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Bonn zoological Bulletin
- Supplementum - Vol. 59 (2014)

formerly: Bonner zoologische Monographien

The Development of Ornithology and
Species Knowledge in Central Europe

Jurgen Haffer (t), Hans Hudde & Brian Hillcoat

An open access journal of organismal zoology, published by

Zoologisches Forschungsmuseum Alexander Koenig - Leibniz Institut fur Biodiversitat der Tiere, Bonn

Editorial

Welcome to the 59th volume of the monograph se-
ries published by the ZFMK. The monograph series of
the ZFMK has been in publication since 1971 , and was
until 2012 known under the name 'Bonner Zoologi-
sche Monographien' (Bonn zoological Monographs),
or BZM.

In order to profit from the experience gained during
the modernization of our journal series, the Bonn zoo-
logical Bulletin, or BzB, monographs will now be pu-
blished as a Supplementum series to the BzB. The Sup-
plementum series will provide a publication outlet for
articles that are too voluminous and 'monographical'
in style to be printed in the BzB. This volume is the first
example of the new Bonn zoological Bulletin, Supple-
mentum. The transition of the BZM to a BzB Supple-
mentum series allows the monograph series to be im-
mediately added to the 'Directory of Open Access Jour-
nals'. We hope that in the future the Supplementum
will achieve an impact factor. Both improvements
would not be possible using the former monograph
format.

The Supplementum series will be free of charge to
the authors, and all recent issues, as well as volumes

55-58, will be accessible online. In the future, we ho-
pe to put all past BZM issues in a freely available on-
line archive, which will further enhance the visibility of
and access to our publications.

This volume has a changed layout and design, to bet-
ter fit the style of our BzB journal series. We thank Eva-
Maria Levermann for her professional layout work and
Uwe Vaartjes for the design of the new cover.

The first contribution to the monograph series in its
new format was written by Jurgen Haffer, Hans Hud-
de and Brian Hillcoat. Until his death, Jurgen Haffer ser-
ved as a long-time member of the journal's Advisory
Board. Brian Hillcoat also worked on numerous pre-
vious BZM issues as an editorial assistant.

We would like to thank Prof. Dr. K.-L. Schuchmann
for serving as editor of the BZM from 2005-201 2 and
wish him the best for his upcoming retirement. We ho-
pe that the transition of the BZM to the BzB-Supple-
mentum series continues to serve readers and authors
in the best possible way.

With best wishes to all readers, the authors, and the
BzB-S team.

Thomas Wesener

Managing Editor, BzB Supplementum,
Myriapoda Curator at the ZFMK
Bonn, October 201 3

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The Development of Ornithology
and Species Knowledge
in Central Europe

Jurgen Haffer (t), Hans Hudde & Brian Hillcoat

Bonn zoological Bulletin - Supplementum 59: 1-116

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Contents Volume 59

Abstract 7

Translator's Preface 7-9

Introduction 9-1 1

Ornithology from the 1 6th to the 1 9th century 1 1 -87

(1 .) Renaissance ornithologists: 16th century 11-14

Hans Sachs - Poet of the Reformation (1531) 1 2-1 3

Eariy writings on European birds - Longoiius and Turner 13

The encyciopedist Conrad Gessner - a Piiny of the Renaissance (1555) 1 3-1 4

(2.) The beginnings of field ornithology: 17th century 14-23

A first iocai naturaiist - Caspar Schwenckfeid (1 603) 1 5-1 6

A compiier - John Jonston (1 650) 1 6-1 7

Ornithoiogists in Engiand - Francis Wiiiughby and John Ray (1676, 1678) 17-23

(3.) The foundations of field ornithology: early and mid- 18th century 23-37

The first ethoiogist - Ferdinand Adam von Pernau (1 702, 1 720) 24-25

Oberjagermeister Hermann Friedrich von Gochhausen (1710, 1732) 25-31

The first book of foiio piates of Centrai European birds -J.L. Frisch (1733-1763) 31-35

The physico-theoiogian Johann Heinrich Zorn and his handbook of ornithoiogy (1742-1743) 35-37

(4.) Standstill in research (1750-1788) 37-38

A compiier - Jacob Theodor Kiein (1 750, 1 760) 38

(5.) The flourishing of field ornithology around 1800 38-63

Johann Andreas Naumann (1789) 38-50

Johann Matthaus Bechstein (1791-1795) 50-56

Johann Andreas Naumann & Johann Friedrich Naumann (1795-1817) 56-61

Friedrich Tiedemann (1 81 0, 1814) 61-63

(6.) "The Golden Age" of field ornithology (1820-1850) 63-76

Johann Friedrich Naumann (1820-1844, 1860) 63-67

Christian Ludwig Brehm (1820-1822, 1823-1824, 1831) 67-68

Friderich Faber (1 822, 1 824-1 827, 1 825-1 826) 68-69

Constantin Wiiheim Lambert Gioger(1833, 1834, 1841-1842) 69-76

(7.) Ornithology of the late 19th and early 20th centuries 76-87

(7a.) Faunistics and iife histories of birds (growth of bird coiiections, ooiogy, 76-80

behaviorai observations, ecoiogy and species richness)

(7b.) Avian systematics (microsystematics and speciation, history of the bioiogicai species concept, 80-87

zoogeography, comparative anatomy, macrosystematics and evoiution)

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Contents Volume 59

Modern ornithology during the 20th century 87-101

(8.) The Stresemann Era 87-101

Erwin Stresemann as an innovator (career, functional morphology, physiology, ethology and 87-96

breeding biology of European birds, migration and orientation, population biology of European birds)

Relations between aviculture and ornithological research 96-97

Applied ornithology 97

Bird artists 97-98

Ornithological research centers in Central Europe 98-99

David Lack and the New Avian Biology in the United Kingdom (1 942-1 947) 99-101

Discussion 101

References 1 02-1 1 5

Online Appendices 115

Endnotes 116

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November 2013

The Development of Ornithology and Species Knowledge in Central Europe

Jiirgen Haffer(t), Hans Hudde" & Brian Hillcoaf"

(f) Tommesweg 60, D-45 149 Essen, Germany
* Rustermark 2, D-45 134 Essen, Germany
** Humboldtstrafie 34, D-14193 Berlin, Germany

Abstract. This work has two distinct aspects, as suggested in its title: an outline of the history of Central European or-
nithology, concluding with the worldwide influence of the “New Avian Biology” of Erwin Stresemann, and an account
of development in the knowledge of European bird species illustrated by extracts from the works of selected ornitholo-
gists up to the 1 9th century. How this increased knowledge was reflected and refined in the gradual rise in the number
of species identified and named is shown in three chronological online Appendices and many illustrations. These tables
of German vernacular names will be a most useful resource for scholars studying early texts. The major figures consid-
ered regarding bird names are Hans Sachs (1495-1576), Marcus zum Lamm (1544-1606), Conrad Gessner (1516-1565),
Caspar Schwenckfeld (1563-1609), John Ray and Francis Willughby (1627-1705) and (1635-1672), H.F. von Gdch-
hausen (1663-1733), J.L. Frisch (1666-1743), J.T Klein (1685-1759), J.H. Zorn (1698-1748), J.A. Naumann (1744-1826),
J.M. Bechstein (1757-1822), and J.F Naumann (1780-1857). As an illustration of one man’s detailed knowledge of field
ornithology, special emphasis is given to a detailed description of J.A. Naumann’s little-known first book Der Vogel-
steller [The Bird-Trapper} of 1789.

The two great research traditions in the discipline - systematic ornithology and field ornithology - arose from the works
of John Ray in England, but after almost three centuries of separate development were only reunited from the early 1 920s,
principally in the work and influence of Erwin Stresemann (1889-1972), especially following the publication of his
in 1927-1934. This could perhaps have taken place earlier, since the “Golden Age” of ornithology in Central Europe
from ca. 1820 to 1850 (J.F. Naumann, C.L Brehm, F. Faber) provided an excellent foundation on which to build further
scientific research, especially after the publication of Charles Darwin’s epochal On The Origin of Species in 1859. But
ornithologists became “distracted” by dealing with the flood of exotic species from the colonies and from sponsored ex-
peditions to the remotest places. Only in the early 20th century did the momentum pick up again with the late accept-
ance of Darwin’s ideas and the work of Stresemann in Berlin. The resulting concentration on avian biology rather than
systematic-faunistic studies meant that gradually the two research traditions came together again. The spread of this New
Avian Biology to the U.S. and U.K. is examined in the lives and work of Ernst Mayr (1904-2005) and David Lack
(1910-1973), respectively.

TRANSLATOR’S PREFACE

Just a few weeks before his shockingly sudden death in
April 2010, Jurgen Haffer contacted me to inquire
whether I would be able to undertake the Job of translat-
ing his latest manuscript. The work was originally to ap-
pear in this journal when it published papers and longer
pieces in German and English, but before he finished his
German manuscript the journaTs policy was changed to
English only. His final illness was beginning to affect his
health to such an extent that he felt he was not able to trans-
late what he had written, something he was normally able
to do better than anyone since he was completely bilin-
gual. His manuscript stops abruptly in the “Return to or-
nithology” part of the section on Gloger in Chapter 6,
though he also completed most of the later Discussion be-
forehand. The Discussion remains as he left it, followed
by his own Acknowledgments.

