1967, Haukioja
& Kalinainen 1968, Haukioja 1971b, Ginn 1975). However, birds
may have ways to mitigate the consequences of wing-gaps during
moult, such as managing the sequence of remige moult (see 3.2.2),
reducing body mass and increasing flight muscle mass. Furthermore,
  The Function and Consequences of Moult
birds may change their wingbeat frequency and kinematics in
flapping flight (e.g. amplitude, wing-tip reversal during the upstroke,
folding and extending the wing during up- and downstroke) and
how they use the tail (Chai 1997, Swaddle et al. 1999, Rayner
& Swaddle 2000). Hence, moulting birds appear to be able to
compensate, at least partly, for wing-gaps and a slow moult (2 – 3
simultaneously growing primaries per wing) may have no effect on
predation risk and predator evasion. However, the costs of impaired
flight during moult and the costs of compensations (e.g. changes in
wingbeat kinematics, follow-up costs of changes in body mass and
composition) remain largely unknown. Obviously, if wing-gaps are
large, complete compensation is not possible. A species’ requirement
for flight capability during moult largely determines moult duration,
and is therefore one of the most important determinants of the
moult strategy.
1.3.3 Conclusions
Assessing the actual costs of moult is very complex. All costs of
moult (energetic, physiological, plumage impairment) depend on
the intensity of moult; i.e. the number of simultaneously growing
feathers, and hence moult duration. Clearly, simultaneous moult of
too many feathers would be especially disadvantageous: (a) during
cold seasons; (b) during periods in which efficient flight is important
(e.g. on migration or when feeding young); (c) during courtship;
and (d) at times when territoriality or social hierarchies operate.
Moult duration varies greatly between species and depends on the
organization of a bird’s annual cycle (see 3.4.1).
The few indications available suggest that feather replacement
per se may not be much more costly than other forms of protein
synthesis. During moult, additional physiological processes give rise
to a considerable increase in metabolic rate during the night at
thermoneutrality, and these may double below the thermoneutral
zone.
Paradoxically, in free-living birds the apparently costly and
inefficient process of moult results in a period of minimal energy
expenditure, at least in those species examined so far. This is due
to many compensating mechanisms and because birds normally
do ‘nothing else’, or are unable to do ‘anything else’, during a
fairly rapid moult. A reduction in activity seems to be an important
measure to offset the energetic costs of moult and to mitigate
reduced or suppressed flight capability. In turn, this relative inactivity
during moult is probably actually caused by the many non-energetic
costs of moult such as impaired flight, reduced aerobic capacity
and impaired plumage functions (waterproofing, display, insulation,
etc.) which have potentially negative consequences for survival. It
therefore seems that in an ecological context these non-energetic
costs are more important than the energetic and nutritional costs.
Therefore, the validity of the common notion that moult is costly,
as stated frequently at the beginning of papers about moult, depends
on: (a) whether the author is looking only at feather replacement, or
all of the processes occurring during the moult period; (b) whether
the author considers overall investment or rate of moult; and (c)
whether the basis of comparison is more physiological or ecological,
either excluding or including compensations.
During a fairly intense moult the added energetic costs seem
to be largely compensated for, but both physical performance and
appearance are severely impaired. At the risk of exaggerating we
could say that moult is not costly, but is constraining.
7
1.4 Environmental effects on
new feather quality
Moult serves to optimize the quality of the plumage by replacing
feathers which have deteriorated since the last moult and, in many
cases, also to modify the plumage to serve new requirements (see 1.2).
However, environmental conditions may vary during moult and this
can lead to the production of poor quality replacement feathers. It is
important to realize that plumage quality and hence the appearance
of a bird, including its ornamental feathers, is partly determined
genetically and partly a product of environmental conditions during
the moult period, as well as how well a bird copes with these
conditions, and these effects are generally poorly understood.
Here we shall briefly discuss the environmental effects during
moult which can affect the quality of non-ornamental and orna­
mental feathers, and the consequences these may have. Again, more
information can be found in Jenni & Winkler (2020: chapter 4).
Various environmental factors acting during moult have been
shown to affect the structural quality and colour of feathers. These
include limits on the time available for moult, suboptimal nutrition,
stress events, parasites, immune challenges, and pollutants. The time
available for moult, which determines the extent and/or the speed
of moult, appears to have an overarching effect on plumage quality.
When time is short, partial moults are less extensive, and complete
moults are accelerated and produce feathers of reduced quality (e.g.
Dawson et al. 2000, Dawson 2004, Serra et al. 2010). In this context,
moult speed and feather growth-rate need to be clearly distinguished.
At the between-individual level, a high moult speed compromises
plumage quality, while a high feather growth-rate, as a consequence
of good individual condition, produces high quality feathers (Jenni &
Winkler 2020: 3.2.10).
The quality of the feathers produced during moult has been shown
to affect flight performance, insulation properties, and signalling
capacity while many other plumage functions have not yet been
examined (e.g. wettability, heat absorbance). Variation in plumage
quality, through its effect on plumage function, also affects survival,
sexual selection, reproduction, and hence fitness. Many studies have
investigated the effects of ornament quality on sexual selection and
the reproductive benefits to females, while studies investigating
plumage quality effects on survival are rare and correlative.
1.5 Fitting moult into the
annual cycle
Moult is commonly regarded as one of the major events in the annual
cycle of every bird, together with breeding, migration, and survival of
periods of food shortage or cold weather (Newton 2008). We have
seen how the process of moult entails costs as well as physiological
and behavioural constraints and therefore conflicts with the other
main stages of the annual cycle of a bird (see 1.3).
The declining quality of the feathers through wear increasingly
compromises fitness, as may a set of feathers adapted to past rather
than seasonally upcoming ecological conditions. Moult is therefore
8 The Function and Consequences of Moult
an inevitable maintenance and repair process which enhances survival
and reproduction, but which also results in competition for time,
energy and other resources between the need to re-establish feather
quality and the other annual life-history stages.
In order to find the optimal balance between moulting and the
other annual activities, a bird must, in effect, make difficult choices:
when and where is the most appropriate time to moult?; which
feathers should be replaced?; at which rate and in which sequence
should feathers be replaced?; and to what extent must plumage
performance be maintained during moult? It must also be able to
adapt the timing and speed of moult to its own particular annual
cycle and to changing environmental conditions. A bird’s particular
moult strategy is the outcome.
The moult strategies adopted by European passerines are actually
rather uniform compared with the diversity shown among birds in
general (see Jenni & Winkler 2020: chapter 5). All European passerines
usually renew the entire plumage at least once a year. Living in a
highly seasonal environment, they moult during distinct, annually
recurring periods. Most renew the entire plumage during a single,
relatively short period while only a few divide complete plumage
renewal between two moult periods. Other moult periods may be
added, usually so that part of the plumage can be renewed more
often than once a year. This is in contrast to large non-passerines
and certain tropical passerines which may moult more slowly over
longer periods, sometimes lasting for much of the year, and in which
the temporal division of a single moult cycle is more frequent (Jenni
& Winkler 2020: 5.3).
Two factors may account for the regular, distinct and restricted
moult periods typical of European passerines. First, due to their size,
small birds can moult completely relatively quickly while maintaining
a high degree of flight capability. Large birds, however, need longer
to perform a complete moult if they are to maintain a similar degree
of flight capability, or else become flightless during a rapid moult
(Jenni & Winkler 2020: 5.3.2).
Second, for birds in temperate areas, the seasons are marked
and predictable. This leads to a single, predictable and rather short
breeding season and, consequently, to a strict, regular organization
of the annual cycle. Furthermore, in a highly seasonal environment,
a favourable period after the breeding season is probably necessary
for successful post-fledging development before the onset of the
cold period or migration. This provides a window in which the adults
of most species can moult having no other major commitments and
in which the energetic costs of moult may be minimal. Alternatively,
long-distance migrants may delay their complete moult until after
reaching the winter quarters. In contrast, tropical species with
prolonged breeding seasons have a less strict organization of the
annual cycle and are more likely to moult slowly over a long and
variable period of the year which occasionally includes part of the
breeding season (Jenni & Winkler 2020: 5.2). Birds with unpredictable
breeding seasons may also moult slowly over much of the year,
including the breeding season, or may interrupt moult whenever
the opportunity to breed arises. In such species, the moult strategy
is less strict and less uniform within members of the species. Among
European passerines, a tendency for variable moult strategies is
apparent in the opportunistic breeder, Red Crossbill Loxia curvirostra
(see p. 265) and in migrants wintering in tropical areas (see 3.3 and
3.4.3).
Therefore, most European passerines have annual cycles with
regular, distinct and relatively short periods for breeding and
moult, rather than prolonged and overlapping ones and moult is
comparatively intense and the costs per day are higher. These intense
moults do not usually overlap significantly with breeding, migration
and wintering and this has often been explained by the competition
between moult and these three other events, notably for energy and
nutrients (Jenni & Winkler 2020: 5.2).
Despite the conflict between moult and breeding, some
overlap between the end of breeding and the onset of moult is
widespread (see 3.4.2; see Jenni & Winkler 2020: 5.2.1 and 5.2.2 for
explanations). In contrast, the temporal separation of flight-feather
moult and migration is rather strict (see 3.4.2), except in some short-
distance migrants and species which must maintain a high degree
of flight capability all year round (e.g. hirundines). This appears
primarily due to the conflict between moult and flight capability
and between opposing metabolic adjustments for moult and (long-
distance) migration (see Jenni & Winkler 2020: 5.2.3). Most European
passerines wintering in cold and temperate climates do not moult
during winter/spring or, at most, renew only a few feathers. The
few species which undergo a substantial pre-breeding moult may,
however, start before winter is over, despite the loss of insulation
involved.
Within a species, it appears that the timing, rate and extent of
moult can be adjusted according to a bird’s individual circumstan­
ces. Within the time window set by the endogenous annual cycle
and modified by the photoperiod, the precise timing and rate of the
post-breeding moult is thought to be fine-tuned by feedback from
perceived environmental factors (e.g. food availability, tempe­rature),
social cues (e.g. breeding success, number of broods, parental care
activities, interactions between mates) and factors modifying moult
once started (e.g. bad weather, unpredictable onset of breeding)
(Jenni & Winkler 2020: 3.5 and 5.3.7).
  Terminology and Methods in Moult Research 9

Chapter 2

Terminology and Methods in Moult Research

2.1 Arrangement of the feathers

In passerines, as in other birds, most of the feathers are contour-
feathers and these form the surface plumage of the full-grown bird
and give it its shape and colour. They number about 3500 in the
House Sparrow Passer domesticus (Staebler 1941) and 3300 in the
Song Thrush Turdus philomelos, but over 4600 in the diving White-
throated Dipper Cinclus cinclus (Glutz von Blotzheim & Bauer 1985)
(see Wetmore 1936 and Schieltz & Murphy 1997 for American
passerines). They are grouped into distinct tracts (pterylae) which
cover about half of the skin surface (Fig. 8). Eight major feather
tracts are usually recognized (Verbeek 1973, Ginn & Melville 1983,
Campbell & Lack 1985). The areas in between (apteria) are devoid
of contour feathers (though down feathers may be present) and are
usually covered by the feathers of the adjacent pterylae.
Dorsal view
capital tract
scapular tract, shoulder
Contour-feathers have closed vanes and a variable basal, downy
(plumulaceous) part (see Jenni & Winkler 2020 for feather structure
terminology). They vary enormously in size, shape and structure,
within and among species, according to their specific function. A
rough classification often used in moult studies distinguishes between
two main types of contour-feather. Small-feathers or body-feathers
cover the head and body, while the flight-feathers comprise the
large feathers of the wing (primaries, secondaries and tertials) and tail
(rectrices) (here we follow Lucas & Stettenheim 1972, Baumel 1993
and Lovette & Fitzpatrick 2016, while other authors only include the
remiges among the flight-feathers, e.g. Campbell & Lack 1985). The
small-feathers are reasonably symmetrical with almost equal vanes
and a plumulaceous part at the base, while flight-feathers usually
have a narrow outer vane, a broader inner vane and no plumulaceous
part. Associated with the flight-feathers, and intermediate in structure,
are the various wing-coverts, the alula and the tail-coverts. These are
usually considered a part of the small-feathers.
For the purpose of this book, we do not usually refer to the
individual feather tracts of the body (see e.g. Clench 1970, Verbeek
1973, Morlion & Vanparijs 1979, Baumel 1993), but use a detailed
classification of the feather rows of the wing, and refer to them as
feather tracts (see e.g. Zeidler 1966, Morlion & Vanparijs 1979 for a
detailed description).

dorsal or spinal tract 2.1.1 Flight-feathers

femoral tract, thigh

crural tract, lower leg The flight-feathers of the wing are called remiges (singular remex;
alar tract, remiges
and wing-coverts caudal tract, rectrices sometimes called wing-feathers). Those inserted onto the metacarpus
and tail-coverts and the digits (hand or manus) are the primaries, those on the ulna
(forearm) the secondaries and tertials (Fig. 9, 10).
Ventral view Passerines have ten primaries (P). The inner primaries, P 1 – 6, are
attached to the metacarpus, and the outer primaries, P 7 – 10, to the
capital tract digits: P 7 to the third digit, P 8 – 9 to the first phalanx of the second
digit and P 10 to the second phalanx of the second digit (Fig. 10;
Zeidler 1966, Stephan 2002, Eck et al. 2011). In some passerine species,
the outermost primary P 10 is about half as long as the adjacent one
(e.g. Eurasian Wren Troglodytes troglodytes), but in most passerine
ventral tract
species P 10 is very much reduced. In the Alaudidae, Hirundinidae,
Motacillidae, Passeridae, Fringillidae and Emberizidae, P 10 is extremely
small and difficult to detect under the outermost primary covert (Hall
crural tract, lower leg alar tract, remiges
and wing-coverts 2005). In Calyptomena broadbills, males lose juvenile P 5 permanently
caudal tract, rectrices and have only nine primaries, while females retain all ten primaries
and tail-coverts
(Wells 2011). The primaries are usually numbered descendently,
i.e. from inside (P 1) to outside, towards the wing-tip (P 10)
Fig. 8. Distribution of the eight major feather tracts (pterylae, dotted) on the dorsal
and ventral side in passerines. From Morlion (1971), with English names of tracts (Fig. 9, 12; Ashmole et al. 1961, Stresemann & Stresemann 1966,
added. Handbuch der Vögel Mitteleuropas by Glutz von Blotzheim et al.
 10 Terminology and Methods in Moult Research

1985 – 1997, Handbook of the Birds of Europe, the Middle East and arise when the outermost primary is reduced, and that the numbering
North Africa by Cramp et al. 1977 – 1994, Eck et al. 2011, and most corresponds to the normal sequence of primary moult which usually
of the other literature on moult), a convention which is followed in progresses descendently. The ascendent system of numbering
this book. In passerines this has the advantage that no difficulties primaries (from outside to inside) is used by some authors, especially
for indicating wing-formulae (e.g. Witherby et al. 1943, Vaurie 1959,
Stephan in Bub 1985, Svensson 1992, Demongin 2016, Shirihai &
Svensson 2018), while a few authors have (erroneously) used the
terms descendent and ascendent in an opposite manner to their
original definitions (see Winkler & Jenni 1996). As the increase or the
reduction of the number of primaries is confined to the metacarpus,
the ascendent numbering system preserves the homology of the four
outermost primaries, which are attached to the hand, in those non-
passerine species with nine or 11 primaries. It is considered more
practical for ringers, despite the difficulty of observing a reduced
outermost primary. Once familiar with the topography of the remiges,
it is not particularly difficult to find the innermost primary at the
carpal joint by manipulating the wing, since the primaries move in a
block. In any case, great care is needed when counting the primaries
of moulting birds, especially at the start of primary moult when just
the innermost primary is missing. It is important to state which system
of numbering is used in every published observation since papers
PC9 P10 PC8
describing exceptional moult patterns are still published without
9
CC
AI 2 3 giving this vital information.
7 8
1 6 The secondaries are numbered ascendently (from outside to inside)
PC
5
MaC
1
4
7 and in passerines there are usually nine, although some species have
2
C
Me 3 2
3 10 or 11. The six outer secondaries usually differ in shape and colour
4 6
5 1 P from the inner secondaries, which cover most of the primaries and the
1
8 7 6
2
GC
3
5 six outer secondaries when the wing is closed. Those inner secondaries
10
5
4 4 are called the tertials (T) (Hickman 2008) and are usually not moulted
9 8 7 6
T9 3 in sequence with secondaries 1 – 6. In this book, we use the term
S
2 secondaries (S) for the six outermost feathers only and always call
T8 1 the inner ones tertials, but number them T 7 to 9 (10, 11). Among
T7
1 European passerines, there are four or five tertials in the Alaudidae,
2
6 5 4 3 Oriolidae, Bombycillidae, and some members of the Corvidae (Stephan
1965, Stephan in Bub 1985). In the genus Lanius, which is reported
to have four tertials (Stephan 1965, Snow 1967), the innermost T 10
Fig. 9. Abbreviations and numbering of the feathers on the wing as used in this book,
demonstrated on the wing of a 2y  Northern Wheatear Oenanthe oenanthe. MaC = is a thin, downy structure hidden under greater covert 10 (own obs.).
marginal coverts. MeC = median coverts 1 – 8. GC = greater coverts 1 – 10. CC = carpal Most passerines have six pairs of tail-feathers or rectrices (R)
covert. Al = alula feathers 1 – 3. PC = primary coverts 1 – 9. T 7, T 8, T 9 = tertials 7 – 9.
S = secondaries 1 – 6. P = primaries 1 – 10. (singular rectrix). Among European passerines, only the Cetti’s Warbler