His life was cut short before he was able to complete
his task, and he had to be satisfied with leaving me a list
of instructions as to how I should continue. These most-
ly took the form of using his own previously published ma-
terial, of which there was an abundance, since the histo-
ry of ornithology, in Central Europe in particular, was his
speciality. The papers he directed me to, from which pas-
sages - at times lengthy - are used in the text, are Haffer
(2006) Blatter aus dem Naumann-Museum 25: 1-55 (in
the section on Tiedemann at the end of Chapter 5);
Glaubrecht & Haffer (2010) Zoosystematics and Evolution
86: 81-115 (in the last part of the section on Gloger in
Chapter 6, where the main manuscript ends); Haffer (2007)
Journal of Ornithology, Supplement 2: S125-S153 (in
Chapter 7); Haffer (2001) Journal fur Ornithologie, Spe-
cial issue 1: 27-93 (in Chapter 8), and Haffer (2008)
Archives of Natural History 35: 76-87 (in the passages
concerning David Eack, etc. in Chapter 8). One “unfortu-

8

Jurgen Haffer (t) et al.

nate” result occurs in Chapter 2, in the section on the great
English naturalist John Ray. Dr. Haffer translated many
pages from Ray’s seminal work The Wisdom of God
(1691) into German, therefore I had no choice but to re-
produce the passages word for word from the English orig-
inal (the facsimile of the modern 1 826 edition), an unusu-
al and not very satisfactory situation. Nevertheless, it
would have been contrary to his intentions to shorten this
passage since the many examples Dr. Haffer quotes from
Ray’s book - one of the two key figures in his argument
- not only show what a brilliant observer Ray was, but
also illustrate Haffer’s extremely important point that all
of the behavioral and physiological phenomena exhaus-
tively listed to show us the Wisdom of God are easily con-
verted into today’s evolutionary adaptations. As his old
friend Ernst Mayr put it (Mayr 1982: 105): “When ‘the
hand of the creator’ was replaced in the explanatory
scheme by ‘natural selection’, it permitted incorporating
most of the natural theology literature on living organisms
almost unchanged into evolutionary biology”.

An additional difficulty for me was that neither an elec-
tronic version of the text nor of any of the illustrations
could be found following Dr. Haffer’s death, entailing a
major reconstruction effort. Only Hans Hudde’s tables of
species names were in an electronic form. With the gen-
erous permission of Herr Hudde, I had to be counted as
a co-author only because of the number of contributions
that I had to undertake, including re-photographing almost
all of the illustrations. I also wrote the Abstract and com-
piled the reference list.

A further consequence of Dr. Haffer’s sudden death was
that he was unable to bring his material up to date. He did
make new insertions in the main text right up until early
2010, but the citations in those sections based on his ear-
lier works are beginning to show their age and many will
have been superseded in recent years. To update these pas-
sages would have entailed an immense amount of work,
so I have confined my additions to citations of newer im-
portant standard publications (handbooks, seminal works
on genetics and other new fields, etc.), or of titles which
I felt Dr. Haffer would have inserted had their absence
been drawn to his attention. Other insertions or remarks
by me in the text are in bold placed inside square brack-
ets. Since the work was originally in German, occasion-
al reference is made to a translation of a publication that
was originally in English. I have left these citations un-
touched, although in general the original will also be in
the reference list.

This indeed, and not the translation itself, has been the
main problem in the preparation of this text for publica-
tion. I feel that it is not acceptable for me as translator to
add any new material to Dr. Haffer’s text, nor indeed to
delete anything substantial, when he is no longer present
to approve or disapprove. The contribution of his trusted
colleague Hans Hudde was restricted to the online Appen-

dices 1-3. 1 have received many useful remarks and sug-
gestions from anonymous referees, but I hope they will
appreciate that for the above reasons I think it would be
inappropriate for me to make substantial alterations.

Having said that, the impressive number of considered
suggestions and corrections I received from Professor Wal-
ter J. Bock of Columbia University deserves special men-
tion. He has gone to great lengths not only in reading my
translation most thoroughly - even to the extent of cor-
recting typos and other small errors - but also in provid-
ing me with quite a few pages of valuable comment. Now
that Jurgen Haffer and Ernst Mayr are no longer with us.
Dr. Bock is probably the leading authority on the history
of ornithology, including the German material, and so his
words carry great weight. Therefore rather than altering
the text that was given to me, or introducing comments
from a third party, I will set out in this Preface the points
made by Dr. Bock.

Walter Bock and Jurgen Haffer were old friends and oc-
casional collaborators, but since they were both active in
the same field it was only natural that they did not agree
with each other all of the time. The facts of history -
names, dates, ideas - are not disputed, but the importance
of ideas and theories for future developments can be ar-
gued about, and not unreasonably Dr. Bock begs to dif-
fer with some of Jurgen Haffer’s interpretations as set out
here. For example, regarding Chapter 8, “The Stresemann
Era”, Professor Bock feels that the influence of Erwin
Stresemann in North America as described there is some-
what exaggerated. He believes that the New Avian Biol-
ogy was well under way in the United States by the time
Stresemann’s work began to have any influence. One rea-
son for this, he asserts, was the less authoritarian struc-
ture of the universities outside Germany, as well as the lack
of the rigid division between museum and university that
hindered career moves in Germany.

Stresemann’s was little read outside of Central Eu-
rope. More than two-thirds of the 2200 printed copies were
in fact destroyed in a warehouse fire at the end of the war
so its physical distribution and influence were necessari-
ly limited. In addition, German research, both in fact and
in reputation, was set back at least a decade by the catas-
trophe of the Second World War. Without the great pros-
elytizing energy of Ernst Mayr in the U.S., history might
have taken a slightly different turn. The influence of
Joseph Grinnell (1877-1939) at the University of Califor-
nia, Berkeley in moving ornithology in the same direction
as Stresemann, whom he greatly admired, was consider-
able but restricted to the academy. His ideas were scat-
tered in papers throughout the scientific literature so
reached a limited number of people. His successor Alden
Miller (1906-1965) had perhaps the greater impact, send-
ing a couple dozen PhD graduates out into the world to
take up successful careers in American ornithology. See
also Bock (2001).

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The Development of Ornithology and Species Knowledge in Central Europe

9

Dr. Bock had much to say on the text as a whole. For
example, he feels that Dr. Haffer did not sufficiently stress
the modernizing role played by Professor Alfred Newton
in the UK at the end of the 19th century. However, al-
though Netvton was a great ornithologist, and his Diction-
ary of Birds (1893-1896) a masterly compendium of
everything that was then known in avian science, and de-
spite the fact that he was one of the first to immediately
recognize the revolutionary power of the theory of evo-
lution by natural selection of Darwin and Wallace, he was
a life-long conservative with little to no interest in field
or laboratory studies. See especially Birkhead & Gallivan
(2012).

Despite the incorporation of older material, this book
is a genuine addition to Jurgen Haffer ’s work in that it
brings together a longer time-span and - with the inclu-
sion of developments in 20th-century Great Britain - a
broader geographical scope than his previous articles. A
fascinating novelty is how the development of species
knowledge is reflected in the gradual increase in the num-
ber of species named. This is made beautifully clear in the
three online appendices of names compiled by his friend
and collaborator of many years, Hans Hudde. These ta-
bles, whose wealth of German vernacular names through
the centuries will be a great help to researchers, represent
a labor of love by Hans Hudde and an admirable achieve-
ment for a man of his advanced years. The slow refining
of species recognition and identification is also made clear
in the many illustrations accompanying the text. A further
aspect of species knowledge that was close to Dr. Haffer’s
heart was to bring to the attention of the wider ornitho-
logical public J.A. Naumann’s first bookDcr Vogelsteller
[The Bird-Trapper^ of 1789. The elder Naumann’s ency-
clopedic knowledge of birds is well illustrated by the
lengthy extracts from this fascinating work.

For their help in a great variety of ways I would like to
express my gratitude to Dr. Karl Schulze-Hagen
(Monchengladbach), Dr. Karl-Ludwig Schuchmann (Zo-
ologisches Forschungsmuseum Alexander Koenig, Bonn),
Dr. Eberhard Mey (Thiiringer Landesmuseum Heidecks-
burg, Rudolstadt), Dr. Sylke Frahnert, Dr. Matthias
Glaubrecht, and Pascal Eckhoff (Museum fur Naturkunde,
Berlin), Hans Hudde (Essen), and naturally to my wife
Northild Hillcoat-Kayser, who checked my translation
most assiduously. I would also like to thank the anony-
mous referees for their constructive suggestions, but in
particular Professor Walter J. Bock for the trouble he took
with his very detailed comments. My deepest thanks go
to Frau Maria Haffer and Frau Amelie Haffer-Penther for
their generous support and encouragement, but above all
for their patience.

Brian Hillcoat Berlin, June 2013

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INTRODUCTION

The beginnings of ornithology in Central Europe lie many
centuries in the past (Fig. 1). Despite this, some ornithol-
ogists, who around 1 800 and early in the 19th century had
published multi- volume handbooks on the birds of Ger-
many, were celebrated in the 20th century as the
“founders”, “creators”, or “fathers” of German-language
ornithology, while by comparison the various achieve-
ments of their important forerunners were ignored. Here
we wish to discuss the work of these early ornithologists
in Europe, especially in the period from the 1 6th to the
1 8th century, when the foundations of European ornithol-
ogy were laid. Until now, the wealth of information on bird
species accumulated by ornithologists of this era has nev-
er been presented in a tabular form or reviewed using com-
parative methods.

Zoologists and botanists began using Einnaean nomen-
clature - in which every species is accorded a Eatin-based
generic name and specific name - in the second half of
the 1 8th century. Thus every animal and plant species had
a unique binomial “label”. At this time almost ah of these
organisms had long possessed one or more German ver-
nacular names, so the new Eatin names for bird species
in the German- speaking areas denoted creatures that were
already familiar to people.