1 1 2 10
2 3
4 5 9
3 6 7 8
4
5 Fig. 10. This left wing of a Carrion Crow Corvus corone,
6
shown from the underside, shows which bones the flight-
11 feathers are attached to. All muscles and tendons have
7 been removed, as well as all of the feathers except the
10
9 8
primaries, secondaries, and longest alula. The humerus
and the radius are not coloured. The secondaries 1 – 11
are attached to the ulna (coloured brown; most passerines
have only 9 secondaries), primaries 1 – 6 to the meta­carpus
(carpometacarpus yellow), primary 7 to the third digit
(red), primaries 8 – 9 to the first phalanx of the second
digit (blue), primary 10 to the second phalanx of the
second digit (blue), and the alula to the first digit (green).
Preparation by R. Winkler.
  Terminology and Methods in Moult Research
Cettia cetti has five pairs. They are numbered centrifugally from the
inside to the outside.
Exceptionally, individuals may have supernumerary flight-feathers
on one or both sides, usually rectrices, but also primaries, secondaries
(Fig. 545) or tertials (Fig. 706) (for European passerines see e.g. Hanmer
1981, 1985, Copete et al. 1992, Stephan 1996, Fernandez Garcia 1999,
editor’s note to Benson 2003, Ottosson & Waldenström 2003, Roper
& Grantham 2005). Very rare are individuals with permanently missing
remiges (e.g. a secondary in Alauda arvensis, own obs.).
2.1.2 Wing-coverts
The gaps between the vane-less bases of the remiges are covered on
both sides of the wing by the tectrices (singular tectrix) or coverts. On
the upperside, the primary coverts (PC) are inserted just distally to
each primary (Fig. 11). PC 9 is very much reduced and covered by PC 8.
PC 10 is extremely small, or absent. A greater covert (GC) is inserted
just proximally to each secondary and tertial (Fig. 11). An additional,
innermost GC 10 has no corresponding tertial in species with three
tertials. Although numbered as the innermost within the row of greater
coverts, GC 10 inserts at a slight angle, is often smaller than the adjacent
greater coverts, and is often moulted out of sequence with the other
greater coverts. It is debatable whether GC 10 is actually a true greater
covert (Zeidler 1966).
Above the greater coverts, there are eight to nine median coverts
(MeC) and several rows of marginal coverts (MaC), the latter often
called lesser coverts in many publications. However, it is a characteristic
of passerines that the lesser coverts are absent or reduced to downy
feathers (Wray 1887, Reichling 1915, Steiner 1917, Zeidler 1966,
Stephan 1970, Winkler & Jenni 1996).
Between the primary and greater coverts lies the carpal covert (CC),
a covert belonging to the carpal remex which is absent in passerines
(Stephan 1974). The alula or bastard wing (Al) inserts at the first
digit and consists of three clearly visible feathers numbered from the
smallest to the largest. A fourth innermost alula feather is present in
all passerines but is much reduced and lies transverse to the three
conspicuous alula feathers (Mayr 1996). It is not considered in this
book. The alula completely covers the very small median primary coverts.
1 6
2 4
3 1 2 3
4
11 5
6
10 7
9 8
11
The underwing-coverts are not treated in this book (see e.g. Stübs
1972). They have been largely neglected in moult studies, except in
a few cases focussing on single species (e.g. Zeidler 1966, Winkler &
Winkler 1985) and in the work by Rymkevich et al. (1990) on a large
number of European passerines.
2.2 Terminology of plumages,
feather generations and
moults
2.2.1 Concepts of moult and plumage
terminologies
Most passerines of northern latitudes have distinct moult seasons
intimately related to the calendar year, resulting in a sequence of
plumages shared by members of a species or population which is
relatively easy to classify into a formal nomenclature of plumages
and moults. All plumage and moult terminologies currently in
use originated from moult studies on the better studied birds of
northern latitudes (e.g. Dwight 1900). However, tropical passerines,
opportunistic breeders and certain Palaearctic migrants wintering in
the tropics have moult periods and plumage cycles which are less or
hardly related to the yearly cycle or which exhibit large variation among
the members of a population (e.g. Stresemann & Stresemann 1966,
Foster 1974, Britton 1978, Zann 1985, Thompson 1988, Wikelski
et al. 2003, Johnson et al. 2012, Jahn et al. 2017, Red Crossbill
Loxia curvirostra, p. 265). Such plumage and moult cycles are much
more difficult to describe using the current terminologies. In non-
passerines, the variations in moult and plumage cycles are so diverse
that a convincing common nomenclature is even more difficult.
A terminology of plumages and moults could relate to the breeding
cycle (e.g. post-breeding, post-nuptial), to age and sexual maturity
(e.g. juvenile), to the season of the year (e.g. winter plumage),
9
78
5
Fig. 11. This right wing of a Eurasian Magpie Pica pica,
shown from the upperside, shows where the wing-
coverts are inserted. All of the marginal coverts have been
removed. The other coverts and the alula feathers are
trimmed leaving only a stump which is coloured red and
numbered for the 11 greater coverts GC (most passerines
have 10 GC) and the 9 primary coverts PC, and coloured
light blue for the 9 median coverts MeC, green for the
carpal covert CC, and yellow for the 3 alula feathers Al.
Preparation by R. Winkler.
 12 Terminology and Methods in Moult Research

to the aspect of the plumage (e.g. immature plumage) or to a
combination of these classifications. A terminology unhindered by
such relationships was introduced by Humphrey & Parkes (1959) and
defines a key plumage or a key moult (usually the complete moult)
from which homologous plumages and moults are derived. This so-
called ‘H-P system’ of naming moults and plumages has since been
debated (Miller 1961, Humphrey & Parkes 1963, Stresemann 1963a,
Amadon 1966, Willoughby 1991, Rohwer et al. 1992, Willoughby
1992, Thompson & Leu 1994, Filardi & Rohwer 2001), and was
subsequently revised (Howell et al. 2003) to produce the ‘H-P-H
system’. The ‘H-P or H-P-H system’ is now the common terminology
among American ornithologists, although it is still heavily debated
(Howell et al. 2004, Jenni & Winkler 2004, Thompson 2004,
Willoughby 2004, Pyle 2005, 2007, Hawkins 2011, Pyle 2013, Wolfe
et al. 2014, Howell & Pyle 2015). Meanwhile, European ornithologists
have largely retained a terminology relating moult and plumages to
the life-cycle of birds (see below).
Most terminologies correlate easily for the simple plumage
cycles of northern latitude passerines (Table 1). Difficulties only arise
when dealing with more complicated moult patterns and here each
terminology has its own drawbacks (see literature cited above).
If the purpose of a terminology for plumages and moults is
purely descriptive, it will usually be defined according to its primary
function. Preference may be given to age, sexual maturity and aspect
of the plumage (e.g. for field guides: juvenile, eclipse, immature
plumage) or to life-cycle stages (e.g. Anonymous 1985, Cramp 1988
p. 34 – 35), which is the system we adopt in this book (see below).
However, if the purpose of a terminology is to denote homologies of
plumages and moults, the explicit aim of the ‘H-P-H system’, major
difficulties arise, because neither the evolution nor the genetics
and control of moult and plumages are well enough known to
determine homologies (Amadon 1966, Jenni & Winkler 2004,
Willoughby 2004, Wolfe et al. 2014). Indeed, there is essentially
no existing evolutionary framework on which to build homologies
among plumages and moults and then to test them. The assumption
of the ‘H-P-H system’ is that the complete (or nearly complete) moult
of adults, which is repeated in the annual cycle, can be considered
homologous across species and that the common starting point
across species is the juvenile plumage, the first set of non-downy
feathers. Additional moults and plumages are then introduced into
the basic cycle, assuming that the simple cycle of one complete moult
annually is the ancestral pattern (Howell et al. 2003, Howell 2010).
However, as these authors admit, the juvenile plumage, as well as
additional plumages and moults, may or may not be homologous
across species. Further problems of finding homologies with the
‘H-P-H system’ are specified in Jenni & Winkler (2004), Thompson
(2004), Willoughby (2004), and Wolfe et al. (2014).
Uncovering homologies of moults and plumages is particularly
difficult, because, up to now, there are no firm and convincing criteria
on which to base a plausible evolution of plumages and moults.
Timing, extent and sequence of moult, and the colour of plumage,
are regulated by different factors and control mechanisms (see Jenni
& Winkler 2020: 3.5), so can be under different selective forces.
Therefore, it is highly debatable whether the evolution of moults is
always the same as the evolution of plumages, as the ‘H-P-H system’
pre-supposes by linking one moult to one plumage (Amadon 1966,
Jenni & Winkler 2004, Willoughby 2004).
An additional problem in establishing homologies of moults
and plumages lies in determining which parts of the plumage one
single moult comprises. While we would accept the idea that the
complete post-juvenile moult of the Common Starling Sturnus
vulgaris is homologous to the partial post-juvenile moult of the
European Robin Erithacus rubecula, there is, in our opinion, no
sound basis on which to determine the homologies between the
complete post-breeding and partial pre-breeding moult of e.g.
the White Wagtail Motacilla alba or the partial post-breeding and
complete pre-breeding moult of the Garden Warbler Sylvia borin.
Several species divide the moult of primaries and/or secondaries
into two temporally separate stages. Therefore, the two stages of
flight-feather moult may be regarded either as one or two moults.
In the terminologies based on homologous moults and plumages,
it is essential to know whether a feather renewal is merely the
continuation of a suspended moult or a separate moult. While this
question may be more easily resolved for a primary moult (e.g. Tawny
Pipit Anthus campestris), it is often much more difficult in secondary
and body-feather moults (see 3.3.2). In some species (e.g. Sedge
Warbler Acrocephalus schoenobaenus), it is still not clear whether
the complete moult in the non-breeding area is a separate moult
or whether it is the continuation of the partial body-feather moult
undertaken in the breeding area, perhaps even overlapping with a
pre-breeding moult. It is also conceivable that a moult split into two
stages in a species’ evolutionary history includes the renewal of some
body-feathers during the second stage which have already been
moulted during the first stage. The moult patterns of certain species
of passerines (e.g. Sylvia warblers) and non-passerines suggest to
us that the division of a formerly single moult into two temporally
separate moults, and the overlap of two formerly separate moults
or moult waves may be more common evolutionary processes than
hitherto recognized (Mulvihill & Rimmer 1997, Jenni & Winkler
2004). In this context, it is interesting that when Great Knots Calidris
tenuirostris were kept in constant environmental conditions, moult
of body-feathers and wing moult, commonly thought to be part of
a single, integrated annual event, become temporally dissociated
(Piersma et al. 2008). A similar, but less extensive dissociation was
also observed in captive Garden Warblers Sylvia borin and Red Knots
Calidris canutus (Gwinner 1986, Karagicheva et al. 2016).
A terminology based on homologies of plumages and moults
would certainly be useful. However, it would require very careful

This book
Growth of juvenile feathers
[of nestlings]
Juvenile plumage
Post-juvenile moult
Post-juvenile plumage
Post-breeding moult
Non-breeding or annual plumage
Pre-breeding moult
Breeding plumage
Humphrey & Parkes (1959) terminology
Growth of juvenile feathers
[of nestlings]
Juvenile or juvenal plumage
First prebasic molt
First basic plumage
Prebasic molt
Definitive basic plumage
Prealternate molt
Definitive alternate plumage
Howell et al. (2003) terminology
Prejuvenal or first prebasic molt
[of nestlings]
Juvenal or first basic plumage
Preformative molt
Formative plumage
Prebasic molt
Definitive basic plumage Table 1. Comparison of the most frequently used plumage
Prealternate molt and moult terms for passerines in this book with those
used by Humphrey & Parkes (1959), later modified by
Definitive alternate plumage Howell et al. (2003) and put into use by Pyle (2008).
  Terminology and Methods in Moult Research
application, and be of greatest value when investigating closely
related species (Rohwer et al. 1992, Thompson 2004), and then
only for species whose moults and plumages have been specifically
studied for homologies (e.g. Rohwer 1986, Thompson 1991, Young
1991), rather than just becoming an automatic replacement for
the old descriptive terminology (e.g. Pyle 1997). In our view, the
means of identifying homologies with any degree of confidence are
still poorly developed, at least for European species with complex
moult patterns. Therefore, adopting a general moult and plumage
terminology based on the homology concept seems premature so,
for general descriptive purposes, we advocate the use of the more
traditional terminology. The challenge for future specialist moult
studies will be to evaluate evolutionary homologies, functional
adaptations, and their various interrelations. For this, we need a
more open and flexible approach than the ‘H-P-H system’ in its
current form, which would allow us to dissociate the evolution of
plumages and of moults, and take into account temporal splits,
fusions and overlaps of moults.
Because we are dealing with passerines of temperate areas, a
terminology which relates the plumages and moults to the life-cycle
of the bird is appropriate. The breeding season is a distinct and
pivotal annual event and we have adopted the general terminology
used by Roselaar (in Cramp 1988 p. 34 – 35; similar to Amadon 1966
with modifications; and similar to the Handbook of the Birds of the
World, del Hoyo et al. 2003 – 2011) which relates the plumages
and moults to the breeding cycle, without any claim to designate
homologies. However, we differ from Roselaar, in all those cases in
which it remains uncertain whether the moult preceding autumn
migration is resumed in the wintering area. In such cases we have
called the moult of long-distance migrants which precedes autumn
migration the post-breeding/post-juvenile moult, and all subsequent
moults in the wintering area the pre-breeding moult.
2.2.2 General terms
The term plumage refers to all the feathers normally covering a bird
for an appreciable length of time (Amadon 1966). During moult, birds
are in transition from one plumage to the next. The change from
one plumage to the next can only occur by moult, not by feather
wear changing the plumage appearance. Usually, only the contour
feathers (flight- and body-feathers) are taken into account, but not
the downy feathers. A plumage may be composed of one or several
feather generations. Plumage is used similarly by Cramp (1988, p.
34 – 35) and many others and is included in the more broadly defined
terms ‘feather coat’ and ‘aspect’ of Humphrey & Parkes (1959) and
‘feathering’ of Palmer (1972).
The term feather generation usually designates the feathers
acquired during a single moult, consisting of anything from only a
few to all the feathers of a bird. After a complete moult the bird’s
plumage consists of one feather generation, while after a partial
moult at least two generations are present (the new one plus at least
one older one). Feather generation is defined similarly by Amadon
(1966) and Palmer (1972) and equals the term ‘plumage’ as used by
Humphrey & Parkes (1959). Difficulties arise when a moult is halted
temporarily and resumed later at the point of interruption. Is each
‘resumption moult’ the continuation of a temporarily halted moult
wave of the same feather generation, or the start of a new feather
generation (i.e. are all feathers produced during a moult period of the
same feather generation)? We advocate that it is the same feather
13
generation that needs two (or more) moult periods to be formed (see
Jenni & Winkler 2020: 3.1.3 for more explanations).
Moult is the natural replacement of feathers by a new feather
generation, i.e. the normal shedding of feathers and replacement
by a new feather generation. More than one feather generation
may be replaced in one moult. A single moult may be discontinuous
(suspended), occurring in two temporally separated stages. However,
because in many species it is not clear whether the second stage of
an interrupted moult is really the continuation of the first rather than
a separate moult event, we sometimes have to give the two parts of
an interrupted moult different names.
We define moult strategy broadly to describe how birds fit moult
into their annual cycle (timing of moult relative to breeding and
migration) and how they perform this moult with regard to extent,
speed, and sequence (see Jenni & Winkler 2020: 3.1.4). The moult
strategy encompasses the moults of adults, as well as the particular
moults of young birds until they adopt the adult moult strategy.
We use the term moult period to refer to the time periods during
which a bird is moulting (e.g. as Shugart & Rohwer 1996). Moult
periods can be only a few weeks long when moult is rapid or of
limited extent, or encompass most of the year when moult is very
protracted. The term moult type is used in the context of ageing
European passerines (see 5.3).
2.2.3 Moult terms
The first true feathers acquired by a nestling passerine bird (the
first set of non-downy feathers) constitute the juvenile plumage.
Therefore, at fledging (the moment at which a feathered young
passerine leaves the nest) it is wearing its juvenile plumage. The
first moult after fledging, which replaces part, or all, of the juvenile
plumage, is the post-juvenile moult. In European passerines,
this usually occurs during the first summer/autumn of a bird’s life.
Later moults start either before the breeding season in winter/
spring or after the breeding season during late summer/autumn
and are termed pre-breeding moult and post-breeding moult,
respectively. In many species, only one of these moults occurs,
usually the post-breeding moult. Normally, the first pre-breeding
and post-breeding moults during the life of a bird are similar to
subsequent pre-breeding and post-breeding moults, respectively. In
certain species, however, they differ in extent or coloration of the
feathers they produce and have to be explicitly termed first pre-
breeding moult and first post-breeding moult.
Any of the above moults may be of very different extent. A moult
comprising the entire plumage is termed a complete moult. We
use this expression in a strict sense and not for cases in which a
regular descendent primary moult began but was not obviously
finished. If, however, only one or a few individual feathers (e.g. a
single alula feather, a few body-feathers, or a single secondary) have
been retained exceptionally, we would still call the moult complete.
A single moult often does not involve the entire plumage and is then
called a partial moult.
Following Harper (1984), the phenomenon by which a moult
stops before the entire plumage has been renewed is called moult
interruption. This term and the related terms suspended and arrested
moult have generally been used only for primary and secondary moult
and, in order to prevent confusion, we retain this usage. Two different
types of primary and secondary moult interruption are recognized
(King 1972, Harper 1984). In the case of suspended moult,
 14 Terminology and Methods in Moult Research
127 mm
t of primaries in passerines. Ascendent is from outside to
d en
en
sc nt
de ge
r
dive nt
rge
n ve
ascendent co
the primary and/or secondary moult is subsequently resumed at the
point of interruption. Therefore, the suspension normally divides a
single moult into two temporally separated stages. Those primaries
and secondaries not moulted during the first stage of the moult
are renewed later during the second stage. In the case of arrested
moult, the next primary and secondary moult starts at the normal
site of initiation or another site other than the point of interruption.
Therefore, an arrested moult cannot later proceed to completion and
the subsequent moult is in fact a different moult entirely. For the use
of the terms seasonally divided moult, split moult and interlaced
moults, see 3.3.3.
In the case of eccentric primary moult, the renewal of the
primaries does not start with the innermost or outermost primary,
but with a central feather and is interrupted (see 4.3.3). The next
primary moult usually starts at the normal site of moult initiation
(usually P 1). Eccentric primary moult is still poorly understood and
may comprise several patterns of primary moult including several
directions of progression. Therefore, we do not know whether it
is always an arrested moult, whether resumption at the point of
interruption may occur, or whether a divergent sequence may even
produce a complete primary moult.
We use the term moult limit to describe the point where two
feather generations meet within or between any of the feather tracts,
since the terms moult interruption and arrested moult are reserved
for primary and secondary moult interruption only. Some authors
use the term moult contrast instead of moult limit (Lewington et al.
1991, Alström et al. 2003, Howell 2010).
Within the feather tracts of the wing, moult may proceed in
various sequences: descendent (from the body towards the wing-tip),
ascendent (from the wing-tip towards the body), convergent (from
both the body and the wing-tip towards the centre) or divergent
(from the centre both towards the body and the wing-tip) (Fig. 12).
For the rectrices, the terms centrifugal (from the central pair of
rectrices, R 1, towards the outside), rather than divergent, and
centripetal (from the outermost pair of rectrices, R 6, towards the
centre), rather than convergent, are used. We prefer the spelling
‘descendent’ and ‘ascendent’, as in Campbell & Lack (1985), versus
descendant and ascendant as occasionally also used.
Fig. 12. Terms used to indicate the direction in which
flight-feathers of a given tract are numbered or moulted.
Descendent is from inside to outside, from the body
towards the wing-tip, i.e. from proximal to distal. The start
is usually at the innermost feather of a tract. Descendent is
the usual direction of numbering and sequence of moult
inside, from the wing-tip towards the body, i.e. from distal
to proximal. Ascendent is the usual direction of numbering
and the most common direction of moult of secondaries
1 – 6 in passerines. Divergent is from a focal point in both
centrifugal
directions, i.e. from somewhere within the tract both
towards the body and the wing-tip, while convergent is
from both the body and the wing-tip towards the centre.
centripetal For the rectrices, the terms used are centrifugal (from the
central pair of rectrices, R 1, towards the outside), rather
than divergent, and centripetal (from the outermost pair of
rectrices, R 6, towards the centre), rather than convergent.
2.2.4 Terms for plumages, feathers and feather
generations
After fledging, all passerines wear the juvenile plumage. If there
are two moults annually, the bird enters its breeding plumage after
the pre-breeding moult and its non-breeding plumage after the
post-breeding moult. If there is only one moult each year these terms
are unnecessary, though the plumage may be described as annual.
For individual feathers or feather generations, we refer to the
moult at which these feathers were produced, e.g. a greater covert
formed during the post-juvenile moult is termed a post-juvenile
greater covert.
Since partial moult is a frequent phenomenon, the plumage of a
bird is frequently composed of two or even three feather generations,
each acquired during a separate past moult. In order to designate
plumages more precisely for the purpose of this book, we prefer to
refer to the extent of the past or current moult e.g. ‘after partial post-
juvenile moult’, ‘after complete post-breeding moult’, ‘in suspended
post-breeding moult, ‘in post-juvenile moult’ etc., and do not use
the plumage terms of Cramp (1988, p. 34 – 35).
2.2.5 Age classes
We use the age classification which refers to the calendar year.
Birds fledged in the breeding season of the current year (up to 31
December) are first-year birds (1y, EURING Code 3, HY in Pyle
1997). Birds fledged in the previous calendar year and now in their
second calendar year (from 1 January) are second-year birds (2y,
EURING Code 5, SY in Pyle 1997). 1y/2y birds are usually recognizable
up to the first complete moult, after which they are called adults
(ad), i.e. birds fledged before the current calendar year (EURING Code
4, AHS in Pyle 1997) and birds fledged before the previous calendar
year (EURING Code 6, ASY in Pyle 1997) but whose precise year of
fledging is otherwise unknown.
Because an age classification referring to the calendar year cannot
readily be used in species with a breeding season that includes
  Terminology and Methods in Moult Research
1 January, age classifications based on moult and plumage criteria
have been developed for such cases (Jackson 2001, Wolfe et al. 2010,
Johnson et al. 2011, Clark & Pyle 2015). This is mainly intended
for tropical and southern hemisphere species. Among European
passerines, only the Red Crossbill Loxia curvirostra has a breeding
season that includes 1 January.
2.3 Recording moult, plumage
and feather properties 
2.3.1 What to record: moult progress or moult
intensity and of which feather tract?
Moult data are usually collected to study one or several of four topics:
(a) seasonal timing of moult; (b) duration of moult, and its speed and
intensity; (c) sequence of moult; and (d) the final extent of moult. In
many cases the aim is to compare species, individuals, or populations
or groups thereof (e.g. sexes, age classes, breeders/non-breeders). For
some studies, knowing that a bird is in full, active complete moult may
suffice, e.g. when comparing the physiology or behaviour of moulting
with non-moulting birds. Depending on the purposes of the study
additional data may help to analyse and interpret moult patterns, such
as the mass of feathers (often required for the analysis of moult data,
see 2.4.1), the degree of feather abrasion, or the colour and physical
quality of feathers (e.g. stiffness, presence of fault bars).
Seasonal timing, duration and sequence of moult are usually
derived from point-in-time records of the state of moult of individuals
examined once or repeatedly, and important decisions need to be
made as to exactly which features to include.
The first is to decide which feathers to record and in what detail.
This entirely depends on the purpose of the study and below we
describe ways to record the moult of remiges, wing-coverts and
15
50
lt
40 ou
f m
sso
og
re
pr
Primary moult score
30
4
Number of growing primaries
20 3
in
te
ns
ity
of 2
m
10 ou
lt
1
0 0
0 1
Proportion of time through moult
Fig. 13. Schematic representation of the progress and intensity of primary moult. The
black line indicates the progress of moult, as indicated by the primary moult score (left
axis). Primary moult score is zero before the first primary is shed and 50 when all ten
primaries are fully grown. Primary moult score typically does not increase perfectly
linearly (see 2.4.1). The grey line indicates the intensity of primary moult, given here
as the number of primaries simultaneously growing (right axis). At the beginning of
moult, inner primaries are generally shed in shorter succession than outer primaries,
which results in an asymmetric curve (see 2.4.1).
body-feathers in various levels of detail, from single feathers to entire
tracts or the entire body. However, it is important to state clearly
which feather tracts have been considered and which have not. There
are still studies published on ‘the moult’ of a particular species, which
only present data on the primaries or the remiges, while the wing-
coverts or the bulk of the feather material of a complete moult, the
body-feathers, are not considered.
Secondly, any moult study should be clear about whether the
progress of moult or its intensity, or both, are of interest. In an actively
moulting bird, a measure of the progress of moult, as defined
here, identifies the proportion of the plumage or a feather tract that
has been renewed (or is still old) at the time of recording. After the
Fig. 14. The Moult Record Card used by the British Trust
for Ornithology (reproduced with permission from BTO).
The scores are defined as: 0 = old feather remaining;
1 = old feather missing or new feather completely in pin;
2 = new feather just emerging from sheath or up to one
third grown; 3 = new feather between one and two thirds
grown; 4 = new feather from two thirds to fully grown, but
with remains of waxy sheath persisting; 5 = new feather
fully developed with no trace of sheath remaining at base;
and 8 = full grown feather, age uncertain.
 16 Terminology and Methods in Moult Research

Fig. 15. The moult card developed by the Swiss Ornithological Institute and the Natural History Museum of Basel. The card is laid out like the two wings of a bird in the hand with the
head pointing away from the ringer, as a means of reducing mistakes in recording. The scores 0 – 5 are defined following the BTO moult card (Fig. 14). Additional scores are:
6 = renewed after autumn migration during (late autumn) winter/spring (‘pre-breeding’ moult).
7 = in winter/spring: Older than 6, either 0 or 5. This code may be used in winter/spring for feathers which appear much older than 6, but where it is uncertain whether they have
been acquired during the post-juvenile/post-breeding moult or earlier. In summer/autumn: Older than 0. This code may be used in late summer/autumn for feathers which have been
retained during the previous pre-breeding moult (e.g. adult Spotted Flycatcher Muscicapa striata, Eurasian Golden Oriole Oriolus oriolus).
8 = older than 6, either 5 or ‘early 6’. This code may be used in winter/spring for feathers which appear somewhat older than 6, but where it is uncertain whether they have been
acquired during the post-breeding/post-juvenile moult before autumn migration or during an early ‘pre-breeding’ moult in late autumn/winter.
9 = impossible to assign.
For body-feathers, marginal and median coverts, the approximate percentage of old, new and growing feathers is given. If body-feathers are composed of three generations, the
additional column may be used with the corresponding explanations in the comments section.

moult, this then becomes the extent of moult, which might
be complete or partial for the entire plumage or a feather tract.
Therefore, the progress of a single moult increases continuously, but
not necessarily linearly, with time from 0 at the start to the proportion
attained at the end of moult (1 in the case of a complete moult) (Fig.
13). A measure of the intensity of moult, as defined here, identifies
the proportion of the feathers of the plumage or a tract that are
growing concurrently at the time of recording. Therefore, the intensity
of a single moult over time starts at 0, attains some value during
the peak of moult and decreases to 0 at the end of moult (Fig. 13).
The distinction between progress and intensity is crucial, yet we
have seen many manuscripts which interpreted measures of intensity
as progress (less often vice versa). It is therefore important to decide
which aspect(s) to focus on. For example, a study on the costs of
moult would probably focus on the intensity of moult, while a study
on the relationship between age and the speed of post-juvenile moult
would probably focus on the progress of moult.
2.3.2 Scoring flight-feather moult
The most commonly used system of recording the moult of remiges
and rectrices was introduced in the 1960s and has been used widely
in Britain and elsewhere (Snow 1967, Ginn 1973, Ginn & Melville
1983, Redfern & Clark 2001). Each feather is scored either 0 (‘old
feather remaining’), 5 (‘new feather fully developed with no trace of
waxy sheath remaining at base’), or from 1 to 4, depending on its
length relative to its full length (Fig. 14). It is evident that great care
is needed to count and identify the feathers correctly, to be aware of
whether P10 is much reduced or not, and to unambiguously identify
missing feathers. For this, both primaries and secondaries need to be
counted to see whether the correct number is reached, even if only
primaries are of interest.
The concept of scoring each individual primary was apparently
developed by Williamson (1957, 1962) for use in the British Trust
for Ornithology Moult Enquiry (Williamson 1960), as well as by
Ashmole (1962) (using slightly different scores), while Miller (1961)
scored primary moult over the entire wing using the number of
remaining old primaries (therefore recording progress of moult, similar
to the primary moult score; see below). At a conference in 1971,
European ornithologists agreed on the scores 0 – 5, which became
the standard, and on a common moult record card (Ginn 1973)
which has subsequently been expanded (Ginn & Melville 1983; Fig.
14). Newton (1966) used a somewhat different scoring system, using
score 5 for feathers of three-quarters to full length (subsequently
used by Newton & Rothery 2000, Rothery et al. 2001, Newton &
Rothery 2005, 2009). We recommend using the standard system with
score 5 exclusively for new, non-growing feathers, because grouping
growing and full-grown feathers into score 5 may cause difficulties
with statistical analyses, which may partly explain differences in the
estimated parameters between various methods of analysis (Newton
& Rothery 2000, Rothery et al. 2001).
Alternatively, growing feathers can be scored directly as proportions
of their full length between 0 and 1 (fractional scores), with
  Terminology and Methods in Moult Research
dropped feathers scored as 0.01 and almost fully grown and full-
grown feathers with sheathing at their bases as 0.99 (Rohwer 2008).
Additional scores need to be added to describe more complicated
moults involving more than two feather generations, suspended
and incomplete moults (Fig. 15). Assigning full-grown feathers
to the correct feather generation in species with multiple feather
generations present (e.g. Common Whitethroat Sylvia communis,
many large non-passerines) can be very difficult, because the flight-
feathers wear differently across the wing. In large species with a
serial moult, the different feather generations present in one wing
are extremely difficult to discern in birds more than a few years old
(e.g. Langston & Rohwer 1995) and the full pattern is best revealed
by examining the same individual over many years (see White Stork
Ciconia ciconia in Bloesch et al. 1977 and Sutter 1984; Eurasian Eagle-
Owl Bubo bubo in Glutz von Blotzheim & Bauer 1980). The use of
black ultraviolet light fluorescence helps in distinguishing feather
generations in species with porphyrin pigments (particularly owls),
which are quickly destroyed by sunlight (Weidensaul et al. 2011).
The moult of the wing-coverts is scored in various ways, either
per individual feather or collectively (Fig. 14, 15), generally as one of
three stages (old, growing, new).
Especially for birds undergoing a complete moult, or for
complicated moult patterns, it is handy to use a moult record card
(Fig. 14, 15). For more simple patterns or for recording a restricted
number of feather tracts, other systems can be devised.
Obviously, the standard scores give only approximations of the
length of a growing feather (and its mass, see 2.3.4) and have been
criticized on this basis (e.g. Seel 1976). In dead birds, individual
feather lengths can be measured accurately but this is usually too
time-consuming and certainly quite stressful for live birds. However,
by correcting the standard moult scores for feather mass, very
satisfactory results can be obtained (see 2.3.4. and 2.4.1).
If the state of moult is recorded for each feather separately, at least
to the distinction of old, growing, and renewed feathers, both moult
progress and moult intensity can be derived. If old and new feathers
are not distinguished (growing versus non-growing), moult progress
cannot be derived, except when birds are examined before and after
moult in clearly differentiated time frames. If feather tracts are scored
collectively, two scores are needed: one estimating the proportion of
growing feathers and a second estimating the proportion of renewed
(growing or fully grown) feathers, if both moult progress and moult
intensity are of interest.
Flight-feather moult is scored most easily in live birds in the
hand or from open-wing preparations, but is much more difficult
to discern in museum skins (see recommendations in Stresemann
91 mm
140
120
100
80
Mass (mg)
60
40
20
0
1 2 3 4 5 6 10 9 8 7 6 5 4 3 2 1 1 2 3 4 5 6 7 8 9 10
Rectrices Tertials Secondaries Primaries
Fig. 16. Mean mass (mg ± s.d.) of primaries, secondaries, tertials, and rectrices of 10
post-juvenile male Eurasian Jays Garrulus glandarius. Data from Seel (1976).
17
& Stresemann 1966). In birds frequently seen flying in the open,
photographs may also allow precise scoring of flight-feathers (Snyder
et al. 1987, Conklin & Battley 2012, Ryan 2013, Vieira et al. 2017,
van Bemmelen et al. 2018, Zuberogoitia et al. 2018). In caged birds,
and other exceptional cases, timing and sequence of moult can be
described by the daily collection of shed feathers that can be safely
attributed to an individual (Ginn & Melville 1983, Winkler et al. 1988).
2.3.3 Scoring body-feather moult
There is no standard method for scoring the moult of feathers of
the body and head (collectively named body-feathers) and various
methods have been used. Because body-feathers cannot be assessed
individually, but are scored collectively per feather tract, two scores are
needed, one estimating moult intensity as the proportion of growing
feathers, and one estimating moult progress as the proportion of
renewed (growing and fully grown) feathers. The distinction between
juvenile and post-juvenile body-feathers can be a challenge in some
species, and therefore may not be practical (e.g. Norman 1981).
The moult of body-feathers can be recorded for the entire bird, or
for only the underparts (Bensch & Lindström 1992), or for a few larger
parts of the body (e.g. head, upperparts, underparts, tail-coverts,
and wing-coverts; Norman 1981, 1990a; see also Herremans 1987),
or in more or less detail for either six (Willoughby et al. 2002), nine
(Newton 1966, Bojarinova et al. 1999), 12 (Seel 1976), 16 (Winkler
& Winkler 1985), 21 (Berthold et al. 1970; plus six for wing-coverts)
or even 28 feather tracts on the body, as well as for the head and
separately for the different tracts of the wing including the underwing-
coverts (Noskov & Gaginskaya 1972, Rymkevich et al. 1990). Indeed,
even more systems have been adopted in various studies. If there is
a distinct sequence of moult of body-feather tracts, different stages
of moult progress can be defined depending on which tract shows
growing feathers (e.g. Newton 1966, Ryzhanovsky 2010).
For studies which do not go into the details of body-feather moult,
a simple system is proposed (Fig. 15) which records the proportion of
old, new (full-grown) and growing feathers for the head, upperparts,
underparts and tail-coverts. This system allows a rough assessment
of moult progress (proportion of new and growing feathers) and
intensity (proportion of growing feathers). The determination of old,
new and growing feathers can however be difficult for the four areas
of the body. In any case, the particular aim of a study dictates the
degree of detail needed. A mixture of codes, mainly for the progress
of body-feather moult, but also including active wing-feather moult,
is proposed by Redfern & Clark (2001).
2.3.4 Mass of feathers
For many analyses of moult, the mass of the feathers is needed, in
particular the mass of the individual primaries (see 2.4.1) and this
can be obtained by weighing plucked feathers from dead individuals
or shed feathers from live birds (Fig. 16). Only full-grown feathers
with little or no wear or damage should be taken, cleaned and dried
to constant mass and weighed immediately (to 0.1 mg or 0.01 mg)
(e.g. Summers et al. 1983, Underhill & Joubert 1995). Because in
many instances only the relative mass is of interest, feathers may
also be allowed to equilibrate to room temperature and humidity
before weighing (e.g. Dawson 2005). For some species the (relative)
mass of the primaries has been published (e.g. Seel 1976, Marks
 18 Terminology and Methods in Moult Research

A B

C D

Fig. 17. Feather wear can be easily scored with the naked eye or a magnifying glass into four classes, demonstrated here using examples of Common Whitethroats Sylvia communis:
A: no indication of wear; B: slight wear at the P-tips, visible on close examination; C: markedly worn feather tips and outer edges (e.g. GC), old feathers bleached and contrasting with
the renewed S 6 and T 7 – 9; D: heavily worn with broken tips of rachis which may affect feather length measurements, in this example the contrast in bleaching between renewed S
6 and T 7 – 9, and older S and P is not very marked. Depending on the species, the feathers affected by wear and the kind of wear (fading, abrasion, breakage) varies.