We wish to pursue several aims in this work:

(1) To identify, by using several correlation tables, the Ger-
man bird names employed in the writings of early or-
nithologists that are no longer current, in order to facil-
itate the study of the original texts;

(2) To document the often impressive biological and field-
ornithological knowledge about Central European birds
that existed in the period 1500-1900. It is not our in-
tention to trace the development of faunistic expertise
regarding the geographical distribution of birds in Cen-
tral Europe;

(3) To illustrate the steady increase in the number of
known species in Central Europe over the last few cen-
turies;

(4) To show how Central European field ornithology flour-
ished in the 1 8th century. To do this most effectively we
will review in detail the contents of important publica-
tions from that time which today are almost forgotten
or are extremely difficult to find in the original. Works
such as Ray (1691, 1717), von Gochhausen (1710),
Frisch (1733-1763), and J.A. Naumann (1789) will be
discussed, with many passages being quoted in full;

(5) To encourage greater interest in the ornithological
thinking of these early masters, rather than concentrat-
ing on biographical details.

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10

Jurgen Haffer (t) et al.

Years
2000 n

1900 -

1800 -

1700 -

1600 -

1 500 ■“

1400

1300 -

1200-1

Field

ornithology

Systematic

ornithology

-1800

Linnaeus

Paintings

Gessner
■W Turner

Thirty Years* War
Aldrovandi

Be Ion

Ortus

sanitatis

I

Albsrtus

Magnus

Friedrich

Years

-2000

1900

1700

-1600

1500

Megenberg
(Book ot Nature)

Cantinprt

1400

1300

^ 1200

Fig. 1. The development of ornithology in Central Europe 1200-2000. Asterisks and open bars denote significant publications.
The word “Paintings” indicates the existence of several collections of bird paintings. Abbreviations are: Sch - Schwenkfeld, Jo -
Johnston, H - Hohfeld, M - Marsili, P - Pernau, Go - Gochhausen, Z - Zorn, F - Frisch, Be - Bechstein, B - Buffon (German
translation), N - Naumann, Bre - C.F. Brehm, Fa - Faber, Br - A.E. Brehm, A - Altum. There is no horizontal scale (slightly al-
tered from Haffer 2007a).

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The Development of Ornithology and Species Knowledge in Central Europe

11

During the 19th and into the early 20th century, ornithol-
ogy in Central Europe was deeply divided into systemat-
ic ornithology and field ornithology (the natural history
of birds). In Berlin during the early 1920s, Erwin Strese-
mann (1889-1972) initiated the integration of both
branches into a unified New Avian Biology (Haffer 2001,
2007a; Birkhead 2008; Birkhead & Charmantier 2009).
Other global histories of ornithology have neglected to dis-
cuss these crucially important developments in Central Eu-
rope (e.g. Farber 1982; Walters 2003; Chansigaud 2009).

The roots of the scientific study of birds in Central Eu-
rope lay in the various activities of people interested in
animals and specifically birds, like hunting, trapping, col-
lecting, the keeping of birds in captivity, or simply tak-
ing aesthetic pleasure in the beauty of birds. It was an im-
portant step towards a scientific attitude if an author also
showed an interest in species that were of significance nei-
ther for the kitchen nor for the cage. Bird collections have
existed since the early 16th century; they consisted of
mounted or mummified specimens whose durability, how-
ever, was poor until the employment of arsenic as protec-
tion against invertebrate pests from around the mid- 17th
century (Schulze-Hagen et al. 2003). Some early authors
were primarily motivated by the medical uses of birds.
Among these people with an early interest in the avian
world were emperors and princes, counts, generals, cler-
gymen, artists, teachers, and many other people.

Brief biographies of almost all the ornithologists men-
tioned here have been published by Gebhardt (1964, with
details of additional literature) and can be read there.
Therefore in this paper we will not go into biographical
details of the early ornithologists, but will instead concen-
trate on the results of their work.

Ornithology from the 16th to the 19th century

As early as the Middle Ages, the Hohenstaufen Emperor
Friedrich II, in the general ornithological chapter of his
handbook on falconry {De arte venandi cum avibus, be-
fore 1248), made remarks on the life histories of about 100
bird species (Henss 1970; Kinzelbach 2008). Around the
same time, the Dominican friar Albertus Magnus dealt
with a similar number of species (Balss 1928), an account
that underpinned the basic learning of the early ornithol-
ogists. If one wanted a strict definition of who the “father”
of Central European ornithology was then Friedrich II and
Albertus Magnus would have to be called its “fathers”.
Those ornithologists who were prominent in later centuries
were variously called “old masters”, “pioneers”, or
“trailblazers” who, through their works, ensured impor-
tant advances in the science (Stresemann 1951; Haffer
2006, 2007a).

While nature in earlier times was simply understood to
be a manifestation of the Mind of God, after the 1 5th cen-

Bonn zoological Bulletin - Supplementum 59: 1-116

tury researchers themselves increasingly studied natural

objects.

(1) In the 16th century several naturalists collated their
knowledge in different fields to create comprehensive
encyclopedias; important among them was Conrad
Gessner in Zurich (see online Appendix 1).

(2) In the 17th century we see the beginnings of field or-
nithology, when Caspar Schwenckfeld (1603) and John
Jonston (1650) published their surveys of birds, while
Ray (1676, 1678) along with Willughby gathered togeth-
er the results of their studies in England and Central Eu-
rope in a first handbook of ornithology.

(3) The foundations of field ornithology were laid down
in the first half of the 1 8th century, with the outstand-
ing research work of four men working independently
of each other in northern, central, and southern Ger-
many: A. von Pernau in Sulzbach near Coburg
(Franken), H.F. von Gochhausen in Sachsen-Weimar
(Thiiringen), J.E. Frisch in Berlin and Brandenburg, and
J.H. Zorn in Pappenheim, Sachsen-Coburg (Franken).
(online Appendix 2).

(4) In the following four decades (1750-1788) hardly any
progress was made in field studies (Stresemann 1941a).
Nevertheless, translations of work from other languages
appeared, such as Statius Muller (1773), Buffon
(1772-1809), Pennant (1787), and the overviews com-
piled by Klein (1750, 1760) and Halle (1760).

(5) The flourishing of field ornithology in the late 1 8th
and early 1 9th centuries is typified by the work of Jo-
hann Andreas Naumann (1789), J.M. Bechstein
(1791-1795), and Johann Andreas Naumann & Johann
Friedrich Naumann (1795-1817). (online Appendix 3).

(6) This period was followed by the blossoming of field

ornithology, its “Golden Age”, when J.F. Naumann
(1820-1844, 1860), C.E. Brehm (1820-1822,

1823-1824, 1831), F. Faber (1824-1827, 1825-1826),
and C. Gloger (1834a,b) were active in Central Europe
and published their great works. See Fig. 2.

(1.) Renaissance ornithologists: 16th centnry

The Renaissance period of the 15th and 16th centuries is
characterized by the discovery and re-appropriation of
Classical learning following the era of the “dark” Middle
Ages. The starting point for natural “scientists” were the
zoological, botanical, and mineralogical writings of Ar-
istotle, Theophrastus, and Dioscorides (Baumer 1991).
This knowledge was supplemented in the zoological works
of the encyclopedists Gessner around 1550 and Aldrovan-
di around 1600 by their own observations, as well as those
of their contemporaries, resulting in an all-encompassing
picture. The endeavors of these decades also led to the
emergence of specialized fields, such as comparative

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12

Jurgen Haffer (t) et al.

anatomy, physiology, ornithology, ichthyology, and ento-
mology. The first works devoted solely to birds were those
by G. Longolius of Cologne (1544), the Englishman W.
Turner (1544), and the Frenchman P. Belon (1555), all of
whom were principally interested in identifying the species
mentioned in Classical literature. Nevertheless, these au-
thors did include their own observations in their writing,
while Belon in particular also used good illustrations,
which were absent from the books of his two colleagues.
The popular books written in German on plants and ani-
mals by Adam Lonitzer (Kreuterbuch, 1557) [Herbal] and
Jacob Horst {Von den wunderbarlichen Geheimnissen der
Natur, 1579) [The wonderful secrets of nature^ also con-
tain descriptions of animals, including birds, and were
reprinted many times over the next hundred years.

For these authors and their readers all living things were
holy, since they had been created by God. To occupy one-
self with them could therefore lead to a deeper understand-
ing of the Creator. The beauty of nature led to God through
the contemplation of His works. This philosophy in Ger-

man learning of the 16th and 17th centuries (Crowther-
Heyck 2003) prepared the ground for the rapid spread of
physico- or natural theology in the German- speaking re-
gions at the beginning of the 1 8th century.

Hans Sachs - Poet of the Reformation (1531). Hans
Sachs (1494-1576) was the most important poet of the
Reformation. He lived in Niirnberg, working initially as
a master shoemaker but later becoming committed to the
ideas of the Reformation and to Martin Luther, as ex-
pressed in his aphoristic verses on the “Wittenberg
Nightingale” (Goetze 1890). Sachs and his witty apho-
risms rapidly became famous in Germany as he published
further poems in dialogue form supporting the aims and
dissemination of the Reformation. In his poetry he also
wrote of his home town Niirnberg, composed farces,
Shrovetide plays, comedies, rhyming explanations of
woodcuts, and also vehemently attacked the scourges of
highway robbery and murder. In the 16th century Sachs
was influential in many areas of cultural life. His poem

Years

2000 -

1900 ^

1800 ’

1700 -

^John Ray

Field

ornithology

Buffon

New Avian Biology

E Stresemann

-)(■ Ftirbringer. Gadow
Reichenow
Cabanis

# R0jchenbach, Kaup
Muller I

)Nitzsch Systematic

* . ornithology

Tiedemann

.Meiremj
Statius Muller

copofi I

_y^Moehrtng
■^Kieln *

Linnaeus

Years

r2000

-1900

“1800

-1700

Fig. 2. The development of ornithology in Central Europe 1700-2000 (detail from Fig. 1).