1993, Underhill & Joubert 1995, Dawson & Newton 2004, Dawson
2005, Oschadleus & Underhill 2006, Buchmann et al. 2009,
Remisiewicz et al. 2009, Young et al. 2009, Boano et al. 2015). In
this context it is interesting to note that the primary forming the
wing-tip and all those distal to it (if any) are heavier than predicted
from the allometric relationship between feather mass and feather
length (Dawson 2005). If the relative feather mass of the primaries
is not available, see Bonnevie (2010a) for the use of mean values for
each wing-shape type.
Because the mass of a feather is not equally distributed along the
length of the feather (one cm of feather tip weighs less than one cm
of the base), more refined analyses use the mass of individual feather
segments (Dawson 2003). However, such refinement is not usually
required for most analyses (see 2.4.1).

127 mm
mm

Total length
Shedding of old feather

50
Fig. 18. Growth curve of a primary, illustrated by a 75 mm
long primary 5 of an Eurasian Skylark Alauda arvensis.
The total length including the part in the skin is given by
Length of sheath the triangles; the surface of the skin is indicated by the
horizontal broken line. The length of the feather sheath is
indicated by open circles; the length of the pulpa (blood
quill) by dots. The arrow indicates the time of shedding
Length of pulpa Surface of the skin of the old feather.
Letters indicate the four stages of growth: (a) a short
0 period of slow growth in length (2 days); (b) a long period
0 10 20 30 days of essentially linear growth in length (ca. 14 days); (c) a
gradual slow-down in growth until almost final length (ca.
a b c d
Stages of growth 4 days); and (d) a period of keratinization of the calamus
with hardly any further growth in length (ca. 5 days).
Duration of feather growth Below the graph, two methods of measuring feather
17 days
as measured by moult scores growth are indicated: duration of feather growth as
measured by the moult scores 1 – 4; and duration of the
Duration of the whole entire measurable growth process. Redrawn from Winkler
24 – 25 days
measurable growth process et al. (1988), modified, with permission from Ala.
  Terminology and Methods in Moult Research
Fig. 19. Growth bars on the tail of a Common Grasshopper Warbler Locustella naevia
which are exceptionally clearly visible. Each pair of light and dark bars on R 1, 2 and 3
has a mean width of 2.7 mm corresponding to the daily growth increment.
2.3.5 Quantification of feather wear
Estimating the degree of wear (which includes damage by sunlight,
friction and use in flight, see Jenni & Winkler 2020: 2.1) may be useful
for two reasons: first to determine when wear occurs and which
feathers on a bird are most exposed to wear, and may therefore be
moulted preferentially; and second to differentiate between feather
generations. Wear of primaries also affects wing length (see Jenni &
Winkler 2020: 2.1.5).
Various systems of scoring wear (also termed abrasion, Ginn &
Melville 1983, Rogers 1990) have been devised for remiges, rectrices,
wing-coverts and body-feathers. Scoring wear with the naked eye
or using a magnifying glass has generally used four classes of wear
from no wear to heavily worn (Prater & Marchant 1977, Willoughby
1986, Rogers 1990, Willoughby 1991, Martin 1996, Willoughby et al.
2002, Vágási et al. 2011 and references therein; Fig. 17). For feathers
suffering from particularly strong wear and breakage (e.g. the long
tail feathers of the Eurasian Magpie Pica pica), an extended scoring
system, from intact to feather stump, has been used (Fitzpatrick &
Price 1997). If dead birds are available or pieces of feathers are cut
off, wear can be examined under a microscope for the degree of wear
of barbs, barbules and hooks on the inner and outer vane (scored
from 1 to 3) (Sæther et al. 1994), and for more details of feather
microstructure (e.g. Willoughby et al. 2002, cf. Butler et al. 2008).
Whether scoring wear to differentiate between feather generations
is successful depends very much on an in-depth appreciation of
variation in wear between individuals, and variation in wear between
feathers of the same generation on the same individual. Exposed
feathers wear much faster and further than feathers protected by
19
A B
Fig. 20. A: Growth bars on a secondary feather of an Indian Peafowl Pavo cristatus (total
length without calamus 305 mm). Each pair of light and dark bands is approximately
5.5 mm in width (four of them indicated with red lines) and corresponds to the daily
growth increment. B: Bars produced by the stepwise retreat of the pulpa and the
production of septa or pulpa cap roofs on a secondary flight-feather of an Andean
Condor Vultur gryphus (total length without calamus 355 mm). The pulpa cap bars are
clearly visible on the outer vane and are about 3 mm in width (four of them indicated
with red lines). They are produced more than once per day so do not represent daily
growth increments. In this example the septa bars are exceptionally clearly visible on the
vane; in most cases, however, they are more easily seen on the rachis, if visible at all. In
the calamus, the septa (also called Federseele) are well-preserved and can be seen by
holding the calamus against a light, or by dissection.
others – and wear also depends on the habitat and behaviour of
the bird (cf. Fig. 3 – 6). Therefore, we think that scoring wear to
differentiate between feather generations can be used in special
cases only.
2.3.6 Feather growth rates
The growth rate of individual feathers is only loosely related to overall
speed of moult (Rohwer & Rohwer 2013, Jenni & Winkler 2020:
3.2.10). Therefore, feather growth rates cannot readily be derived
from moult speed and must be measured separately.
The growth of an individual feather, as shown by the primaries and
other flight-feathers, shows essentially four stages: (a) a short period
of slow growth in length when the new feather is still mostly buried
in the skin; (b) a long period of essentially linear growth in length;
(c) a gradual slow-down in growth until almost final length; and (d)
a period of keratinization of the calamus with hardly any further
growth in length (Fig. 18). Depending on the method of measuring
growth rate, different stages of feather growth may be recognised
(see Fig. 18) and growth rates obtained using different methods are
therefore not necessarily comparable.
The most common method of determining feather growth rate is
to measure a growing feather repeatedly over time with a thin ruler
or folded millimetre paper inserted distally of the feather (see e.g.
Jenni & Winkler 1989). If this is done during stage (b), near-maximum
growth rates are obtained and these probably represent the majority
of growth rates reported in the literature (e.g. Prevost 1983, Rohwer
et al. 2009). If feathers are monitored until full-length is attained, only
 20 Terminology and Methods in Moult Research
stage (a) is normally missed (see e.g. Newton 1967). Be aware that the
length of a feather measured on a live bird is from the skin at feather
insertion to the tip and misses the part which is buried in the skin
(ca. 10 % of the full length of flight-feathers in corvids; Seel 1976).
Another method of determining feather growth rate is based
on counting the daily growth bars on full-grown feathers. Growth
bars, also termed growth bands, are pairs of alternate light/dark
bands which run almost perpendicular to the rachis (actually at a
slight backward angle) across both the vanes and the rachis (Riddle
1907, 1908; Fig. 19). A pair of light/dark bands is produced every
24 h, the dark band during the day and the light band during the
night (Riddle 1907, 1908, Michener & Michener 1938, Wood 1950,
Brodin 1993, Grubb 1995). However, approximately two pairs of
light/dark bars per day have also been reported (Langston & Rohwer
1996, Kern & Cowie 2002, Jovani et al. 2011, Elderbrock et al. 2012)
although these may have been confused with a second set of bars
across the feather produced by the stepwise retreat of the pulpa and
the production of septa, also termed pulpa cap roofs (Lüdicke 1965,
1966, Lucas & Stettenheim 1972, Alibardi 2009; Fig. 20). This process
usually occurs more than once per day and the daily growth bars and
septa bars are visible under different light angles (Winkler et al. 1988).
Instructions on how to measure growth bars usually recommend
measuring 10 bars in the middle of the feather under reflecting light
(Grubb 1989, 2006), or viewing and digitalizing an image produced
using a gel documentation system (Shawkey et al. 2003). This again
gives a near-maximum growth rate of stage (b) of feather growth. If
all growth bars from the tip to the base of the vanes can be counted,
and the growth time of the calamus is estimated from the ratio of the
septa bars (visible on the calamus) to the daily growth bars, almost
the entire duration of growth can be estimated (Winkler et al. 1988).
However, growth bars in naturally grown or moulted feathers, and in
worn feathers, are often very difficult or impossible to see (Michener
& Michener 1938, Wood 1950, Murphy 1992, Grubb 1995; own
obs.); growth bars are apparently much more easily seen in feathers
induced to regrow after plucking in winter in ptilochronology studies
(Grubb 1989, 2006). In summary, growth bars offer the possibility
of estimating feather growth rates of full-grown feathers, but this
method is often difficult to apply in practice.
Growth rates vary between different feathers of the same
generation (e.g. the different primaries) and also between different
feather generations (see Jenni & Winkler 2020: 3.2.6. and 3.2.7).
Therefore growth rates cannot be easily generalized across feather
types.
2.3.7 Physical properties of feathers and
fault bars
For many studies it is of interest to evaluate the physical qualities of
feathers produced by a given moult or to see how physical qualities
deteriorate with wear over time. Size (length, width) and mass of
feathers are obvious parameters, as well as rachis diameter and
density of barbs and barbules. Flexural stiffness or bending stiffness
(Worcester 1996, Dawson et al. 2000, Weber et al. 2005, de la Hera
et al. 2010a) is of particular interest for primaries. Tensile strength
has been tested for the rachis of flight-feathers and for the barbs of
body-feathers (DesRochers et al. 2009, Pannkuk et al. 2010). A whole
suite of mechanical properties has been measured for the barbs of
primaries (Butler & Johnson 2004).
91 mm
50
40
Primary moult score
30
20
10
0
June July August September
Fig. 21. Primary moult scores of Eurasian Treecreepers Certhia familiaris against
time of year (redrawn from Ginn & Melville 1983, with permission from BTO). Lines
connect primary moult scores of individuals examined twice and give an indication
of individual moult speed. Outliers may be a problem for some statistical analyses.
The number of fault bars (see Jenni & Winkler 2020: 4.1.5) can
be recorded for each feather and scored as: light (absence of some
barbules producing a visible discontinuity in the feather structure);
medium (a < 1 mm translucent line across the feather); or strong (a
> 1 mm translucent line across the feather) as well as whether the
fault bar has resulted in breakage of the barbules (damage) or not
(e.g. Jovani & Blas 2004, Sarasola & Jovani 2006). The prevalence
(proportion of feathers with fault bars) and abundance (mean number
per feather) of fault bars can then be calculated. Depending on the
study species, modified scoring systems might be useful, e.g. when
some of the fault bars do not cross the entire feather vane while
others do (Machmer et al. 1992, own obs.). If for a particular study
fault bars must be related to feather growth or particular events in the
life of the bird, the length of the feather examined (which together
with growth rate provides growth time) and the distance of each
fault bar from the tip of the feather (which allows determination
of the time of fault bar formation, provided feather growth time
is known) can be recorded (e.g. Negro et al. 1994, Bortolotti et al.
2002, Jovani & Blas 2004).
Recording the colour of the feather helps to document the effects
of wear, in particular that resulting from exposure to sunlight, and
the quality of new feathers and their effect on the appearance of the
bird with its many implications for inter- and intraspecific interactions.
Depending on the purpose of the study, feather colours can be
recorded by eye with reference to colour standards (e.g. for descri­
bing age differences visible to human observers) or, if their effects
on birds is the focus, by using a spectrophotometer as an objective
and quantitative comparison to known avian visual capacity. For
further details of methods used to record feather colours, the reader
is referred to the vast literature (e.g. Endler & Mielke 2005, Hill &
McGraw 2006, Quesada & Senar 2006, Vaquero-Alba et al. 2016,
Hubbard et al. 2017).
The analysis of stable isotopes in full-grown feathers is frequently
used to estimate where within its geographical range a bird was when
growing the feathers analysed. This technique can also be used to
shed light on moult processes. For example, stable hydrogen isotope
ratios in feathers can be used to determine whether birds moult
feathers before, during, or after migration (Bridge et al. 2011, Rohwer
  Terminology and Methods in Moult Research
et al. 2011) or to detect moult interruptions and determine
approximately where the feathers before and after suspension were
moulted (Neto et al. 2006, Pérez & Hobson 2006, Chabot et al.
2018). If adults moult their feathers far from the breeding area where
the juvenile remiges were grown, age classes can be discriminated
using stable hydrogen isotope ratios (Franks et al. 2009). Carbon and
nitrogen stable isotopes from several sections of primaries were used
to elucidate the complex moult pattern of Western Ospreys Pandion
haliaetus and to suggest that moult continues during migration
(Zelanko et al. 2011), as well as to show that some Greylag Geese
Anser anser migrate to specific areas to moult (Rohwer et al. 2015).
2.4 Analysing moult data
2.4.1 Seasonal timing and duration (progress)
of moult
A dataset of birds undergoing a complete moult is usually used to
determine the mean date of onset of moult and its duration, and
therefore its end. Most studies on the timing of a complete moult
are restricted to the primaries and this is justified by the fact that in
many species the primaries are moulted during (almost) the entire
period of a complete moult of an individual. Some species moult
body-feathers and secondaries beyond primary moult (e.g. Flegg
& Cox 1969, Traylor 1971, Seel 1976, Newton & Rothery 2005),
most notably the White-throated Dipper Cinclus cinclus (see p. 218).
Therefore, depending on the purpose of the study, secondaries may
60 mm
1 result that the estimates obtained with various regression methods
Mean score/mass
‘moult’ (Erni et al. 2013). This model uses a likelihood approach to
0 ‘moult’ (Erni et al. 2013): Type 1 in which birds are classified as moult
0 1
Proportion of time through moult
Type 3 in which each bird is given a moult score, but non-moulting
Fig. 22. Increases in primary moult score and moult index over the duration of primary
moult in 16 Common Starlings Sturnus vulgaris (means ± s.d.). The increase in primary
moult score (originally from 0 to 45, is standardized here to lie between 0 and 1)
is non-linear (open circles). When the primary moult scores are corrected using the
total mass of each primary to obtain the proportion of feather mass grown, an
almost linear trend in moult index is obtained (black dots). The solid line represents
the linear regression of feather mass from the time when P 2 started growing until
P 8 had finished growing, i.e. excluding points at the beginning and end of moult
Types 4 and 5 are useful if birds arrive just before moult or depart
when only P 1 or P 9 were growing. The increase in feather mass is almost linear
during most of the moult. Redrawn from Dawson (2003), with permission from Wiley.
21
also need to be considered. There have been relatively few attempts
to estimate quantitatively the progress of the renewal of all of the
feathers (including body-feathers) or the progress of partial moults
(e.g. Newton 1966, Seel 1976, Rymkevich et al. 1987, Mogilner &
Rymkevich et al. 1990, Rymkevich & Bojarinova 1996, Bojarinova et
al. 1999). In these studies the estimated mass of new feathers was
compared to the final feather mass of all tracts.
If data are available from individuals checked only once, the
starting point of most analyses is to sum the moult scores of the
individual primaries to obtain the primary moult score. A primary
moult score of 0 indicates that all primaries are still old, while a score
of 50 (or 45 for species with a much reduced P 10 which is not scored)
indicates that all primaries of one wing have been renewed and are
fully grown. For some statistical models, only birds before, during,
and after (primary) moult need to be distinguished.
Before statistical analysis of moult data, graphical inspection of the
data is always recommended (Fig. 21), e.g. to detect late-moulting
individuals (outliers) which some statistical models find difficult to
deal with (Erni et al. 2013). If some birds have been retrapped during
moult, connecting their data points on the graph helps to obtain a
first indication of moult speed (Fig. 21).
In early studies, primary moult scores (only of birds in moult)
against time were analysed using linear least squares regressions (e.g.
Pimm 1976, Ginn & Melville 1983). The problem with this method
is that it describes the average progress of moult in the sampled
population, rather than of the individual, because the date of onset
of moult varies between individuals. This problem could be rectified
if some of the individuals are captured at least twice during their
moult and explicit subject-specific modelling techniques are used
(see e.g. Zeger et al. 1988). Furthermore, least squares regressions
of data of birds captured only once are sensitive to the distribution
of the sampling dates, and violate the assumptions of a standard
linear regression (see Underhill & Zucchini 1988 for details) with the
may differ considerably (Summers et al. 1983). Therefore fitting
linear regression models to moult data of birds captured once is not
appropriate for estimating individual moult speed.
Today, the model developed by Underhill & Zucchini (1988) and
extended by Underhill et al. (1990) and Brandão (1997), the ‘UZ-model’,
is generally used, and is now conveniently available in the R package
specifically estimate the parameters of interest: the mean start date
of moult and its standard deviation as well as the average duration of
moult of an individual. It overcomes the difficulties that moult scores
are bound to lie between 0 and 50 (45) and that observations on start
and duration are not directly available (Erni et al. 2013).
Underhill & Zucchini (1988) and Underhill et al. (1990) provide
the likelihood equations for estimating the parameters of the model
using one of five data types; these are implemented in the R package
not started, during moult, moult completed; Type 2 in which each
bird is classified as in Type 1, but moulting birds receive a moult score;
birds are ignored; Type 4, similar to Type 2, but birds before moult are
absent, and only birds during and after completed moult are available;
and Type 5, similar to Type 2, but birds with completed moult are
unavailable, and only birds before and during moult are available.
just after moult. The UZ-model is not suited to more complicated
moults such as arrested, suspended, partial and stepwise moults.
 22 Terminology and Methods in Moult Research
The UZ-model is based on several assumptions (Underhill & Zucchini
1988, Erni et al. 2013): (a) the moult index increases linearly over time
for each individual (data Types 2 – 5); (b) the duration of moult, i.e.
the rate of change in moult index, is the same for every individual; (c)
the start date of moult of individuals is normally distributed around
the mean start date of the population (other distributions may be
implemented); (d) the individual observations are independent; (e)
the birds examined are a random sample from the population studied,
i.e. the probability of capture on a given day does not depend on
the stage of moult or whether a bird is in moult or not (although
it may vary between capture days due to trapping effort, weather,
etc.); and (f) there is no emigration or immigration, except for Types
4 and 5, which are specifically designed for situations where rapid
immigration just before moult or emigration just after moult occurs.
Furthermore, sampling effort should be spread out over the entire
moult period to obtain precise estimates (e.g. the start of moult is
difficult to estimate when only birds in the final stages of moult
have been caught). It is important to realize that the inclusion of
non-moulting birds before and after moult greatly improves the
precision of the estimates (Underhill & Zucchini 1988). Therefore,
recording non-moulting birds is important. One might get a feel for
the importance of imbalances in the distribution of data over the
moult period by applying various data type models, i.e. by omitting or
including non-moulting birds. For further discussion of the problem of
having many non-moulting birds in the sample see Bonnevie (2010b).
Some of the assumptions of the UZ-model may be violated to
certain extents in many datasets, resulting in biased parameter
20
Grey Plover (The Wash, UK)
P10
15
mass %
10 P5
5 In summary, the recommended method to obtain a linear moult
0 the mass of individual feathers (see 2.3.4) in three to six (or even only
Aug Sep Oct Nov
20
Ruddy Turnstone (Scotland) P10
15
mass %
10 P5
5
0
Aug Sep Oct Nov
Fig. 23. Timing of moult of each of the ten primaries (P 1 – P 10) of Grey Plovers
Pluvialis squatarola from The Wash, UK and of Ruddy Turnstones Arenaria interpres
from Scotland. The start date of moult is given on the x-axis for each primary (P 5
and P 10 are labelled). The y-axis represents the percentage of total primary mass.
The height to which the line of a given primary rises is the percentage of mass
of this primary of the total primary mass. This endpoint lies directly above the
completion date on the x-axis. Therefore the slope of the line gives the rate at which
feather material is deposited. Steep slopes represent rapid growth. The two examples
demonstrate two different strategies: fast growth rates and almost simultaneous
moult of the inner five primaries in Ruddy Turnstones; and slow growth rates and
spaced shedding intervals in Grey Plovers. Redrawn from Underhill (2003), with
permission from Springer Nature.
estimates and incorrect inference, but transformations may help to
reduce the effects of such violations.
In most cases it is advised to transform the moult score into a
linear moult index, which is scaled to range between 0 and 1.
In most cases, primary moult score does not increase linearly with
time, because the outer primaries are usually longer and have longer
shedding intervals, and therefore take more time to be renewed
than the inner primaries (e.g. Gwinner 1966, Zeidler 1966, Seel
1976, Summers et al. 1983, Newton & Rothery 2000, Rothery et al.
2001, Dawson 2003; Fig. 22). The deviation of the moult score from
linearity is more pronounced in birds whose outer primaries are much
longer than their inner primaries (waders, swifts, swallows). It has
been shown in several species that transforming the moult scores of
the individual primaries into percentage of feather mass grown
(using the mass of the corresponding feathers, see 2.3.4) results in
a linear moult index (Fig. 22; Summers 1980, Summers et al. 1980,
1983, Underhill & Summers 1993, Underhill & Joubert 1995; but see
Underhill et al. 1990 for a case needing a power transformation).
More recent detailed investigations have shown that the increase
in total mass of new primary feather material is indeed linear over
time (except at the very beginning and end of primary moult) in
Common Starling Sturnus vulgaris, Carrion Crow Corvus corone,
Eurasian Magpie Pica pica, European Greenfinch Chloris chloris and
Eurasian Bullfinch Pyrrhula pyrrhula – species with widely differing
wing shapes (Dawson 2003, Dawson & Newton 2004; see also Seel
1976 for Western Jackdaw Coloeus monedula). The analyses by
Dawson (2003) and Dawson & Newton (2004) not only corrected
moult scores by the mass of the different primaries, but also by the
unequal distribution of mass along a primary (because the tip of the
feather is lighter than the base), a correction already suggested by
Seel (1976) and Redfern (1998). However, the inaccuracy introduced
by assuming a uniform distribution of mass along the length of a
feather is negligible (Dawson & Newton 2004). Rothery et al. (2001)
modified the UZ-model for non-linear increases in moult index, but
this requires determination of the shape of the non-linear curve from
samples of captive birds or birds caught repeatedly during moult.
index (e.g. for use in data Types 2 – 5 of the UZ-model) is: (a) determine
one) individuals (Underhill & Joubert 1995, Dawson & Newton 2004);
(b) calculate the relative contribution of the mass of each primary
feather to total primary feather mass; (c) transform the standard moult
score of each feather into percentage of feather mass grown (e.g. by
using the proportions 0, 0.125, 0.375, 0.625, 0.875 and 1 for scores
0 – 5, respectively; Underhill 2003); and (d) finally sum these up to
obtain the proportion of new primary feather mass grown, i.e. the
moult index ranging between 0 and 1 (Summers et al. 1980, Dawson
& Newton 2004).
Violations of other assumptions of the UZ-model cannot easily be
removed from a given dataset. Importantly, the probability of capture
should not depend on moult stage (assumption (e) above). However,
many species behave differently when moulting and are less likely
to be caught compared to when they are not in moult (e.g. Newton
1966). Multistate capture-recapture models with ‘moult stage‘ as a
state could show whether capture probability depends on moult stage.
Violation of this assumption could bias estimates obtained from the UZ-
model (Underhill et al. 1992, Bensch & Grahn 1993) and may be partly
removed by assuming data Type 3, 4, or 5.
Certain assumptions of the UZ-model, such as equal moult duration
among individuals and normal distribution of the moult start date