Bonn zoological Bulletin - Supplementum 59; 1-116

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The Development of Ornithology and Species Knowledge in Central Europe

13

of 1 53 1 , Das Regiment der anderthalb hundert Vogel [The
regiment of one-and-a-half hundred birds], reproduced by
Suolahti (1909: 462^72), provides a good overview of
the vernacular names of birds common in the Niirnberg
area. According to our count, 107 species are referred to
(online Appendix 1) which we have correlated with their
modern names using Suolahti (1909). What is missing in
the poem is any mention of ducks or wetland birds, for
which there must have been many names in southern Ger-
many, and only a few raptors appear in the work.

Early writings on European birds - Longolius and
Turner. Gilbertus Longolius (1507-1543) was the first
Renaissance zoologist to publish an exclusively ornitho-
logical book (Baumer 1991 ; Kinzelbach 2012). The man-
uscript, published in 1544 after his death by his friend
William Turner in Cologne, was in the form of a dialogue,
a very popular literary genre at the time. Longolius dis-
cusses with his opposite number Pamphilus the names or
identity of the birds described in Classical literature. The
birds dealt with are Peacock, Jay, Pheasant, Capercaillie,
Hazel Grouse, Black Grouse, Red-legged Partridge,
Guineafowl, Quail, snipe or godwit, plover, thrush, lark,
sparrow, and pigeons. The Red-legged Partridge is men-
tioned as occurring in the central Rhine Valley near Land-
skron Castle (lower Ahr Valley), not far from Remagen,
where this species has not been recorded for hundreds of
years. The death of the author prevented a continuation
of the work.

William Turner (1508-1568), being a non-conforming
Protestant, was several times forced to flee from England
for religious and political reasons, and lived for many
years in Cologne (Kinzelbach 2012). Here his book
praecipuarum, on the identity of birds in the works of Ar-
istotle and Pliny the Elder, appeared in 1544. His com-
mentaries on the individual species are based on his own
observations, for instance the account of the Great Grey
Shrike (Lanius excubitor) quoted by Stresemann (1951:
14-15). A new ornithological science begins with Turn-
er. He treated the following species:

Goshawk, Kingfisher, Mallard, geese, eagle, heron.
Goldfinch, Hazel Grouse, Sylvia warbler. Eagle Owl,
Nightjar, Blue Rock Thrush, White Stork, Hawfinch, pi-
geons, Quail, Carrion Crow, Raven, Pied/Grey Wagtail,
Cuckoo, Black-winged Stilt, Shelduck, Blackcap,
Chaffinch, Brambling, Yellow Wagtail, Coot, terns. Sky
Eark, domestic chickens. Woodcock, Jackdaw, Crane,
martins/swallows, Rarmigan, Greenfinch, Redpoll/Siskin,
Nightingale, gulls. Bee-eater, Blackbird, harrier. Great
Grey Shrike, Bearded Vulture, owl, swan. Com Crake,
Barn Owl, tit, plover, sparrow. Peacock, Grey Partridge,
Pheasant, Phoenix, Magpie, woodpecker, parrot, pelican.
Purple Gallinule, Wren, Robin, Common Redstart,
Nuthatch, Black Grouse, Goldcrest/Firecrest, Moorhen,
thmsh. Hoopoe, and vulture.

Bonn zoological Bulletin - Supplementum 59: 1-116

The encyclopedist Conrad Gessner* - a Pliny of the
Renaissance. In Zurich, after the fashion of the Roman
Pliny the Elder, the universal scholar Conrad Gessner
(1516-1565) (Fig. 3) wrote a fom-y oXurnsNaturgeschichte
der Tiere [Natural history of animals], an encyclopedic
work of reference (“thesaurus” or “pandecf ’) that was de-
signed to replace an entire library (Baumer 1991). For this
he requested information from friends in several European
countries, stressing that his Historia animalium would
contribute to the glorification of the works of God, whose
craftsmanship could especially be discerned in the very
smallest animals, like the ants. The species are ordered al-
phabetically, though with certain divergences from the sys-
tem in the raptors and game-birds, where species that are
obviously closely related are placed together. The avian
volume appeared in 1555 (Historiae animalium liber III:
De Avium Natura). Gessner described many species for
the first time and introduced the Eatinized form of their
names into science. Two hundred years later Einnaeus took
over most of Gessner’s Eatin nomenclature, which is why
many are still in use today, or until very recently (e.g.
Parus cristatus, P. ater, P. palustris, P major). Gessner
named as forerunners of his bird book those of Eongolius
and Turner (see above). He had heard of the ornitholog-
ical publication by the Frenchman Pierre Belon (1555) but
had not been able to see a copy, he wrote.

Das Vogelbuoch (1557) [The book of birds] is a short-
ened German translation in 526 pages of Gessner’s bird
volume of 1555 (779 pages). Since this book was intend-
ed as a popular compendium appealing to the broadest
possible audience, all of the philological embellishment
and mythical creatures were left out.

In his great encyclopedia Ornithologia (1599-1603), the
Italian Ulisse Aldrovandi deliberately turned away from
the idea of listing species alphabetically, using systemat-
ic (including anatomical) concepts to arrive at a “natur-
al” order so that related species would not be artificially
separated from each other. His work is more voluminous
than Gessner’s.

Gessner described 1 82 bird forms fairly precisely, so that
they can be more or less identified today (Ziswiler 1969);
see Figs. 4a-d. Occasionally he classed birds in male and
female or juvenile and adult plumages as belonging to dif-
ferent species. The most difficult identifications are those
of ducks, geese, seabirds, and waders, since Gessner had
very little personal experience of them. On the other hand,
those species that are accurately described (apart from the
very common ones like chickadees (tits), etc.) are the
Alpine birds Wallcreeper, Alpine Accentor, Citril Finch,
Snow Sparrow, Nutcracker, as well as many others, includ-
ing Serin, Waxwing, Stone-curlew, and Water Rail. The
Waldrapp, which still bred in Switzerland in Gessner’s day,
can be identified as today’s Bald Ibis (Geronticus eremi-
ta), now completely extirpated in Europe. Ziswiler
(1969) created a synoptic table of Gessner’s bird names,

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14

Jurgen Haffer (t) et al.

I). Co.VR AI1V3 CkS \'£.RV S ,

£1*3’ at y^irT'tn , j

^t-afruets It^cr .T.ft4t-i3^ J^r'ir/. J^Ai. .

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jc4rx<ii^i*in. # W Jttr-ot&ia i^ua^u-c j-fintUr -i t

.ijr.tiT ji. ^^3 .

Fig. 3. Portrait of Conrad Gessner (from Stresemann 1951;
Bildarchiv des Museums fur Naturkunde, Berlin).

which we have used to compile the list in online Appen-
dix 1.

Springer & Kinzelbach (2009) have treated Gessner as
a “scientist” of the early modern era and have collated a
total of 241 bird species from his books, the most impor-
tant collection of avifaunistic data from 16th-century Cen-
tral Europe.

Numerous bird illustrations by Gessner (1555), Belon
(1555), Cyganski (1585), Aldrovandi (1599-1603), and in
the Thesaurus Picturarum of Marcus zum Lamm
(1577-1606; see Kinzelbach & Holzinger 2000) were
made using preserved birds as models. In Gessner (1555)
in particular, the great majority of the illustrations are of
mounted or mummified specimens (Schulze-Hagen et al.
2003).

Mateusz Cyganski published a popular handbook in
1585 on game-birds. Over 150 species are mentioned and
79 of them depicted in simple woodcuts (Nowak 1985).
The Protestant theologian Johannes Wigand (1586) wrote
about the Common Cuckoo, observed European Nightjars
on the nest, listened to the drumming of the Common
Snipe, and described some details from the life of the Ea-
gle Owl (Hildebrandt 1930).

Bonn zoological Bulletin - Supplementum 59: 1-116

(2.) The beginnings of field ornithology: 17th centnry

The first extensive local avifauna (for Silesia) in Europe
was published at the beginning of the 17th century
(Schwenckfeld 1603), and also at this time several natu-
ralists were collecting color illustrations of Central Euro-
pean birds, but these were never published, or only as se-
lections centuries later (Hackethal 1992, 2001; Haffer
2007a). See especially Mey & Hackethal (2012).

Marcus zum Lamm was a Protestant cleric in Heidel-
berg, where from 1570 onwards he collected three vol-
umes of watercolor paintings of birds from the general re-
gion of the upper Rhine Valley, consisting of 207 species,
of which 60 are exotic species from abroad (see online Ap-
pendix 1). These pictures cover the time span of the cli-
mate change around 1580, when the “Little Ice Age” be-
gan. Marcus zum Lamm recognized the causal relation-
ships between this climatic change and the disappearance
of several “Mediterranean” bird species from the Rhine
Valley (Fig. 5) (Lauterborn’s Theory). Kinzelbach
(1995a,b,c, 1998, 2004), Kinzelbach & Springer (1999),
and Kinzelbach & Holzinger (2000) trace invasions into
Central Europe of Bohemian Waxwing (Bombycilla gar-
rulus). Common Rosefinch {Carpodacus erythrinus).
Common Redpoll (Carduelis JJammea), and European
Serin (Serinus serinus). See online Appendix 1.