 Terminology and Methods in Moult Research
can sometimes be overcome by using explanatory variables (e.g. Burman
2016). A major advance by Erni et al. (2013) was not only to implement
the UZ-model in R, but also to allow for an examination of the effects
of explanatory variables (such as different populations, sexes, migration
distance, photoperiod, year, etc.) on start date of moult and its standard
deviation as well as the duration of moult.
As an alternative to the UZ-model for Type 1 data, the start date
can also be estimated using a probit analysis (Rothery & Newton 2002),
which is equivalent to the UZ-model, while the end date is estimated
in a second probit analysis (see Erni et al. 2013 for implications of this)
without any assumption about the form of change in moult index over
time. This allows for a different standard deviation for the start and end
date. The duration is the difference between average end and start
dates. This method uses only two states of moult; not started moult
(0) and started or completed moult (1).
If the moult scores of individual primaries (transformed into
percentage of feather mass grown) are analysed feather by feather using
the UZ-model, a detailed picture of the timing and growth duration of
the individual primaries, including shedding intervals and number of
simultaneously growing primaries, can be obtained from birds caught
once (Fig. 23; but large sample sizes of moulting birds are needed;
Serra 2000, Underhill 2003, Underhill et al. 2006, Remisiewicz et al.
2009, 2010, Barshep et al. 2013, Burman 2016). The daily increment in
proportion of feather mass grown over all primaries can be estimated
from the estimated daily growth rates of the individual primaries and
their timing. This overall flexible moult curve, which should be similar
to the moult index curve over time, can then be inspected for linearity
(Remisiewicz et al. 2009, 2010, Burman 2016).
An entirely different method from the UZ-model to calculate moult
duration and its standard error was proposed by Rohwer & Broms
(2012) and applied by Rohwer (2013). The duration of a complete
primary moult is estimated as the length of P 1 when P 2 is shed, plus
the length of P 2 when P 3 is shed, and so on through P 9. The full
length of P 10 is then added and this sum is divided by the daily growth
rate of the primaries. For this method, the mean length of each primary,
the moult score of each primary and an estimate of growth rate from
recaptures or from growth bars are needed. An advantage of this
method over the UZ-model is that moult duration can be estimated
for partial moults and for moults that start at different primaries but
with a consistent direction of sequence, and for species which moult
asynchronously, such as many tropical species.
Surprisingly, in all of the statistical analyses mentioned to this point,
no integrated use is made of birds examined more than once during
moult, although they provide quite important and precise information
about individual moult speed (Fig. 21). It is therefore desirable that
further developments of the statistical models used to analyse timing
and speed of moult allow inclusion of more than one moult score per
individual from more than one day, e.g. by embedding the UZ-model
into mixed models.
Because the rate of new feather material accumulation in primaries
is linear over much of its range (Fig. 22), two moult scores taken a
few days apart during moult (not at the very beginning and end)
from the same individual give a reliable indication of moult speed
(and therefore duration) for that individual, provided that the moult
scores are converted to percentage of feather mass grown. Furthermore,
Johnson (1989) proposes a linear regression model to estimate the
time intervals between full-growth of each successive primary from
retraps (and by adding them up total moult duration), as well as the
start date of moult. However, recaptures possibly constitute a biased
sample, because capture probability may be influenced by moult speed;
91 mm
23
5
4
Rate of increase in dry mass (g/15 days)
3
2
1
0
May Jun Jul Aug Sep Oct Nov Dec Jan
Fig. 24. Estimates of the intensity of the renewal of the entire plumage, including
body-feathers, expressed as the rate of new feather material produced per 15 days,
for five species of corvids: dots Carrion Crow Corvus corone; open circles Rook C.
frugilegus; open squares Western Jackdaw Coloeus monedula; crosses Eurasian
Magpie Pica pica; and triangles Eurasian Jay Garrulus glandarius. Redrawn from Seel
(1976), with permission from Wiley.
individuals with many growing wing-feathers are reluctant to fly and
might be under-represented in mist-net captures.
If individuals can be examined several times during moult or even
systematically from beginning to end, very detailed information
about the moult of each feather can be obtained (e.g. Gwinner 1966,
Zeidler 1966, Seel 1976, Winkler & Winkler 1985, Winkler et al. 1988,
Hegelbach 2014). The findings can be presented either as bars (e.g.
Fig. 26) or as growth trajectories (similar to Fig. 23). Such studies are
generally done on captive birds and it therefore remains to be shown
whether moult in captive birds differs from free-living conspecifics.
2.4.2 Moult intensity and moult speed
For wing-feathers, moult intensity (expressed as the number of feathers
growing simultaneously) can easily be derived from the standard moult
scores of each feather or from analyses such as those presented in Fig.
23 and 26. The changes in the number of simultaneously growing
wing-feathers during the course of moult can be presented graphically
against time, or against the progress of moult (Serra & Underhill
2006, Remisiewicz et al. 2009; see Rohwer & Rohwer 2013 for the
calculation of average moult intensity). If moult scores of wing-feathers
are translated into new feather material added (see 2.4.1), estimates
of the rate of wing-feather material deposited can be obtained (e.g.
Dawson 2003).
Moult intensity of body-feathers is usually directly recorded as the
proportion of growing feathers in the different tracts (see 2.3.3) and
there are various ways of combining tracts to produce a single score
(e.g. Newton 1966, Seel 1976, Rymkevich et al. 1987, Mogilner &
Rymkevich et al. 1990, Rymkevich & Bojarinova 1996, Bojarinova et al.
1999). The conversion into the rate of body-feather material deposited
has only rarely been undertaken. Given its importance in the study of
the costs and physiology of moult, there are surprisingly few attempts
to estimate the intensity of the renewal of all of the feathers, including
body-feathers which could then be expressed as the rate of new feather
material produced (e.g. Newton 1966, Seel 1976, Dietz et al. 1992, Fig.
24; however, feather sheaths should also be taken into account; see
Murphy & King 1986).
 24 Terminology and Methods in Moult Research
00
00
0
0
2
3
3 2.4.3
2
3
2
4 presented graphically (e.g. Fig. 23 and 26). In describing the sequence
1
5 between tracts, the primaries are usually taken as a reference against
5 0
0
3
1 2
0 0 0 0 4 complicated moult sequences, such as serial moults (see also Rohwer
0
0 0 0 & Rohwer 2018). The tables yield quantitative results, from which
Fig. 25. The raggedness score is a measure of the wing area missing (pink) because of
shed or growing flight-feathers. The raggedness score is the sum of the raggedness
values (in red) which are the differences between the moult score (in black) for missing
and growing feathers (1 – 4) and 5; old and new feathers have a raggedness value of
0. Primaries and secondaries 1 – 6 are taken into account. Therefore in a passerine
with 9 scored primaries, raggedness varies between 0 (no growing or missing P and
S) and 60 (all nine P and six S shed). In this example P 3 – 6 and S 1 – 2 are growing
and have reached different lengths compared to full-grown, giving a raggedness
score of 14. After Bensch & Grahn (1993).
The speed of primary moult can be derived from analyses which
produce estimates of moult duration (see 2.4.1), but another method
exists which can be used in some birds. The speed of primary moult
is determined mainly by their shedding intervals, i.e. the number of
simultaneously growing feathers, rather than by feather growth rates
(Rohwer & Rohwer 2013). This prompted the use of the difference in
length between adjacent actively growing primaries (Green 1974, but
not developed further), or the gap in the wing caused by incompletely
grown wing-feathers, as an estimate of the speed of moult. The gap in
the wing is measured as a raggedness score (Haukioja 1971a; Fig. 25).
Because the raggedness score is positively correlated with primary moult
score, the residuals of this relationship were taken and proved to be
correlated with the speed of moult (Bensch & Grahn 1993). Estimating
individual speed of moult from wing raggedness is possible when data
from birds in full active moult are available (>30 birds, without birds
in early and late moult stages) and for species which grow more than
one or two wing-feathers simultaneously (Bensch & Grahn 1993). The
residual raggedness value is a relative measure of moult speed that
can be translated into actual moult speed by reference to retraps with
known moult speed. An advantage of this method is that the speed of
moult can be estimated for individuals, while the UZ-model provides
sample means.
The gap in the wing, as measured by the raggedness score, can be
transformed into a measure of relative wing surface area, or the wing
area missing because of moult (Hedenström 1998), important variables
when examining the effects of wing-feather moult on flight capacity.
Similarly, by taking into account the relative masses of the primaries the
raggedness score can be transformed into the proportion of feather
mass missing (Remisiewicz et al. 2009).
Sequence of moult 
The sequence of moult is usually described in words but can also be
which the progress of other tracts is plotted.
Rohwer (2008) developed a system of tables to detect more
groups of feathers that form separate moult series, feathers that form
nodes, moult directions within these series, and multiple moult waves
can be derived. The tables use data obtained from birds caught once.
If individuals can be examined repeatedly over entire moult cycles
or several moults, complicated moults can be followed directly (e.g.
White Stork Ciconia ciconia in Bloesch et al. 1977 and Sutter 1984;
Eurasian Eagle-Owl Bubo bubo in Glutz von Blotzheim & Bauer 1980;
see Jenni & Winkler 2020: 5.3.4).
2.4.4 Extent of moult
While a description of the sequence of moult requires the examination
of birds with growing feathers, assessing the extent of moult requires
observations on birds after the completion of moult and is usually
performed for partial moults. Obviously, the difficulties lie in assigning
the feathers to the different feather generations. The extent of moult
is usually described in quantitative terms and presented graphically for
each feather or tract, while the interdependence in extent between
tracts is often given relative to the greater coverts (see the graphs
in the species accounts, explained on p. 100). Again, the extent of
moult can be translated into feather material produced during a
given moult using the mass of the feathers (e.g. Herremans 1988b,
Gargallo 1996, 2013).
The metric ‘moult topography’ quantitatively addresses the spatial
configuration of moult using two parameters: extent and vector
(Guallar et al. 2014). A schematic wing is used with the single feathers
(or feather blocks for the marginal coverts) shown on an orthogonal
plane with their Cartesian coordinates in an anterior-posterior
and proximal-distal dimension. The extent quantifies the number
of feathers replaced, while the vector gives the mean direction of
moult extent from an arbitrary origin. This illustrates the spatial
configuration of feathers replaced at the end of a partial moult (it
is not intended to reveal moult sequence) and is most suitable for
species with relatively simple moult patterns.
  The Moult of Adults
Chapter 3
The Moult of Adults
Adult European passerines all renew their plumage at least once
a year, raising three main questions: (a) when should moult be
fitted into the annual cycle of other events (breeding, migration,
survival of the cold period; see 1.5), including in migratory birds
the question of whether to moult in the breeding or non-breeding
area?; (b) should all of the feathers be moulted once annually,
or should some or all of the feathers be moulted twice or even
three times, so increasing the number of moult periods to be fitted
in the annual cycle?; and (c) in which sequence should feathers
be moulted during a moult period so that the resulting plumage
impairment is minimised and the feathers are in optimal condition
for the imminent requirements.
In this chapter, we start with a summary of the moult strategies of
adults (see 3.1) to enable understanding of the subsequent sections.
Then we describe the sequence of moult (see 3.2), which is rather
uniform. Then we describe the various moult strategies adopted
by adult European passerines (when and where to moult, and how
many times) (see 3.3) which are uniform among species wintering in
temperate or Mediterranean areas, but more variable and complex
in long-distance migrants. Finally, we take a closer look at the timing
and duration of moult, in particular how moult is fitted into the
annual cycle when time is limited, and in sub-Saharan Africa (see 3.4),
and try to discern the factors shaping the various moult strategies
(see 3.5).
3.1 Introduction to the moult (e.g. Svensson 1992), and to various ways of describing how moult
strategies
European passerines live in a highly seasonal environment, so that
their annual cycle is well structured and fairly well synchronized
between individuals in a population. They usually have a restricted
and well-defined breeding season and invest heavily in reproduction
(large clutch size compared to tropical species; worn plumage
after breeding). Therefore, moult does not generally overlap with
breeding, except in larger species and, in some birds, at the end of
the breeding season. While all adult passerines normally replace the
entire plumage at least once a year, many renew part of, and some,
the entire plumage twice annually (for cases of the renewal of some
feathers three times a year see 3.3.1 and 3.3.2; for the occasional
retention of some feathers for more than a year see 3.3.3).
In resident species and short-distance migrants (wintering in
Europe or North Africa), the cold winter precludes moult at that time.
In these species, moult is restricted to one distinct period, between
the breeding season and autumn migration or the onset of winter
(see 1.4). In some of these species, some feathers may be moulted
during a secondary moult period in late winter/spring.
25
Long-distance migrant European passerines (most wintering in
sub-Saharan Africa, and a few in subtropical and tropical Asia) need a
considerable period for autumn and spring migration, and possibly for
itinerant movements within the wintering area. Therefore, the moult
period is time-limited if carried out in the breeding area, i.e. before
autumn migration (ways in which moult can be inserted between
breeding and the autumn migration are treated in 3.4). However,
long-distance migrants can transfer part of, or the entire, main moult
into the non-breeding area if conditions there are better than in
the breeding area after breeding. Under such benign ‘wintering’
conditions, migrants may considerably lengthen the potential period
for moulting, and may even subdivide a moult or add another moult.
Four moult strategies have long been generally recognized in
passerines, based on primary moult only: (a) complete moult before
autumn migration or the onset of winter; (b) complete moult after
autumn migration; (c) two complete moults annually; and (d) moult
suspension during autumn migration (e.g. Stresemann & Stresemann
1966, Ginn & Melville 1983). However, a more realistic insight into
the moult strategies results from considering the moult of the
remainder of the plumage as well. In particular, it enables a better
understanding of the sometimes bewildering variation between
members of the same species. Also, unusual moult patterns can be
more easily understood as transitions between the more common
moult strategies. The usefulness of considering the moult of all parts
of the plumage when ageing birds has been recognized for some time
and has led to a rough classification of moult according to season
(summer or winter) and extent (partial, complete, suspended, etc.)
is fitted into the annual cycle (e.g. Newton 2011, Kiat et al. 2019a,
Jenni & Winkler 2020: 5.3).
Because of the variability within species with many intermediate
stages, moult strategies are not always clearly distinguishable. Here
we distinguish five main moult strategies with several sub-categories
(Table 2; Fig. 43 – 47 on p. 49 – 53).
All adult sedentary and migrant European passerines which winter
in cold and temperate climates, as well as about half of the long-
distance migrants, perform a complete moult each year between the
breeding season and the autumn migration or winter. The majority do
not moult again during winter/spring and only perform a complete
post-breeding moult (moult strategy 1, Fig. 43). About one third
of the species which perform a complete post-breeding moult in
the breeding area renew part of or all of the body-feathers (and
occasionally also the tertials and rectrices, rarely some secondaries)
again during winter/spring, and thus perform a complete post-
breeding moult and a partial pre-breeding moult (moult strategy
2, Fig. 44).
In northern latitudes, late winter/spring seems to be a rather
unfavourable season for an extensive moult, probably because of the
demands of territorial defence and courtship, generally scarce food
resources, and the spring migration. Hence, Palaearctic passerines
 26 The Moult of Adults