The Thirty Years’ War (1618-1648) between Catholic
and Protestant rulers ravaged Central Europe, and it was
to be many decades before countries recuperated from the
catastrophe. No real progress in natural history research,
including of course ornithology, was made during that cen-
tury in the region. Aitinger’s report on bird-trapping
(1626-1631) contains some biological information regard-
ing birds, but the author himself was killed in the chaos
of the war. The text of John Jonston’s Historia naturalis
(1650-1653) was compiled from the handbooks of Gess-
ner and Aldrovandi and contained barely any new infor-
mation, but because it included many plates with simpli-
fied illustrations it was widely read, thus contributing to
a new interest in the natural world after the horrors of war,
as did the last edition of Gessner ’s Vogelbuoch in 1669.

A new dawn in natural history studies, and in ornithol-
ogy in particular, rose with the work of the English nat-
uralist John Ray (1627-1705), who, together with his
young friend and co-worker Francis Willughby
(1635-1672), published a survey of all bird species known
at the end of the 17th century (Ray 1676, 1678). These
books represented a foundation on which the systematic
ornithology of the following centuries could be built. This
seminal work was studied throughout the whole of Europe,
which is why Willughby and Ray’s list of European bird
species is included here. Ray was the first biologist to in-
troduce the concept of successful sexual reproduction as
a criterion for defining a species. Concerning plants, he
wrote: “One species never springs from the seed of an-

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The Development of Ornithology and Species Knowledge in Central Europe

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a

cun

d

Ctftciiacn. Ccxxeu

^PenirrtKtlKiilEininmflrt,

Untb^iK^albaj

&CT

Einbcn^ preM:nfi**

Figs 4a-d. Four plates from Gessner (1582 edition). 4a: Great Crested Grebe; 4b: Eurasian Eagle Owl; 4c: Western Capercail-
lie; 4d: Snow Bunting (top), Rock Bunting.

other nor vice versa.” Species are reproductive commu-
nities that have existed unchanged since the Creation. As
a first account of biological research on flora and fauna
using field studies, Ray’s book The Wisdom of God man-
ifested in the Works of the Creation (1691) is of special
importance. It went into 13 editions and was translated in-
to French and German: Gloria Dei oder Spiegel der Weifi-
heit und Allmacht Gottes: offenbahret in denen Wercken
der Erschaffung ( 1 7 1 7). In this publication, Ray examines
the problems of form and function in birds (and other an-
imals) as well as their adaptation to their environment, dis-
cusses the skill with which birds build their nests (differ-
ently depending on species), points out their lightweight
and hollow bones and their system of air sacs, and de-
scribes details of feather structure and the function of the
preen gland. Physiological aspects of this kind need to be
studied by scientists, Ray stressed.

Bonn zoological Bulletin - Supplementum 59: 1-116

A first local naturalist - Caspar Schwenckfeld (1603).

Schwenckfeld was a pious Protestant, strict believer in the
literal truth of the Bible, doctor, and naturalist, in partic-
ular a botanist (student of Bauhin in Basle and Gessner
in Zurich), and lived in Silesia from 1563 to 1609. He was
the town physician, an official post, firstly in Hirschberg
and at the end of his life in Gorlitz. In his free time he ex-
plored, in the course of many long walks, the world of
plants and animals in the high hills of the Riesengebirge
(Karkonoski in Polish, Krkonose in Czech), also collect-
ing fossils and minerals. In 1600 his Stirpium et fossili-
um Silesiae Catalogus (407 pages) was published in
Leipzig, the very first scientific description of nature in
Silesia, containing the first floristic and paleontological
study of the region. The plants are listed alphabetically ac-
cording to their Latin names. His most important work ap-
peared in 1603 in Bratislava, a fauna of Silesia with the

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Jurgen Haffer (t) et al.

Fig. 5. Male Red-legged Partridge (Marcus zum Lamm; from
Kinzelbach & Holzinger 2000).

title Therio-tropheum Silesiae (563 pages), the “Silesian
zoological garden”, dealing with the mammals, reptiles,
birds, fish, and insects of his homeland - the first local
fauna known anywhere in the world, in which animals are
separated according to where they live, their way of life,
and how they organize themselves. In 1607 he published
a further work, a monograph in German on the spa at
Warmbrunn in Silesia (Cohn 1888; Gebhardt 1964).

Of the 1 54 native birds known to him (online Appen-
dix 1), he described the voice, food, distribution, nest,
number and color of the eggs. The species are in alpha-
betical order, are not illustrated, and are arranged accord-
ing to the following scheme: name (Latin, Greek, Ger-
man), brief description, remarks on reproduction and use-
fulness. The book is a lexicon for reference purposes, in
which some exotic animals also appear (e.g. elephant,
leopard, guinea pig, monkeys, parrot, ostrich). All the
species had already been mentioned in the contemporary
works of Gessner, Aldrovandi, Wotton, Belon, and Ron-
delet. Schwenckfeld probably took many species and the
broad systematics from these authors. He described no
new bird species, while some mythical animals like uni-
corn and phoenix were apparently regarded by him as re-
al. For the compilation of the Central European bird
species known to Schwenckfeld we used a copy of his
book in the Martin Opitz Library in Heme.

Bonn zoological Bulletin - Supplementum 59: 1-116

The opinion widely held in his time that “no organism
is so small that it does not radiate something of the divine”
(Baumer 1996: 186) was also stressed by Schwenckfeld.
Outstanding men of God, mighty kings, and the clever-
est doctors had all investigated the world of nature in or-
der to praise the Creator. Knowledge of nature leads to
knowledge of God was the prevailing philosophy within
which the naturalists pursued their studies.

A compiler - John (Jan or Johannes) Jonston (1650).

He was a Polish doctor, naturalist, and polymath of Scot-
tish extraction (Fig. 6). Following the end of the Thirty
Years’ War (1648), his Historia Natural is Animalium (with
248 plates and almost 3000 illustrations from the work-
shop of the Merian family in Frankfurt a. M.) promoted
the new and growing study of the animal world. The vol-
ume covering the birds was published in 1650 (Fig. 7).
They are divided, as was usual at the time, into landbirds
(birds of prey, insect-eaters, seed-eaters) and waterbirds
(with webbed or unwebbed feet). Their method of feed-
ing is the most important characteristic. Book 1 deals with
the birds of prey. Book 2 with the plant-eaters (seed- and
berry-eaters). Book 3 with the insect-eaters (songbirds and
non-songbirds). Book 4 with the web-footed waterbirds
(meat-, insect- and vegetation-eaters). In Book 6 “exotic
birds” are presented, with an appendix on mythical birds
such as griffin or phoenix.

According to Baumer (1991), “the book on birds is in
general a readable abridgement of the corresponding vol-
ume of Aldrovandi (1599-1603), although supplemented
by quotes from Gessner and contemporary descriptions of
birds from America, especially Brazil. By its appealing
style and restriction to basic scientific details, the impres-
sion given by Jonston’s treatment of birds is that it is al-
most like a forerunner of today’s biological field-guides”
(p. 139), and “by being limited to natural history facts,
fleshed out by a few references to the medicinal uses of
some species, a practical compendium of animals was pro-
duced, illustrated for the first time by copperplate engrav-
ings” (p. 149), which were in the main taken from Al-
drovandi and Gessner (Fig. 8). Jonston’s compendium was
published in many editions in Latin, German, English,
Dutch, and French, and hence was very widely read. Jon-
ston did not include any observations of his own in the
text, which was purely a compilation, the main reason why
Stresemann’s (1951: 24) later judgement was so devas-
tating: “The worst animal books have always found the
most purchasers!”.

Jonston’s scholarship extended to many spheres of learn-
ing, and he spoke 12 languages. He published writings on
medicine, pharmacology, botany, zoology, and history, and
his great desire was to produce textbooks with the aim of
disseminating general knowledge. In this he always fo-
cused on the animal itself, leaving all semantic and em-
blematic aspects to one side. In the spirit of the later physi-

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The Development of Ornithology and Species Knowledge in Central Europe

17

. -Jeh j<er orfrntc

'3nniim^rJj oj'e/r .i

Imrxiiil JOiSJTONli iM . ion jUm tK ur

Jurutsm l/lr ,

j.C' frf

Portrait of John Jonston (from Miller 2008).

Fig. 7.

Title page of Jonston (1650).

co-theologians, he recommended that his readers concern
themselves with natural phenomena because this encour-
aged the adoration of the power and goodness of the
Almighty. Further publications on the character of Jon-
ston’s work are those of Matuszenski (1989) and Miller
(2008).

Ornithologists in England - Francis Willnghby and
John Ray (1676, 1678). Early omithlogists classified birds
in an ecological fashion, on the basis of their preferred
habitats (water, fields, woodland), without finding satis-
factory solutions to the problem of howto categorize birds
in groups. In their important work Ornithologiae Libri
Tres, which initially appeared in Latin (1676) (Fig. 9) and
then in English (The Ornithology of Francis Willughby,
1678) (Fig. 10), Willughby and Ray separated birds into
units that often agree with modern groupings. For their
classification they used the form of birds: bill shape, foot
structure, and body size, similar to Linnaeus in the 1 8th
century. This system was Ray’s idea, while Willughby col-
lected numerous notes on individual species during their
joint travels in England and in western and Central Eu-
rope. See online Appendix 1.

In the years 1663 to 1666 Willughby and Ray journeyed
through Holland and the lower Rhine region, followed the
Rhine Valley to Strasbourg, and then traveled eastwards

Fig. 6.

Fig. 8. Plate from Jonston (1650) with various flycatchers,
thmshes, and warblers.

Bonn zoological Bulletin - Supplementum 59: 1-116

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18

Jurgen Haffer (t) et al.

to Vienna from where they moved south to Italy. In their
textbook, Ornithology, many records of species that they
found on markets in Germany and Italy can be found. In
addition, they purchased color illustrations in Strasbourg,
Niimberg and other cities (Charmantier & Birkhead 2008).
In this ornithological handbook, which was largely con-
ceived by Ray, they described every species so exactly that
they can be easily recognized, and repeat descriptions of
male and female or juvenile and adult were avoided. Ray
was dissatisfied with the quality of the illustrations, al-
though he still regarded them as better than other contem-
porary copperplate engravings of birds (Figs. 11a, b). The
plates are black and white, though at least one copy ex-
ists with hand-colored plates, in the library of the famous
diarist Samuel Pepys (Montgomerie & Birkhead 2009).
Here, the colors of most of the depictions are good, and
the species easily identifiable, though the toucan on Plate
20 (“Pica Brasiliensis Toucan”) resembles no existing bird,
appearing to be a conglomeration of several species.
Raven (1942, 1950) provides a clear and detailed descrip-
tion of Ray’s work.

Many 17th-century theological scholars believed that the
study of nature could not be approved of for Christians:
nature represents a temptation leading away from the path
to redemption and salvation for mankind. John Ray in
Cambridge preached the reverse: that the study of plants
and animals is pleasing to God, revealing his plan in the
Creation. At the age of 60, Ray brought his sermons and

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the results of his decades of nature study together in his
most popular book, written in English rather than the Lat-
in of his other works, which appeared in London in 1691 :
The Wisdom of God manifested in the Works of the Cre-
ation (Fig. 12). The German translation of 1717 carried
the title Gloria Dei oder Spiegel der Weifiheit und All-
macht Gottes: offenbahret in denen Wercken der Erschaf-
fung. In this book, Ray sets out his concept of natural or
physico-theology, in which living organisms are in a
miraculous way adapted to their environment and hence
are suitable objects of study, with the aim of praising God
and recognizing the plan of the Creator. By stressing the
significance of form and function, with his extensive
works on plants, birds, mammals, fish, and insects be-
tween 1676 and 1713, Ray laid the intellectual foundations
for the ornithologists, entomologists, and other specialists
in 18th-century Europe. He encouraged naturalists to
closely observe plants and animals in their natural habi-
tats in order to celebrate the wisdom of the Creator of the
world. He saw the order of the universe, the life of plants
and animals, and the functioning of nature as a manifes-
tation of the Holy Spirit (Raven 1942).

What was new in The Wisdom of God was that Ray
moved from the simple tasks of a naturalist (identifica-
tion, description, classification) to an interpretation of the
biological significance of structural characters and phys-
iological processes, and an investigation of the adaptations
of organisms to their environment, their behavior and in-

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Bonn zoological Bulletin - Supplementum 59: 1-116

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Figs 11a, b. Two plates from Ray (1678). 11a; ducks and Anhinga sp.; 11b: thrushes and Common Starling.

stincts (Raven 1942: 452^53). In order to stress the cru-
cial importance of this naturalist to the development of or-
nithology we feel it appropriate to quote at length from
his pioneering work. In the following passages we sum-
marize his most important conclusions relating to birds,
using the 1 826 edition that was reprinted by the Ray So-
ciety in 2005. Key statements are emphasized in italics,
while any added comments of ours appear in square brack-
ets.

In the Preface (p. ix) Ray assures the reader that he has
“been careful to admit nothing for matter of fact, or ex-
periment, but what is undoubtedly true, lest [he] should
build upon a sandy and ruinous foundation; and by the ad-
mixture of what is false, render that which is true suspi-
cious”, and continues: (1) “The belief of a deity being the

foundation of all religion must be demonstrated by

arguments drawn from the light of nature and works of
creation you may hear illiterate persons affirm-

ing that they need no proof of the being of a God, for that
every pile of grass, or ear of corn, sufficiently proves that:
for ... all the men of the world cannot make such a thing
as one of these; and if they cannot do it, who can, or did
make it but God? To tell them that it made itself, or sprung
up by chance, would be as ridiculous as to tell the great-
est philosopher so” (pp. x-xi). (2) This argument here
“serves not only to demonstrate the being of a Deity, but

Bonn zoological Bulletin - Supplementum 59: 1-116

also to illustrate some of his principal attributes; as name-
ly, his infinite power and wisdom the sun and moon,

and all the heavenly hosts, are effects and proof of his
almighty power the adapting all the parts of ani-

mals to their several uses: the provision that is made for
their sustenance” (p. xi). By the “Works of the Creation”
of his title, Ray means ""the works created by God at first,
and by him conserved to this day in the same state and
condition in which they were at first made"? So for Ray
there were no changes and no evolution, only the constan-
cy of the universe and all plant and animal species: “Con-
servation is a continued creation”.

“Animate bodies are divided into four great genera or
orders, beasts [chiefly quadrupeds, with mammals, rep-
tiles, etc.], birds, fishes, and insects.”

“The species of beasts, including also serpents, are not
very numerous; of such as are certainly known and de-
scribed, I dare say not above 150 1 reckon all dogs

to be of one species, they mingling together in generation,
and the breed of such mixtures being prolific [= defini-
tion of a species!].

“The number of birds known and described may be near

500 we may suppose the whole sum of beasts and

birds to exceed by a third part, and fishes by one half, those
known” (p. 24). Regarding the plants, “there are a great
many, I might say some hundreds, put down for different

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Jurgen Haffer (t) et al.

Fig. 12. Title page of John Ray’s Wisdom of God ... (1701 edi-
tion).

species, which in my opinion are but accidental varieties'"
in respect of their characters, which are not “sufficient to
constitute a specific difference” (p. 27).

“If the number of creatures be so exceeding great, how
great, nay, immense, must needs be the power and wis-
dom of him who formed them all!” (p. 27). “God can and

doth by different means produce the same effect the

various ways of extracting the nutritious juice out of the
aliment, in several kinds of creatures. (1) In man and the
viviparous quadrupeds, the food moistened with the spit-
tle (saliva) is first chewed and prepared in the mouth, then

swallowed into the stomach (2) In birds there is no

mastication or comminution of the meat in the mouth; but
in such as are not carnivorous, it is immediately swallowed
into the crop or craw where it is moistened and mol-
lified by some proper juice And thence transferred

into the gizzard or musculous stomach, where by the

working of the muscles and by the assistance of small

pebbles (which the creature swallows for that purpose) it
is, as it were, by millstones ground small, and so trans-
mitted to the guts” (p. 30). “The works of God are

all very wisely contrived and adapted to ends both par-
ticular and general” (p. 31).

Preservation of species by sexual reproduction. “It is

the great design of Providence to maintain and continue
every species Why can we imagine all creatures

Bonn zoological Bulletin - Supplementum 59: 1-116

should be made male and female but to this purpose?

Why in viviparous animals, in the time of gestation, should
the nourishment be carried to the embryon in the womb,
which at other times goeth not that way? When the young
is brought forth how comes all this nourishment then to

be transferred from the womb to the breasts or paps ?

a great proof and instance of the care that is taken, and
provision made for the preservation and continuance of
the species” (pp. 101-102).

“That birds should all lay eggs, and none bring forth

live young, is a manifest argument of divine providence,

designing thereby their preservation For if they had

been viviparous, the burthen of their womb, if they had
brought forth any competent number at a time, had been
so great and heavy, that their wings would have failed
them, and had become an easy prey to their enemies: or
if they had brought but one or two at a time, they would
have been troubled all the year long with feeding their
young, or bearing them in their womb” (p. 103).

To facilitate the transition of a chick from liquid nour-
ishment in the egg to solid food after hatching “nature hath
provided a large yolk in every egg, a great part whereof
remaineth after the chicken is hatched, and is taken up and
enclosed in its belly, and by a channel made on purpose
received by degrees into the guts, and serves instead of
milk to nourish the chick for a considerable time” (p. 104).
“That birds which feed their young in the nest, though in
all likelihood they have no ability of counting the num-
ber of them, should yet (though they bring but one morsel
of meat at a time, and have no fewer .... than seven or
eight young in the nest together, which at the return of their
dams, do all at once with equal greediness, hold up their

heads and gape) do not omit or forget one of them

This seems to me most strange and admirable, and beyond

the possibility of a mere machine to perform When

they have laid such a number of eggs, as they can conve-
niently cover and hatch, they give over and begin to sit.""
However, that this behavior can be circumvented, as in do-
mestic chickens, Ray learned from his friend Dr. Lister:
“one and the same swallow, by the subtracting daily of her
eggs, proceeded to lay nineteen successively and then gave
over”. On the subject of the growth of the young, Ray ob-
served: “the marvellous speedy growth of birds that are

hatched in nests till they be fledged and come almost

to their full bigness; at which perfection they arrive with-
in the short term of about one fortnight, seems to me an
argument of providence designing thereby their preserva-
tion, that they might not lie long in a condition exposed
to the ravine of any vermin that may find them, being ut-
terly unable to escape or shift for themselves” (p. 105).
In building their nests, the adult birds “search out a se-
cret and quiet place, where they may be secure and undis-
turbed in their incubation; then they make themselves
nests, every one after his kind, that so their eggs and young
may lie soft and warm, and their exclusion and growth be

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promoted. These nests, some of them so elegant and ar-
tificial, that it is hard for man to imitate them and make

the like. I have seen nests of an Indian bird which they

hang on the end of the twigs of trees over the water, to
secure their eggs and young from the ravage of apes and
monkeys After they have laid their eggs, how diligent-

ly and patiently do they sit upon them till they be hatched

with such an ardent and impetuous desire of sitting

are they inspired, that if you take away all their eggs, they

will sit upon an empty nest After their young are

hatched, for some time they do almost constantly brood
them under their wings, lest the cold and sometimes per-
haps the heat should harm them. All this while also they

labour hard to get them food with what courage they

are at that time inspired, that they will even venture their
own lives in defence of them. The most timorous, as hens
and geese, become then so courageous, as to dare to fly
in the face of a man that shall molest or disquiet their

young so eminent pieces of self-denial, must needs

be the works of Providence, for the continuation of the
species and upholding of the world: especially if we con-
sider that all this pains is bestowed upon a thing which
takes no notice of it, will render them no thanks for it, nor
make them any requital or amends: and also, that after the
young is come to some growth, and able to shift for itself

it takes no further care of it, but will fall upon it, and

beat it indifferently with others.”