Table 2. The principal moult strategies of adult European passerines based on trochilus regularly performs two complete moults a year (for other
the seasonal occurrence and extent of their moults. Moult strategies of species
species elsewhere see Jenni & Winkler 2020: 5.3.2).
with variable breeding seasons (e.g. Loxia curvirostra) are not included. For details,
subgroups and special cases see 3.3. Species with a very variable breeding season avoid, or at least slow,
moult during breeding and, consequently, cannot follow a fixed moult
Adult Summer/autumn Winter/spring cycle (see 1.4). They interrupt moult at various stages and this results
moult strategy (before autumn migration) (after autumn migration)
in individually variable and sometimes complex moult patterns which
1 complete
(post-breeding) are not covered by the moult strategies given above. The only such
species treated in this book is Loxia curvirostra. We entirely omit this
2 complete partial
(post-breeding) (pre-breeding) species from the following general account and refer the reader to
3 none or partial complete (pre-breeding or
the detailed species account (p. 265). We also omit the peculiarities
(post-breeding) post-breeding) of the moult of Cisticola juncidis (see 4.5.4).
4 first part of remiges second part of remiges
(seasonally divided) (seasonally divided)

3.2 Sequence of moult

5 complete or almost complete
complete (post-breeding) (pre-breeding)

3.2.1 Basic sequence of the complete moult

Compared with non-passerines, the sequence of the complete moult

usually do not perform a complete moult during spring, except while
still in their tropical wintering areas. Exceptions include some of
the corvids which start moult in May while breeding (Corvus spp.)
or in March (Corvus corax, Nucifraga caryocatactes; e.g. Glutz von
Blotzheim & Bauer 1993, Cramp & Perrins 1994a), and Yellow-
breasted Greenfinch Chloris spinoides which has an unusually late
breeding season in August in the Himalayas and moults completely
in May and June after spring migration (Whistler 1940).
The remaining three strategies include the renewal of extensive
parts of the plumage during the northern winter and occur only in
birds wintering in the tropics. Some long-distance migrants enjoy
stable, favourable conditions for moult in the tropics and take the
opportunity to perform their main moult after the autumn migration
(complete moult in the non-breeding area, moult strategy 3, Fig.
45). This strategy is adopted by about one half of the long-distance
migrant European passerine species. Many perform a partial moult
before autumn migration, which may be interpreted either as the
beginning of the complete post-breeding moult performed mainly in
the non-breeding area or as a separate partial post-breeding moult
(see 3.3.2).
of passerines is rather uniform between species and is referred to
as the basic sequence (Ginn & Melville 1983). The complete post-
juvenile moult follows the same sequence and is, therefore, included
in this section. A full description of the moult sequence includes
both the sequence within each feather tract and the relationships
between feather tracts.
Flight-feathers
Within the flight-feather tracts, the basic sequence is as follows (Fig.
26, 27 and 28). The primaries are moulted descendently beginning
with P 1, and the secondaries ascendently beginning with S 1. The
tertials follow their own sequence and are usually renewed in the
order T 8-9-7. The rectrices are renewed in pairs, centrifugally,
starting with R 1.
Primary moult extends over virtually the entire moult period and
is usually taken as a reference for the progress of moult in the other
9
Primaries

7
Certain long-distance migrants moult part of the remiges before
5
autumn migration and part during winter in the non-breeding area
(seasonally divided moult of remiges, moult strategy 4, Fig. 46). 3

This strategy is defined for the sake of simplicity and actually includes 1

several rather different moult strategies which are detailed in section 1

3.3.3. The majority of the plumage may be renewed before or after
Secondaries

3
autumn migration, and the body-feathers, tertials, and rectrices once
5
or twice annually. This seasonal division of the moult of remiges is the
7
normal strategy in only a few species. In many species, only a minority
9
of the individuals follow strategy 4, the majority moulting according to
strategy 2 or 3. For moult research, the patterns subsumed in strategy 1
Rectrices

4 are of particular interest because they show that moult strategies 3

are variable, not clear cut or mutually exclusive, and because they 5
suggest ways in which moult strategies may have evolved.
While birds moulting according to strategies 1 – 4 may renew 10 30 50 70 90 110 130
only the body-feathers, tertials, and rectrices twice annually, birds Days

belonging to moult strategy 5 (Fig. 47) also renew all of the remiges,
or almost all, twice each year (biannual complete moult; post-
breeding moult arrested within remiges and complete pre-
breeding moult). Only one European passerine species, Phylloscopus
Fig. 26. Sequence of flight-feather moult in a captive Pyrrhocorax graculus. The bars
indicate the growth duration of individual flight-feathers, divided into the growth
of the vane (grey bar) and the final growth and keratinization of the calamus which
results in only a slight increase in feather-length (white bar). Redrawn from Winkler
et al. (1988), with permission from Ala.
  The Moult of Adults 27

A B C

D E F

Fig. 27. Progress of the complete moult of Acanthis cabaret following the basic sequence in six snapshots, according to data collected in the Swiss Alps by the authors. Half of the birds
pass through the six stages within the dates shown. Each feather tract has its own colour, which is light in shade when old and darker in shade when renewed. A (10 – 16 August): The
complete moult starts with the successive shedding of the innermost two primaries (P 1 and 2). B (17 – 26 August): P 1 and 2 are full grown while P 3 and 4 are growing. The greater
coverts grow under cover of the old median coverts. The central tertial (T 8) is being replaced, while the old T 7 and 9 protect the closed wing. The renewal of the rectrices starts. The
body-feathers are still mostly old. C (27 August – 7 September): The gap in the primaries has progressed distally and increased in size, now comprising three growing primaries (P 4 – 6).
Body-feathers and marginal coverts start to be replaced. The median coverts are renewed while being protected by the undermost row of old marginal coverts. The renewed T 8 protects
the closed wing while T 7 and 9 grow. All rectrices are growing. Flight is certainly impaired. D (8 – 15 September): The moult of the secondaries starts with the outermost, which creates
a second wing-gap. However, the two wing-gaps are separated from each other by a block of about five renewed primaries. Flight impairment has probably reached its maximum,
but the rectrices are soon fully restored. The undermost row of marginal coverts grows under protection of the renewed upper ones. The intensive moult of body-feathers now also
includes the head. E (16 – 25 September): The gap in the secondaries has increased, while the outermost primaries are growing. The rectrices are full-grown, as well as most of the
body-feathers. F (26 September – 2 October): The primaries and wing-coverts are full-grown. The last growing feathers are secondaries 5 and 6, and some feathers on the head which,
on the average bird, will be full grown by 10 October. In this stage of moult, flight capability is hardly impaired and migrants may start autumn migration. Drawings by Niklaus Heeb.

feather tracts. In an average passerine, the renewal of the tertials and
rectrices starts once primary moult has reached about P 4 (Fig. 26,
27 and 28). Secondary moult begins during the growth of P 5. The
tertials and rectrices are full-grown by the time that P 8 and 9 are
growing. Due to its small size, P 10 is usually full-grown before P 9 or
P 8. The last flight-feathers to complete growth are usually S 5 and S
6. Normally, moult proceeds symmetrically on both wings.
Body-feathers and wing-coverts
Moult of the body-feathers usually starts after the beginning of primary
moult and often lasts longer than the renewal of the flight-feathers.
The feathers on the head are normally moulted last. The greater and
carpal coverts are shed soon after the start of primary moult and are
well or almost full-grown before the secondaries are renewed (Fig.
28). The greater coverts are all shed at once, or in a variety of more
complex patterns: e.g. within short intervals; descendently in groups
(e.g. Bährmann 1964, Zeidler 1966, Mester & Prünte 1982, Alonso
1984); ascendently (Rogge 1966, Dhondt 1973); convergently (Winkler
& Winkler 1985); in an ascendent and divergent wave (Dorka 1971);
or divergently with a very variable focus around GC 4 – 5 (Kasparek
1979b). While the carpal covert is moulted in sequence with the
greater coverts, GC 10 is not renewed until the other greater coverts
are almost full-grown (Zeidler 1966, Kasparek 1979b, Alonso 1984,
Winkler & Winkler 1985).
The median coverts are renewed while the new greater coverts are
growing, or are full-grown. The marginal coverts start moulting soon
after P 1 is shed and are renewed throughout much of the period
of primary moult. The alula and the underwing-coverts are replaced
during the second half of primary moult. The primary coverts are
renewed together with the corresponding primary, slightly earlier or
later (Rogge 1966, Zeidler 1966, Kasparek 1979b, Winkler & Winkler
1985, Dittberner & Dittberner 1987). There is only limited variation
in the moult sequence of the wing-coverts between species, but
detailed descriptions are only available for a small number of European
passerines (e.g. Corvidae, Dorka 1971, Seel 1976; Pyrrhocorax
graculus and P. pyrrhocorax, Winkler et al. 1988; Parus major, Dhondt
1973; Sturnus vulgaris, Bährmann 1964; Erithacus rubecula, Rogge
1966; Passer domesticus, Zeidler 1966; P. hispaniolensis, Alonso 1984;
Montifringilla nivalis, Winkler & Winkler 1985; Prunella modularis,
Ginn 1975; Pyrrhula pyrrhula, Newton 1966, 1967; Acanthis cabaret,
Winkler & Jenni 2007; Emberiza schoeniclus, Kasparek 1979b).
3.2.2 Functional aspects of the basic sequence
of moult
Many aspects of the basic sequence of moult can be interpreted as
functional adaptations to mitigate the impairment of the plumage
during the time feathers are missing or growing (Fig. 27). However,
since no one has yet performed a detailed study on the effects
of wing-feather moult and its sequence on flight performance in
 28 The Moult of Adults

MaC AI A MaC AI F less affected by aerodynamic forces. Secondary moult starts only
MeC CC MeC CC

GC PC PC
after the innermost primaries are full-grown, thereby avoiding a
GC

P P
widening of the gap during replacement of the innermost primaries
T/S T/S
(Fig. 27). Another explanation posits that the outer primaries have a
10 9 8 7 6 5 4 3 2 1 1 2 3 4 5 6 7 8 9 10 10 9 8 7 6 5 4 3 2 1 1 2 3 4 5 6 7 8 9 10

R R
stronger effect on flight ability and suffer higher wear and therefore
should be moulted closer to the winter than the inner primaries,
while secondaries are more uniform and can therefore have a more
variable moult sequence (Barta et al. 2006).
MaC AI B MaC AI G
MeC CC MeC CC The tertials have a protective function while the secondaries serve
GC PC GC PC mainly as aerofoils during flight, and each follow their own moult
T/S
P
T/S
P sequence. The protective tertials and greater coverts are already
10 9 8 7 6 5 4 3 2 1 1 2 3 4 5 6 7 8 9 10 10 9 8 7 6 5 4 3 2 1 1 2 3 4 5 6 7 8 9 10 moulted, or at least well grown, before secondary renewal commences.
R R
Among the tertials, T 7 and 8 have the greatest protective function
and their moult is separated in slow moulting passerines (e.g. Corvus
frugilegus, Dorka 1971; Pyrrhocorax graculus and P. pyrrhocorax,
MaC AI C MaC AI H Winkler et al. 1988; Passer domesticus, Zeidler 1966; Montifringilla
MeC CC MeC CC

PC PC
nivalis, Winkler & Winkler 1985). The old median coverts protect
GC GC

P P
the first stages of greater covert growth. They themselves are
T/S T/S
moulted under the protection of the old undermost row of marginal
10 9 8 7 6 5 4 3 2 1 1 2 3 4 5 6 7 8 9 10 10 9 8 7 6 5 4 3 2 1 1 2 3 4 5 6 7 8 9 10

R R
coverts (Fig. 27). The primary coverts are shed together with the
corresponding primary, but since the base of each primary is covered
by the preceding proximal primary covert, some protection of the
primary pin at least is ensured.
MaC AI D MaC AI I
MeC CC MeC CC The alula feathers are replaced during the renewal of the outer
GC PC GC PC primaries, when their function of stabilizing airflow over the outer
T/S
P
T/S
P primaries at high angles of attack is obsolete (Evans 1966). The
10 9 8 7 6 5 4 3 2 1 1 2 3 4 5 6 7 8 9 10 10 9 8 7 6 5 4 3 2 1 1 2 3 4 5 6 7 8 9 10 rectrices are renewed before the outermost primaries, so that at least
R R
one of these sets of feathers, which are both important in landing,
have no gaps during moult (Fig. 27, Evans 1966).
Many studies report that the speed of primary moult, as measured
MaC AI E MaC AI J by the increase in moult score, slows after the shedding of the
MeC CC MeC CC
central primaries, until all the primaries are full-grown, so that the
GC PC GC PC

P P
inner primaries are replaced faster than the outer ones. This is mainly
T/S
T/S
due to longer shedding intervals, but also to the slower growth in
10 9 8 7 6 5 4 3 2 1 1 2 3 4 5 6 7 8 9 10 10 9 8 7 6 5 4 3 2 1 1 2 3 4 5 6 7 8 9 10