“One necessary to the conservation of the species of an-
imals; that is, the keeping up constantly in the world a due
numerical proportion between the sexes of male and fe-
male, doth necessarily infer a superintending Providence.
For did this depend only upon mechanism, it cannot well
be conceived, but that in some ages or other there should
happen to be all males, or all females; and so the species
fail” (pp. 106-107).

Breeding times. Mr Boyle pointed out in his writings, “the
conveniency of the season (or time of year) of the pro-
duction of animals, when there is proper food and enter-
tainment ready for them” (p. 108).

Instincts in animals. “I shall take notice of the various
strange instincts of animals; which will necessarily
demonstrate, that they are directed to ends unknown to
them, by a wise superintendant. As, 1 . That all creatures
should know how to defend themselves, and offend their
enemies; where their natural weapons are situate, and how
to make use of them A boar knows the use of his tush-

es; a dog of his teeth; a horse of his hoofs; a cock of his

spurs 2. That those animals that are weak, and have

neither weapons nor courage to fight, are for the most part
created swift of foot or wing, and so being naturally tim-
orous, are both willing and able to save themselves by
flight. 3. That poultry, partridge, and other birds, should
at the first sight know birds of prey, and make sign of it

Bonn zoological Bulletin - Supplementum 59: 1-116

by a peculiar note of their voice to their young, who

presently thereupon hide themselves 4. That young

animals, as soon as they are brought forth, should know
their food. As for example: such as are nourished with
milk, presently find their way to the paps, and suck at them

5. That such creatures as are whole-footed, or fm-toed,

viz. some birds and quadrupeds, are naturally directed to
go into the water and swim there, as we see ducklings,
though hatched and led by a chicken, if she brings them
to the brink of a river or pond of water, they presently
leave her, and in they go, though they never saw any such
thing done before; and though the hen clucks and calls,
and doth what she can to keep them out: this Pliny takes

notice of every part in animals is fitted to its use, and

the knowledge of this use put into them 6. Birds of

the same kind make their nests of the same materials, laid
in the same order, and exactly of the same figure, so that
by the sight of the nest one may certainly know what bird

it belongs to. And this they do though taken out of

the nest and brought up by hand; neither were any of the
same kind ever observed to make a different nest, either

for matter or fashion And therefore, as Dr. Cudworth

saith well, they are not masters of that wisdom according
to which they act, but only passive to the instincts and im-
presses thereof upon them” (pp. 110-113)

The migrations of birds. “I know not how to give an ac-
count of it, it is so strange and admirable. What moves
them to shift their quarters? You will say. The disagree-
ableness of the temper of the air to the constitution of their
bodies, or want of food. But how come they to be direct-
ed to the same place yearly, though sometimes but a lit-
tle island . . . ? The cold or the heat might possibly drive

them in a right line from either, but that they should im-
pel land birds to venture over a wide ocean, of which they
can see no end, is strange and unaccountable: one would
think that the sight of so much water, and present fear of
drowning, should overcome the sense of hunger, or dis-
agreeableness of the temper of the air. Besides, how come
they to steer their course aright to their several quarters,
which before the compass was invented was hard for man

himself to do Think we that the quails, for instance,

could see quite across the Mediterranean sea? And yet, it
is clear, that they fly out of Italy into Africa: lighting many
times on ships in the midst of the sea, to rest themselves

when tired and spent with flying How these

[salmon], when they have been wandering a long time in
the wide ocean, should again find out and repair to the
mouths of the same rivers, seems to me very strange, and
hardly accountable, without recourse to instinct, and the
direction of a superior cause” (pp. 113-114).

Nest hygiene. “In young birds that are fed in the nest, the
excrement that is voided at one time is so viscid, that it

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hangs together in a great lump, as if it were enclosed in
a film, so that it may easily be taken up, and carried away
by the old bird in her bill. Besides, by a strange instinct,

the young bird elevates her hinder parts so high that

she seldom fails to cast what comes from her clear over
the side of the nest. So we see here a double provision
made to keep the nest clean ” (pp. 115-116).

“No birds of prey are gregarious. Again, that such crea-
tures do not greatly multiply They for the most

part breeding and bringing forth but one or two, or at least
a few young ones at once: whereas they that are feeble
and timorous are generally multiparous, or if they bring
forth but few at once, as pigeons, they compensate that
by their often breeding, viz. every month but two
throughout the year” (p. 121).

Adaptations of the body to the way of life. “I shall note
the exact fitness of the parts of the bodies of animals to
every one’s nature and manner of living woodpeck-

ers .. have a tongue which they can shoot forth to a very
great length, ending in a sharp stiff bony tip, dented on
each side; and at pleasure thrust it deep into the holes,
clefts, and crannies of trees, to stab and draw out coflfi [lar-
vae], or any other insects lurking there, as also into ant-
hills, to strike and fetch out the ants and their eggs. More-
over, they have short, but very strong legs, and their toes
stand, two forwards, two bacl<wards, which disposition (as
Aldrovandus well notes) nature, or rather the wisdom of
the Creator, hath granted to woodpeckers, because it is
very convenient for the climbing of trees, to which also
conduces the stiffness of the feathers of their tails, and their
bending downward, whereby they are fitted to serve as a

prop for them to lean upon The swallow, whose

proper food is small beetles, and other insects flying about

in the air is wonderfully fitted for the catching of these

animalcules; for she hath long wings, and a forked tail,
and small feet, whereby she is, as it were, made for swift

flight and she hath also an extraordinary wide mouth,

so that it is very hard for any insect that comes in her way

to escape her The colymbi, or douchers, or loons,

whose bodies are admirably fitted and conformed for div-
ing under water: being covered with a very thick
plumage; and the superficies of their feathers so smooth
and slippery, that the water cannot penetrate or moisten
them: whereby their bodies are defended from the cold,
the water being kept at a distance; and so poised, that by
a light impulse they may easily ascend in it. That their feet
are situate in the hindmost part of their body, whereby they
are enabled, shooting their feet backwards, and striking
the water upwards, to plunge themselves down into it with
great facility, and likewise to move forwards therein. Then
their legs are made flat and broad, and their feet cloven
into toes with appendant membranes on each side; by

which configuration they easily cut the water Their

bills are also made straight and sharp for the easier cut-

Bonn zoological Bulletin - Supplementum 59; 1-116

ting of the water, and striking their prey. Could we see the
motions of their legs and feet in the water, then we should
better comprehend how they ascend, descend, and move
to and fro; and discern how wisely and artificially their
members are formed and adapted to those uses” (pp.
125-127).

“In birds all the members are most exactly fitted for the
use of flying. First, the muscles which serve to move the
wings are the greatest and strongest, because much force
is required to the agitation of them; the underside of them
is also made concave, and the upper convex, that they may
be easily lifted up, and more strongly beat the air, which
by this means doth more resist the descent of their body
downward. Then the trunk of their body doth somewhat
resemble the hull of a ship; the head, the prow, which is
for the most part small, that it may the more easily cut the

air, and make way for their bodies That the train serves

to steer and direct their flight, and turn their bodies like
the rudder of a ship, is evident in the kite, who by a light
turning of his train, moves his body which way he pleas-
es those that have long legs, have for the most part

short tails; and therefore whilst they fly, do not, as oth-
ers, draw them up to their bellies, but stretch them at length
backwards, that they may serve to steer and guide them
instead of tails. Neither doth the tail serve only to direct
and govern the flight, but also partly to support the body,
and keep it even” (pp. 127-128).

“ the bodies of birds are small in comparison of

quadrupeds, that they may more easily be supported in the
air during their flight; which is a great argument of wis-
dom and design: else why should not we see species of
pegasi, or flying horses, of griffins, of harpies, and a hun-
dred more, which might make a shift to live well enough,

notwithstanding they could make no use of their wings

Besides, their bodies are also hollow and light; nay,

their very bones are light: for though those of the legs and
wings are solid and firm, yet have they ample cavities, by

which means they become more rigid and stiff Then

the feathers also are very light, yet their shafts hard and
stiff, as being either empty or filled with a light and spongy
substance; and their webs are not made of continued mem-
branes, for then had a rupture by any accident been made
in them, it could not have been consolidated, but of two
series of numerous plumulae, or contiguous filaments, fur-
nished all along with hooks on each side, whereby catch-
ing hold on one another, they stick fast together And

for their firmer cohesion, the wise and bountiful Author
of nature hath provided and placed on the rump two glan-
dules, having their excretory vessels, round which grow
feathers in form of a pencil, to which the bird turning her
head, catches hold upon them with her bill, and a little
compressing the glandules, squeezes out and brings away
therewith an oily pap or liniment, most fit and proper for
the inunction of the feathers, and causing their little fila-
ments more strongly to cohere And because the bird

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is to live many years, and the feathers in time would, and
must necessarily be worn and shattered, nature hath made
provision for the casting and renewing of them yearly.
Moreover, those large bladders or membranes, extending
to the bottoms of the bellies of birds, into which the breath
is received [= the air-sac system], conduce much to the
alleviating of the body, and facilitating the flight: for the
air received into these bladders, is by the heat of the body
extended into twice or thrice the dimensions of the exter-
nal, and so must needs add a lightness to the body. And
the bird, when she would descend, may either compress
this air by the muscles of the abdomen, or expire as much
of it as may enable her to descend swifter, or slower, as
she pleases. I might add the use of the feathers in cher-
ishing and keeping of the body warm And for this rea-
son we see, that water-fowls, which were to swim and sit
long upon the cold water, have their feathers very thick
set upon their breasts and bellies, and besides a plentiful
down there growing, to fence against the cold of the wa-
ter, and to keep off its immediate contact.”