R R
Fig. 28. Progress of the complete moult of the wing and tail, following the basic
sequence. The progress of moult is represented schematically by ten stages (A–J), one
for each primary to moult. The primaries (P), primary coverts (PC), alula feathers (Al),
secondaries (S), tertials (T), greater coverts (GC), carpal covert (CC), median coverts
(MeC) and marginal coverts (MaC) of the right wing as well as the right half of the
rectrices (R) are represented by rectangles. The body of the bird lies at the left edge
of each diagram. Old feathers are shown white, new feathers are grey, growing
feathers are black, and all are shown at their approximate length. The example
species is Passer domesticus, modified after Zeidler (1966).
passerines (cf. Jenni & Winkler 2020: 3.4.4) much that follows
remains speculative.
The primaries take the most time to be moulted, since shedding
many, or all, at once or in short succession, would severely reduce
flight capability, and because the daily growth-rate of an individual
primary is limited (see Jenni & Winkler 2020: 3.2). The body-feathers
of small passerines comprise 72 – 82 % of the total plumage-mass
(see Jenni & Winkler 2020: 3.2.2) and consequently, require most
energy and nutrients for replacement. Therefore, it would seem
adaptive to spread both primary and body-feather moult over the
entire moult period.
The descendent replacement of the primaries is thought to provide
better protection for the growing primaries against upward bending
by lift forces during flight, than an ascendent sequence (Noordhuis
1989). The secondaries are shorter, overlap more and are apparently
length of the outer compared to the inner primaries. Therefore, the
outer primaries not only take longer to grow because they are longer
than the inner primaries, but also because they grow more slowly.
Specifically, in species in which primaries increase in length, and/
or are relatively heavier, from inside to outside, shedding intervals
become longer towards the outside, while in species with round
wings, the shedding intervals are longest for the longest primaries
(e.g. Gwinner 1966, Zeidler 1966, Newton 1967, Dorka 1971, Seel
1976, Winkler & Winkler 1985, Winkler et al. 1988, Dawson 2003).
Moreover, the primaries distal to the wing-tip are proportionally
heavier than the primaries proximal to the wing-tip (Dawson 2005)
and hence have a lower growth-rate by length than proximal
primaries of comparable length (e.g. Newton 1967, 1969, Seel
1976, Winkler & Winkler 1985, Winkler et al. 1988, Dawson 2003,
Pap et al. 2008). The longer shedding intervals and lower growth-
rates by length of outer primaries result in the generally constant
production of feather material during the replacement of primaries
in a sequential complete moult, except at the very beginning and
end when only one primary is growing (Seel 1976, Dawson 2003,
2004, Dawson & Newton 2004).
Three explanations may account for the decelerated moult of the
outer primaries. First, the moult of primaries may be constrained by
physiological circumstances limiting synthesis and/or deposition of
primary feather material, as suggested by the constant rate of primary
feather material production (Dawson 2003; see Jenni & Winkler
2020: 3.2.8). Second, the deceleration of primary moult is especially
  The Moult of Adults
marked when tertials, rectrices, and secondaries start moulting and,
consequently, may be explained by overall metabolic constraints
(e.g. Zeidler 1966, Spina & Massi 1992). However, the phenomenon
also occurs in species with a long moult duration and, hence, low
metabolic requirements. Moreover, Muscicapa striata with its inverse
moult sequence (see p. 190) also shows prolonged shedding intervals
of the outer primaries, although few other feathers are growing at
that time (Dorka 1971). Third, and most likely, moult of the outer
primaries is decelerated because gaps among the outer primaries
impair flight capability more than those among the inner primaries,
and thus should be minimized, especially in species for which
continuous flight capability is important (Gwinner 1966, Sach 1968,
Dorka 1971, Dawson 2003; see Jenni & Winkler 2020: 3.4.4).
In contrast, secondary moult usually starts with long shedding
intervals. If the primaries are almost full-grown, secondary moult may
speed up considerably, i.e. while shedding S 4 – 6 (e.g. Dwight 1900,
Evans 1966, Newton 1966, Rogge 1966, Evans et al. 1967, Dorka
1971, Seel 1976, Kasparek 1979b). This may be interpreted as a way
to minimise overall wing raggedness while the outer primaries are
being replaced. In Corvus frugilegus, secondaries are shed during the
intervals between two primary sheddings, rather than at the same
time as a primary, which also minimizes wing raggedness to some
extent. In addition, the intervals between shedding of the remiges are
very constant between individuals, except for the intervals between
P 5/6, P 9/10, S 1/2 and S 5/6. These exceptional, variable shedding
intervals apparently serve to re-establish the coordination between
primary and secondary moult and to accelerate the moult in late
moulting birds (Dorka 1971). There are apparently no other detailed
studies of the variation between individuals in the shedding intervals
between remiges and its functional importance in coordinating the
moult of flight-feathers.
3.2.3 Variations and exceptions to the basic Secondaries
moult sequence
Primaries
Among European passerines, the moult sequence of the primaries
is generally strictly descendent. However, in a few individuals and
species the focus of a complete primary moult may be shifted from
P 1 to a central primary (divergent sequence) or to P 9 (ascendent
sequence). Species normally showing the basic descendent sequence
may exceptionally start with P 2 or P 3 (e.g. Corvus frugilegus,
Garrulus glandarius, Chloris chloris, Linaria cannabina, Serinus
syriacus; Bährmann 1971, Dorka 1971, Kiat 2017). A small proportion
of Lanius cristatus show a divergent complete post-breeding or pre-
breeding primary moult starting with P 3, 4 or 5, while the majority
exhibit the normal descendent sequence (Stresemann & Stresemann
1971). A considerable proportion of Locustella luscinioides moult
the primaries divergently with a variable focus around P 3, 4 or 5,
but complete descendent and eccentric descendent primary moult
also occurs (Steiner 1970, Stresemann & Stresemann 1970b, Mead
& Watmough 1976, Thomas 1977, Müller 1981, Trias et al. 1982,
Bensch et al. 1991, Márton et al. 2000, Neto et al. 2006, Neto &
Gosler 2006, Kulaszewicz & Jakubas 2015, own obs.). Some Anthus
richardi have been observed moulting from P 1, as well as from a
central primary, and may also show other irregularities (Stresemann
& Stresemann 1968a). A divergent sequence may speed up primary
moult (Kiat 2017). In all Muscicapa striata, the primaries are always
29
moulted ascendently, but P 10 is usually shed only shortly before or
after P 7 (details see p. 190, Williamson 1960, Diesselhorst 1961,
Stresemann 1963b, Williamson 1972, Salewski et al. 2002b).
Birds known, or supposed, to renew some of their primaries twice
a year follow an eccentric or ascendent sequence during the partial
moult, so that the most exposed primaries are renewed twice a
year. Eccentric primary moult has been observed exceptionally in
Sylvia borin (P 1 + 8 – 9 renewed, see p. 156), S. nisoria (see p. 159,
Hasselquist et al. 1988, Lindström et al. 1993) and Phylloscopus
sibilatrix (Montalto 1988). Locustella fluviatilis regularly moults the
outer one to five long primaries, and usually also P 10, in autumn
before a complete moult in winter which follows the basic sequence;
this partial primary moult probably follows an ascendent sequence
and may be accompanied by the moult of the alula (see p. 150,
Pearson & Backhurst 1983).
In those species with a single primary moult each year, one or a
few primaries may exceptionally be retained. This usually involves
the last primaries to be moulted (P 9 or 10; e.g. Hirundo rustica,
Oenanthe oenanthe, Anthus trivialis, Fringilla coelebs, Spinus spinus,
see part II). In Sturnus vulgaris, however, the remiges to be moulted
last (the outermost primaries) or first (the innermost primaries if
moult is eccentric) may be retained (Schleussner et al. 1985, Evans
1986, Schleussner 1990, Meijer 1991); in the case of an eccentric
primary moult, the suggested sequence is descendent according to
Schleussner (1990), but ascendent according to Evans (1986).
As far as is known, birds with a seasonally divided primary moult
usually follow the normal descendent sequence and resume primary
moult at the point of interruption (suspended primary moult; see
3.3.3). However, Sylvia c. communis may retain some central primaries
before the autumn migration and may show various irregular moult
sequences (Hall & Fransson 2001, G. Nikolaus in litt.; see p. 165 and
3.3.3 for a possible explanation).
Exceptions to the normal ascendent sequence of a complete
secondary moult are few, but frequent in partial secondary moult.
In the course of a complete moult, a few individuals of many
species may shed S 6 before S 5. The secondary moult of Locustella
luscinioides may show a variety of sequences, such as ascendent,
convergent, or irregular (Steiner 1970, Thomas 1977, Müller 1981,
Trias et al. 1982, Neto & Gosler 2006). In Cinclus cinclus, the
secondaries are moulted in a variable sequence, but S 6 usually
before S 4 (see p. 218). Convergent secondary moult was found
in all individuals of Muscicapa striata (see p. 190), Cisticola juncidis
(sequence 1 - 2 - 6 - 3 - 5 - 4; Gauci & Sultana 1981) and in 2.5 % of
Sturnus unicolor (Peris 1988). In Sylvia communis, secondaries are
also moulted convergently (often in the sequence 1 - 2 - 6 - 5 - 3 - 4)
or in a more irregular sequence (see p. 165).
Species which normally renew all of the secondaries during the
complete post-breeding moult may exceptionally retain one, or a
few, of the last secondaries (usually S 6 or S 5 – 6, e.g. Motacilla flava,
Anthus trivialis, Fringilla montifringilla, Acanthis cabaret, Carduelis
citrinella, see part II). Species in which a small percentage of the
individuals retain one, or a few, secondaries during the post-breeding
moult usually start with S 1, but may also renew S 6 in addition to the
outer secondaries. In Spinus spinus and Ficedula hypoleuca, which
retain at least one secondary, 9.3 % and 7.0 % of individuals moult S 6,
respectively (Fig. 29). In Phylloscopus trochilus, this percentage is about
40 % (Fig. 29, cf. Norman 1991b), and in Emberiza hortulana, which
usually retains part or all of the secondaries during the post-breeding
 30 The Moult of Adults

Spinus spinus, autumn Phylloscopus trochilus, autumn

100 100
N=55 N = 44
% of individuals

80 80
60 60
40 40
20 20
0 0
1 2 3 4 5 6 1 2 3 4 5 6
Ficedula hypoleuca, autumn Ficedula hypoleuca, spring
100 100
N=106 N = 182
% of individuals

80 80
60 60
40 40
20 20
0 0
1 2 3 4 5 6 1 2 3 4 5 6

Emberiza hortulana, autumn Sylvia nisoria, spring

100 100
N=41 N = 63
Fig. 30. Some long-distance migrants with a complete prebr moult in the non-breeding
% of individuals

80 80
area may previously perform a rather extensive post-breeding moult in the breeding area.
60 60
In Locustella naevia, it remains unknown whether or not the feathers renewed in Europe
40 40 are moulted again in Africa. Locustella naevia ad after partial postbr moult, 1 September.
20 20 MaC mostly prebr, MeC and GC mostly postbr. T postbr. S 1 – 5 prebr, 6 postbr. Remainder
0 0 of wing prebr. This bird had also renewed the whole body-plumage and R 1 – 6.
1 2 3 4 5 6 1 2 3 4 5 6
Sylvia c. communis, autumn Sylvia c. communis, spring
100 100
N=129 N = 39
% of individuals

80 80
60 60
40 40
20 20
0 0
1 2 3 4 5 6 1 2 3 4 5 6

Muscicapa striata, spring Muscicapa striata, autumn

100 100
N=39 N = 18
% of individuals

80 80
60 60
40 40
20 20
0 0
1 2 3 4 5 6 1 2 3 4 5 6
Secondary position Secondary position

Fig. 29 . Percentage of adults which have retained a given secondary after an otherwise Fig. 31. In certain species performing an extensive pre-breeding moult which includes all
complete moult (dark grey) or have moulted a given secondary after completion of a of the tertials, the innermost secondaries may also be moulted, as shown in this Ficedula
partial moult (light grey), of those individuals (N) with at least one retained (dark grey) albicollis ad  after partial prebr moult, 12 May. MaC and MeC prebr. GC 1 postbr, 2 – 10
or moulted (light grey) secondary. In Muscicapa striata in spring, it is difficult to judge prebr. T prebr. S 1 – 4 postbr, 5 – 6 prebr. Remainder of wing postbr.
whether or not S 6 has been retained (cf. p. 190). Sources: Spinus spinus, Ficedula
hypoleuca, Emberiza hortulana, Sylvia c. communis spring: own data; Phylloscopus
trochilus: Mead & Watmough 1976, Swann & Baillie 1979, own data; Sylvia c. communis
autumn: Mead & Watmough 1976, Swann & Baillie 1979, T. Fransson in litt., own data;
Sylvia nisoria: Hasselquist et al. 1988, Lindström et al. 1993; Muscicapa striata: Mead &
Watmough 1976, Hansen 1985, Rymkevich et al. 1990, own data.

moult, 23 % moult S 6. Sylvia nisoria, which usually retains all, or most,
of the secondaries, may moult S 1 or, if more secondaries are moulted,
sheds S 1 + 2 + 6 almost simultaneously, followed by S 5, although other
(irregular) sequences occur as well and rarely all of the secondaries
are moulted following the basic sequence (Hasselquist et al. 1988,
Rymkevich in Rymkevich et al. 1990, Lindström et al. 1993). In Sylvia
communis, the moult wave which starts at S 6 may often proceed
further, resulting in a convergent sequence (Fig. 29). Long-distance
migrants with a partial post-breeding moult including the tertials
may also replace S 6 along with the tertials (e.g. Anthus campestris,
Locustella naevia, Oriolus oriolus, Lanius collurio, see part II; Fig. 30).
Species which renew the secondaries during an extensive partial
moult do not usually follow the basic ascendent sequence. In Ficedula
hypoleuca, the renewal of secondaries seems to proceed strictly
descendently during the partial pre-breeding moult (all birds which
renewed at least one secondary moulted S 6, Fig. 29) and probably
also in F. semitorquata and F. albicollis (Fig. 31, Roselaar in Cramp &
Perrins 1993). A descendent sequence during the pre-breeding moult
was also observed in Hippolais polyglotta (Pinilla 2001) and actively
moulting Sylvia curruca (see p. 162). Other species which regularly
renew tertials during the pre-breeding moult may also replace S 6
along with the tertials (Phylloscopus collybita, Motacilla flava, Anthus
trivialis, A. spinoletta, see part II; Sylvia cantillans/subalpina, own data).
In Sylvia c. communis, the sequence during the pre-breeding moult
may be descendent, convergent or start with a central secondary
(31 % renewed all the secondaries, 13 % only S 6, 16 % two to five
innermost secondaries, 13 % one or two innermost as well as one
to three outermost, and 3 % one central secondary, N = 39, own
data). A study of actively moulting Sylvia nisoria in Kenya revealed
that secondary moult usually proceeds convergently starting either
with S 6 or S 5 and S 1 or S 2, but irregular sequences may occur as
well (Hasselquist et al. 1988, Lindström et al. 1993). The secondary
moult of Anthus richardi may also show irregularities (Stresemann
& Stresemann 1968a). Muscicapa striata may retain the central
secondaries during the pre-breeding moult and may moult some
central secondaries during the post-breeding moult.
  The Moult of Adults 31

Phylloscopus collybita, N = 49 Sylvia borin, N = 31

Phylloscopus collybita Sylvia borin Muscicapa striata
100 100
100 100 100
N=105 N=62 N=77
% of individuals

80 80 80 80 80

% of individuals
60 60 60
60 60
40 40 40
40 40
20 20 20
0 0 0 20 20
7 8 9 7 8 9 7 8 9
0 0
Motacilla cinerea Motacilla alba Anthus pratensis 1 2 3 4 5 6 1 2 3 4 5 6
100 100 100
N=34 N=109 N=69
Motacilla cinerea, N = 15 Motacilla alba, N = 60
% of individuals

80 80 80
100 100
60 60 60
80 80
40

% of individuals
40 40
20 20 20 60 60
0 0 0
40 40
7 8 9 7 8 9 7 8 9
Tertial position Tertial position Tertial position
20 20

Fig. 32. Percentage of individuals which have moulted a given tertial after completion 0 0
of a partial moult, of those individuals (N) which have moulted at least one tertial 1 2 3 4 5 6 1 2 3 4 5 6
(own data). The individuals are subdivided into those with all three tertials renewed
(black), those with only one tertial renewed (white) and those with two tertials Anthus pratensis, N = 48 Anthus trivialis, N = 161
renewed (grey). In all Motacillidae and in Phylloscopus collybita (pre-breeding moult), 100 100
adults and second-year birds are included. In Sylvia borin and Muscicapa striata (post-
breeding moult) only adults are shown. 80 80

% of individuals
60 60

40 40
In summary, the descendent moult of one or a few innermost 20 20
secondaries only occurs if the tertials have been moulted. Probably, S 6
0 0
or S 5 – 6 are exposed while the protecting tertials are being renewed 1 2 3 4 5 6 1 2 3 4 5 6
and suffer additional wear which may justify their replacement. Rectrix position
Anthus spinoletta, N = 213
Species retaining some secondaries during an otherwise complete 100
moult tend to moult their secondaries convergently, retaining the well
80 other combinations
% of individuals

protected central secondaries. Therefore, they have two small moult

60 R 1 + 6 renewed
gaps within the secondaries which may allow a faster replacement.
R 1 + 2 renewed
During the complementary partial moult, the sequence may be strictly 40
one R only renewed
descendent, convergent, or variable. 20
all R renewed
0
Tertials 1 2 3 4 5 6
Although the most frequent sequence of the tertial moult is T 8-9-7, Rectrix position

it is often varied (usually to 8-7-9), especially in species which shed

them at short intervals. In those species which moult the tertials
during the course of a partial pre-breeding or post-breeding moult,
T 8 is the most frequently moulted, in particular if only one tertial
is renewed (Fig. 32). This suggests a divergent sequence within the
tertials, although some individuals may renew only T 9 or T 7. In
Motacilla alba, individuals renewing only T 7, the longest tertial, are
relatively frequent. Motacilla flava usually moults all the tertials during
the partial pre-breeding moult, and the most frequent sequence is T
8-7-9 (Wood 1976), but the few individuals known to have renewed
only one tertial all replaced T 7 (see p. 224). Rymkevich (in Rymkevich
et al. 1990) reports that Sylvia nisoria may renew only T 7 or T 7 – 8
during the post-breeding moult.
Rectrices
During a complete moult, the rectrices may be shed either: (a) almost
simultaneously; (b) in an irregular sequence (e.g. many individuals of
Acrocephalus dumetorum, Regulus regulus, R. ignicapillus, Sturnus
vulgaris, S. unicolor, Turdus torquatus, Luscinia megarhynchos, L.
luscinia, Oenanthe oenanthe, Cinclus cinclus, Prunella modularis,
Motacilla flava, M. alba, Anthus trivialis, A. spinoletta, A. richardi,
Calcarius lapponicus, Plectrophenax nivalis); (c) in a convergent
sequence within each half of the tail (1-6-2-5-3-4, often modified:
Motacilla alba, M. flava; Glutz von Blotzheim & Bauer 1985); (d)
Fig. 33. Percentage of individuals which have moulted a given rectrix after completion
of an extensive partial pre-breeding or post-breeding moult, of those individuals (N)
which have moulted at least one rectrix (own data). The individuals are subdivided
into those with all six rectrices renewed, those with only one rectrix renewed, those
with R 1 and R 2 renewed, those with R 1 and R 6 renewed, and those with other
combinations. In all Motacillidae and in Phylloscopus collybita (pre-breeding moult),
adults and second-year birds are included. In Sylvia borin (post-breeding moult),
only adults are shown.
R 6 before R 1 (Motacilla cinerea); or (e) R 6 before R 5 or R 4
(Hirundo rustica, Ptyonoprogne rupestris, Anthus pratensis). In the
two Certhia species, R 1 is shed only after the simultaneously shed R
2 – 6 are well grown. As in woodpeckers, this appears to be because
of the importance of the supporting function of the tail (Stresemann
& Stresemann 1966). In Muscicapa striata, moult of the rectrices
proceeds centripetally from R 6 (see p. 190). A centripetal sequence
is also found in some individuals of Sturnus unicolor (Peris 1988),
while S. vulgaris may show nearly centripetal, alternating, or irregular
sequences (Bährmann 1964, 1970). A divergent sequence within
each half of the tail (3-4-2-5-1-6) has been found in some individuals
of six species (Passer moabiticus, Chloris chloris, Linaria cannabina,
Carduelis carduelis, Serinus syriacus, S. serinus, S. pusillus) and may
allow a higher speed of rectrix moult with reduced impact on flight
performance, because first two and later three small gaps are formed
Another random document with
no related content on Scribd:
 COMMON CHERRY CHEESE.

Stone the fruit, or if this trouble be objected to, bruise and boil it
without, until it is sufficiently tender to press through a sieve, which it
will be in from twenty to thirty minutes. Weigh the pulp in this case,
and boil it quickly to a dry paste, then stir to it six ounces of sugar for
the pound of fruit, and when this is dissolved, place the pan again
over, but not upon, a brisk fire, and stir the preserve without ceasing,
until it is so dry as not to adhere to the finger when touched; then
press it immediately into small moulds or pans, and turn it from them
when wanted for table. When the cherries have been stoned, a good
common preserve may be made of them without passing them
through a sieve, with the addition of five ounces of sugar to the
pound of fruit, which must be boiled very dry both before and after it
is added.
Kentish or Flemish cherries without stoning: 20 to 30 minutes.
Passed through a sieve. To each pound of pulp (first boiled dry), 6
oz. sugar. To each pound of cherries stoned and boiled to a dry
paste, 5 oz. sugar.
 CHERRY PASTE. (FRENCH.)

Stone the cherries; boil them gently in their own juice for thirty
minutes; press the whole through a sieve; reduce it to a very dry
paste; then take it from the fire, and weigh it; boil an equal proportion
of sugar to the candying point; mix the fruit with it; and stir the paste,
without intermission, over a moderate fire, until it is again so dry as
to form a ball round the spoon, and to quit the preserving-pan
entirely; press it quickly into small moulds, and when it is cold, paper,
and store it like other preserves.
 STRAWBERRY JAM.

Strip the stalks from some fine scarlet strawberries, weigh, and
boil them for thirty-five minutes, keeping them very constantly stirred;
throw in eight ounces of good sugar, beaten small, to the pound of
fruit; mix them well off the fire, then boil the preserve again quickly
for twenty-five minutes.
Strawberries, 6 lbs.: 35 minutes. Sugar, 3 lbs.: 25 minutes.
Obs.—We do not think it needful to give directions with each
separate receipt for skimming the preserve with care, and keeping it
constantly stirred, but neither should in any case be neglected.
 STRAWBERRY-JELLY.

A very Superior Preserve. (New Receipt.)

The original directions for this delicious jelly, published in the
earlier editions of this work, were the result of perfectly successful
trials made in the summer of their insertion; but, after much
additional experience, we find that the receipt may be better adapted
to our varying seasons, which so much affect the quality of our fruit,
and rendered more certain in its results by some alterations; we
therefore give it anew, recommending it strongly for trial, especially
to such of our readers as can command from their own gardens
ample supplies of strawberries in their best and freshest state. Like
all fruit intended for preserving, they should be gathered in dry
weather, after the morning dew has quite passed off them, and be
used the same day. Strip away the stalks, and put the strawberries
into an enamelled stewpan if at hand, and place it very high over a
clear fire, that the juice may be drawn from them gently; turn them
over with a silver or wooden spoon from time to time, and when the
juice has flowed from them abundantly, let them simmer until they
shrink, but be sure to take them from the fire before the juice
becomes thick or pulpy from over-boiling. Thirty minutes, or
sometimes even longer, over a very slow fire, will not be too much to
extract it from them. Turn them into a new, well-scalded, but dry
sieve over a clean pan, and let them remain until the juice ceases to
drop from them; strain it then through a muslin strainer, weigh it in a
basin, of which the weight must first be taken, and boil it quickly in a
clean preserving-pan from fifteen to twenty minutes, and stir it often
during the time: then take it from the fire, and throw in by degrees,
for every pound of juice, fourteen ounces of the best sugar coarsely
pounded, stirring each portion until it is dissolved. Place the pan
again over the fire, and boil the jelly—still quickly—for about a
quarter of an hour. Occasionally it may need a rather longer time
than this, and sometimes less: the exact degree can only be
ascertained by a little experience, in consequence of the juice of
some varieties of the fruit being so much thinner than that of others.
The preserve should jelly strongly on the skimmer, and fall in a mass
from it before it is poured out; but if boiled beyond this point it will be
spoiled. If made with richly-flavoured strawberries, and carefully
managed, it will be very brilliant in colour, and in flavour really equal
if not superior to guava jelly; while it will retain all the delicious odour
of the fruit. No skimmer or other utensil of tin should be used in
making it; and an enamelled preserving-pan is preferable to any
other for all red fruit. It becomes very firm often after it is stored,
when it appears scarcely set in the first instance; it is, however,
desirable that it should jelly at once.
Fruit kept hot to draw out the juice, 1/2 hour or longer. Boiled
quickly without sugar, 15 to 20 minutes. To each pound 14 oz. of
sugar: 12 to 15 minutes.
 TO PRESERVE STRAWBERRIES OR RASPBERRIES, FOR
CREAMS OR ICES, WITHOUT BOILING.

Let the fruit be gathered in the middle of a warm day, in very dry
weather; strip it from the stalks directly, weigh it, bruise it slightly,
turn it into a bowl or deep pan, and mix with it an equal weight of fine
dry sifted sugar, and put immediately into small, wide-necked bottles;
cork these firmly without delay, and tie bladder over the tops. Keep
them in a cool place, or the fruit will ferment. The mixture should be
stirred softly, and only just sufficiently to blend the sugar and the fruit.
The bottles must be perfectly dry, and the bladders, after having
been cleaned in the usual way, and allowed to become nearly so,
should be moistened with a little spirit on the side which is to be next
to the cork. Unless these precautions be observed, there will be
some danger of the whole being spoiled.
Equal weight of fruit and sugar.
 RASPBERRY JAM.

This is a very favourite English preserve, and one of the most

easily made that can be. The fruit for it should be ripe and perfectly
sound; and as it soon decays or becomes mouldy after it is gathered,
it should be fresh from the bushes when it is used. That which grows
in the shade has less flavour than the fruit which receives the full
warmth of the sun.
Excellent jam for common family use may be made as follows:—
Bruise gently with the back of a wooden spoon, six pounds of ripe
and freshly-gathered raspberries, and boil them over a brisk fire for
twenty-five minutes; stir to them half their weight of good sugar,
roughly powdered, and when it is dissolved, boil the preserve quickly
for ten minutes, keeping it well stirred and skimmed.
Raspberries, 6 lbs.: 25 minutes. Sugar, 3 lbs.: 10 minutes.
 VERY RICH RASPBERRY JAM OR MARMALADE.

No. 1.—Weigh the finest fruit that can be procured, and bruise it
with the back of a wooden spoon after it is put into the preserving-
pan. Boil it gently, keeping it well turned, for about five minutes, then
stir to it gradually nearly or quite its weight of dry pounded sugar, and
continue the boiling rather rapidly for a quarter of an hour or twenty
minutes, and be careful to remove all the scum as it rises. The
preserve will be clear, smooth, and very thick when it is sufficiently
boiled, and should then be taken from the pan without delay, as it will
very quickly set.
No. 2.—Draw gently from the smallest of the raspberries from half
to a whole pound of juice, and boil down in this three pounds of the
fruit, after it has been crushed with a spoon as usual. In ten minutes,
if the fruit be quite ripe, the sugar may be added. Three pounds to
four of the raspberries and their juice, will make a quite sweet
preserve. It should be gradually stirred in until dissolved, and not be
allowed to boil during the time. Ten or fifteen minutes will then suffice
generally to bring it to the proper degree for jellying firmly.
No. 1.—Fine raspberries: 5 minutes. Sugar, nearly or quite equal
weight: 15 to 20 minutes.
No. 2.—Raspberry-juice, 1 lb.; ripe raspberries, 3 lbs. (or 4): 10
minutes. To each pound of fruit and juice, sugar 3/4 lb.: 10 to 15
minutes.
Obs.—All fruit jams are much improved by the addition of a certain
portion of juice to the fruit which is boiled down; they then partake
more of the nature of jelly.
 GOOD RED OR WHITE RASPBERRY JAM.

Boil quickly, for twenty minutes, four pounds of either red or white
sound ripe raspberries in a pound and a half of currant-juice of the
same colour; take the pan from the fire, stir in three pounds of sugar,
and when it is dissolved, place the pan again over the fire, and
continue the boiling for ten minutes longer: keep the preserve well
skimmed and stirred from the beginning.
Raspberries, 4 lbs.; currant-juice, 1-1/2 lb.: 20 minutes. Sugar, 3
lbs.: 10 minutes.
 RASPBERRY JELLY FOR FLAVOURING CREAMS.

Take the stalks from some quite ripe and freshly-gathered

raspberries, stir them over the fire until they render their juice freely,
then strain and weigh it; or press it from them through a cloth, and
then strain it clear; in either case boil it for five minutes after it is
weighed, and for each pound stir in a pound and a quarter of good
sugar reduced quite to powder, sifted, and made very hot; boil the
preserve quickly for five minutes longer, and skim it clean. The jelly
thus made will sufficiently sweeten the creams without any additional
sugar.
Juice of raspberries, 4 lbs.: 5 minutes. Sugar, made hot, 5 lbs.: 5
minutes.
 ANOTHER RASPBERRY JELLY.

(Very Good.)
Bruise the fruit a little, and place it high above a clear fire, that the
juice may be gently drawn from it: it may remain thus for twenty
minutes or longer without boiling, and be simmered for four or five;
strain and weigh it; boil it quickly for twenty minutes, draw it from the
fire, add three-quarters of a pound of good sugar for each pound of
juice, and when this is dissolved place the pan again on the fire, and
boil the preserve fast from twelve to fifteen minutes longer; skim it
thoroughly, and keep it well stirred: the preserve will then require
rather less boiling. When it jellies in falling from the spoon or
skimmer, it is done. Nothing of tin or iron should be used in making it,
as these metals will convert its fine red colour into a dull purple.
Fruit, simmered 5 to 6 minutes. Juice of raspberries, 4 lbs.: 20
minutes. Sugar, 3 lbs.: 12 to 15 minutes. Or: juice of raspberries, 4
lbs.; juice of white currants, 2 lbs.: 20 minutes. Sugar, 4-1/2 lbs.: 10
minutes, or less.
 RED CURRANT JELLY.

With three parts of fine ripe red currants freshly gathered, and
stripped from the stalks, mix one of white currants; put them into a
clean preserving-pan, and stir them gently over a clear fire until the
juice flows from them freely; then turn them into a fine hair-sieve,
and let them drain well, but without pressure. Pass the juice through
a folded muslin or a jelly-bag; weigh it, and then boil it fast for a
quarter of an hour; add for each pound, eight ounces of sugar
coarsely powdered, stir this to it off the fire until it is dissolved, give
the jelly eight minutes more of quick boiling, and pour it out. It will be
firm, and of excellent colour and flavour. Be sure to clear off the
scum as it rises, both before and after the sugar is put in, or the
preserve will not be clear.
Juice of red currants, 3 lbs.; juice of white currants, 1 lb.: 15
minutes. Sugar, 2 lbs.: 8 minutes.
Obs.—An excellent jelly may be made with equal parts of the juice
of red and of white currants, and of raspberries, with the same
proportion of sugar and degree of boiling as in the foregoing receipt.
 SUPERLATIVE RED CURRANT JELLY.

(Norman Receipt.)
Strip carefully from the stems some quite ripe currants of the finest
quality, and mix with them an equal weight of good sugar reduced to
powder; boil these together quickly for exactly eight minutes, keep
them stirred all the time, and clear off the scum—which will be very
abundant—as it rises; then turn the preserve into a very clean sieve,
and put into small jars the jelly which runs through it, and which will
be delicious in flavour, and of the brightest colour. It should be
carried immediately, when this is practicable, to an extremely cool
but not a damp place, and left there until perfectly cold. The currants
which remain in the sieve make an excellent jam, particularly if only
part of the jelly be taken from them. In Normandy where the fruit is of
richer quality than in England, this preserve is boiled only two
minutes, and is both firm and beautifully transparent.
Currants, 3 lbs.; sugar, 3 lbs.: 8 minutes.
Obs.—This receipt we are told by some of our correspondents is
not generally quite successful in this country, as the jelly, though it
keeps well and is of the finest possible flavour, is scarcely firm
enough for table. We have ourselves found this to be the case in
cold damp seasons; but the preserve even then was valuable for
many purposes, and always agreeable eating.
 FRENCH CURRANT JELLY.

Mix one-third of white currants with two of red, and stir them over a
gentle fire until they render their juice freely; pour it from them, strain
and weigh it; for every four pounds break three of fine sugar into
large lumps, just dip them into cold water, and when they are nearly
dissolved boil them to a thick syrup; stir this without ceasing until it
falls in large thick white masses from the skimmer; then pour in the
currant juice immediately, and when the sugar is again dissolved,
boil the whole quickly for five minutes, clear off the scum perfectly,
pour the jelly into jars or warm glasses, and set it in a cool place.
Red currants, two-thirds; white currants, one-third; juice, 4 lbs.;
sugar boiled to candy height, 3 lbs.: jelly boiled, 5 minutes.
Obs.—A flavouring of raspberries is usually given to currant jelly in
France, the preserve being there never served with any kind of joint,
as it is with us.
 DELICIOUS RED CURRANT JAM.

This, which is but an indifferent preserve when made in the usual

way, will be found a very fine one if the following directions for it be
observed; it will be extremely transparent and bright in colour, and
will retain perfectly the flavour of the fruit. Take the currants at the
height of their season, the finest that can be had, free from dust, but
gathered on a dry day; strip them with great care from the stalks,
weigh and put them into a preserving-pan with three pounds of the
best sugar reduced to powder, to four pounds of the fruit: stir them
gently over a brisk clear fire, and boil them quickly for exactly eight
minutes from the first full boil. As the jam is apt to rise over the top of
the pan, it is better not to fill it more than two-thirds, and if this
precaution should not be sufficient to prevent it, it must be lifted from
the fire and held away for an instant. To many tastes, a still finer jam
than this (which we find sufficiently sweet) may be made with an
equal weight of fruit and sugar boiled together for seven minutes.
There should be great exactness with respect to the time, as both
the flavour and the brilliant colour of the preserve will be injured by
longer boiling.
Red currants (without stalks), 4 lbs.; fine sugar, 3 lbs.: boiled
quickly, 8 minutes. Or, equal weight fruit and sugar: 7 minutes.
 VERY FINE WHITE CURRANT JELLY.

The fruit for this jelly should be very white, perfectly free from dust,
and picked carefully from the stalks. To every pound add eighteen
ounces of double refined sifted sugar, and boil them together quickly
for eight minutes; pour it into a delicately clean sieve, and finish it by
the directions given for the Norman red currant jelly (page 559).
White currants, 6 lbs.; highly refined sugar, 6-3/4 lbs.: 6 minutes.
 WHITE CURRANT JAM, A BEAUTIFUL PRESERVE.

Boil together quickly for seven minutes an equal weight of fine

white currants, stalked with the greatest nicety, and of the best sugar
pounded and passed through a sieve. Stir the preserve gently the
whole time, and be careful to skim it thoroughly.
White currants, 4 lbs.; best sugar, 4 lbs.: 7 minutes.
 CURRANT PASTE.

Stalk and heat some red currants as for jelly, pour off three parts of
the juice, which can be used for that preserve, and press the
remainder, with the pulp of the fruit, closely through a hair sieve
reversed; boil it briskly, keeping it stirred the whole time, until it forms
a dry paste; then for each pound (when first weighed) add seven
ounces of pounded sugar, and boil the whole from twenty-five to
thirty minutes longer, taking care that it shall not burn. This paste is
remarkably pleasant and refreshing in cases of fever, and acceptable
often for winter-desserts.
Red currants boiled from 5 to 7 minutes, pressed with one-fourth
of their juice through a sieve, boiled from 1-1/2 to 2 hour. To each
pound 7 oz. pounded sugar: 25 to 30 minutes.
Obs.—Confectioners add the pulp, after it is boiled dry, to an equal
weight of sugar at the candy height: by making trial of the two
methods, the reader can decide on the better one.
 FINE BLACK CURRANT JELLY.

Stir some black currants over the fire until they have yielded their
juice; strain, weigh, and boil it for twenty minutes; add to it three
pounds and a half of sifted sugar of good quality, made quite hot,
and when it is dissolved boil the jelly for five minutes only, clearing
off the scum with care. This, though an excellent preserve, is too
sweet for our own taste, and we think one made with less sugar
likely to be more acceptable in cases of indisposition generally.
Juice of black currants, 4 lbs.: 20 minutes. Sugar, 3-1/2 lbs.: 5
minutes.
 COMMON BLACK CURRANT JELLY.

Boil from three to six pounds of the juice rapidly for twenty
minutes, stirring it well; then mix with it off the fire, half a pound of
sugar for each pound of juice, and continue the boiling for ten
minutes.
Juice of black currants, 3 to 6 lbs.: 20 minutes. To each pound
juice 1/2 lb. good sugar: 10 minutes.
Obs.—This jelly may be made with Lisbon sugar, but will then
require rather more boiling.