“That the tails of all birds in general do not conduce to
their turning to the right and left, according to the com-
mon opinion, but rather for their ascent and descent, some
modem philosophers have observed and proved by exper-
iment but in those that have forked tails, autopsy con-

vinceth us, that it has this use For it is manifest to

sight, that the fork-tailed kite by turning her train side-
ways, elevating one horn, and depressing the other, turns
her whole body. And doubtless the tail hath the same use
in swallows, who make the most sudden turns in the air
of any birds, and have all of them forked tails” (pp.
129-131).

“I shall now add another instance of the wisdom of na-
ture, or rather the God of nature, in adapting the parts of
the same animal one to another, and that is the proportion-
ing the length of the neck to that of the legs the necks

of birds and quadrupeds are commensurate to their legs,
so that they which have long legs have long necks, and
they that have short legs short ones .... Only in these too
there is an exception, exceeding worthy to be noted, for
some water-fowl, which are palmipeds, or whole-footed,
have very long necks, and yet but short legs, as swans and
geese wherein we may observe the admirable provi-

dence of Nature. For such birds as were to search and gath-
er their food, whether herbs or insects, in the bottom of
pools and deep waters, have long necks for that purpose,
though their legs, as is most convenient for swimming, be

but short Whereas there are no land-fowl to be seen

with short legs, and long necks, but all have their necks

commensurate to their legs For Nature makes not a

long neck to no purpose” (pp. 136-138).

“Lastly, another argument of providence and counsel re-
lating to animals, is the various kinds of voices the same
animal uses on divers occasions, and to different purpos-
es. Hen-birds, for example, have a particular sort of voice,

Bonn zoological Bulletin - Supplementum 59: 1-116

when they would call the male; which is so eminent in
quails, that it is taken notice of by men, who by counter-
feiting this voice with a quail-pipe, easily draw the cocks
into their snares. The common chicken all the while she
is broody sits, and leads her chicks, uses a voice which
we call clucking. Another when upon sight of a bird of
prey, or apprehension of any danger, she would scare them,
bidding them, as it were, to shift for themselves, where-
upon they speedily run away, and seek shelter among

bushes, or in the thick grass These actions do indeed

necessarily infer knowledge and intention of, and direc-
tion to, the ends and uses to which they serve; not in the
birds themselves, but in a superior agent, who hath put an
instinct in them of using such a voice upon such an oc-
casion; and in the young of doing that upon hearing of it

Other voices she hath when angry, when she hath

laid an egg, when in pain, or great fear, all significant

which yet are all argumentative of Providence, intending
their several significances and uses” (pp. 138-139).

Leonhard Baldner (1612-1694) was a Strasbourg fish-
erman and councillor who commissioned paintings from
Johann Jakob Walther, a well-known artist of that city, for
his Vogel-, Fisch- und Thierbuch (1646-1687) [Bird, fish,
and animal book~\, in which he made a zoological inven-
tory of the Rhine and its surroundings that included 72
birds, 45 fish and crustaceans, and 52 other animals. When
Willughby and Ray passed this way in 1663 they pur-
chased copies of the text and some paintings by Baldner
that can still be seen in London. See Baldner (1973).

Birkhead & van Balen (2008) demonstrated the impor-
tant, but largely unrecognized role of European bird-keep-
ing in the development of several different areas of bird
study, especially song acquisition, function and anatomy,
territorial behaviory, breeding biology, external genitalia,
migration, instinct, and learning.

(3.) The foundations of field ornithology: early and
mid-18th century

Physico-theological or natural theological ideas spread at
the beginning of the 18th century from England (see
above) and Holland to Germany, mainly to the Protestant
areas. Their supporters endeavored to attain knowledge of
God, independent of scripture but through the study of na-
ture, regarded as untainted by human activity (Krolzik
1980, 1996; Waschkies 1988; Jahn 1989, 1998: 7-8, 2000;
Baumer 1996: 197). On the basis of their investigations
of useful, orderly, or simply beautiful structures (adapta-
tions) or processes in nature, the existence and properties
of God, His power, wisdom, and goodness, could be
demonstrated or inferred. These concepts induced large
numbers of European naturalists in the 1 8th century to
leave their libraries and go into the field to carry out stud-
ies on living molluscs, insects, frogs, birds, and other an-

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imal groups, as well as plants. They were, following John
Ray, physico-theologians; in Germany for instance, the
well-known naturalists J.A. Fabricius, F.C. Lesser, A.J.
Roesel von Rosenhof, H.S. Reimaais, C.K. Sprengel, J.R.
Forster; ornithologists like A. von Pernau, J.L. Frisch, and
J.H. Zorn. The physico-theological philosophical frame-
work of these researches led to important discoveries con-
cerning adaptations of plants and animals clearly “de-
signed” for a purpose, the ways of life of an assortment
of animals, and also the close adaptive relationships be-
tween flowering plants and insects with respect to polli-
nation (C.K. Sprengel). The clear “adaptedness” and har-
mony of many natural phenomena were taken as proofs
of the existence of a wise Creator (“Argument from De-
sign”). Nature study was divine worship, and nature was
“read” as the book of God’s creation. Later, the wealth of
biologically significant results gathered by these enthusi-
asts could simply be transferred to modern biology, fol-
lowing the replacement of the physico-theology of the
field naturalists of the 1 8th and 1 9th centuries with the new
evolutionary paradigm (Mayr 1984: 86).

William Derham, a student and friend of John Ray in
England, published in 1713 an important book on Physi-
co-Theology (see References for full title), which appeared
in a German translation in 1730: Physiko-Theologie oder
Natur-Zeitung zu GOTT [Physico-Theology or Nature Re-
ports to GOD]. It contains observations on the Earth, on
animals in general, on mankind, quadrupeds, birds, insects,
crawling animals, and on plants. In the chapter Observa-
tions of Birds, the clear fitness for purpose of the form and
structure of the body, and of the wings for flying, is dis-
cussed. The legs of wading bird are naked and long; the
feet of swimming species have webs; those of raptors have
curved talons and also the bill is hooked, both for seizing
and holding prey; a supporting tail and short legs serve
woodpeckers well in climbing trees. The interlocking bar-
bules of feather vanes, with their booklets, which Derham
observed through a microscope, are precisely described
and illustrated. He wrote that migratory birds know when
to start their journey and when to return, and that the Cre-
ator has determined these times and given the birds an in-
stinct when to begin migration and which route to fly,
across alien seas and countries. The artistic and purpose-
ful structure of their nests, and the great variety that ex-
ists, the different nest sites in trees, on the ground, on
rocks, etc., and the natural instinct to choose the appro-
priate site at the right time of year are all described in great
detail. All these works “shine forth with the great Creator’s
infinite wisdom, counsel, and providence” (p. 902, 1741
edition).

The tendency of ornithologists in the 1 8th and 19th cen-
turies to concentrate either on the systematics of birds or
on their natural history (way of life) led to a fundamen-
tal bifurcation in the science, which was to persist for a
period of over 200 years (Haffer 2006, 2007a; see Fig. 2

Bonn zoological Bulletin - Supplementum 59; 1-116

above). However, Blasius Merrem did attempt, in his frag-
mentary outline of ornithology in 1788, to unite both re-
search traditions. There were field ornithologists who al-
so occupied themselves with problems of systematics,
(e.g.) J.E. Frisch in the text to his large folio work of
1733-1763, J.M. Bechstein, who introduced many scien-
tific names into the literature, and in the 19th century C.E.
Brehm, the discoverer of several European sibling species
(Firecrest/Goldcrest, MarshAVillow Tit, Eurasian/Short-
toed Treecreeper, Crested/Thekla Eark), who thought
much about intraspecific variation. But such connections
were insufficient to effect an integration of the two branch-
es of ornithology, which remained widely separated un-
til into the 20th century (Haffer 2007a, 2008).

In the following section we will look at the species
knowledge of the field ornithologists J.F.A. von Pernau,
H.F. von Gochhausen, J.E. Frisch, and J.H. Zorn, as well
as of the systematist J.T. Klein. It must be borne in mind
that the field ornithologists mentioned here dealt with the
local fauna of their respective native regions, and de-
scribed the habits of only those species that they had per-
sonally observed, while Klein’s ambition was to compile
from the literature a list of the bird species of the world,
without himself having had much field experience. He had
no personal knowledge of most of the species on his list.

The first ethologist - Ferdinand Adam von Pernan
( 1702 , 1720 ). Von Pernau lived from 1660 to 1731. He
did not wish to create a complete avifauna of southern Ger-
many with his books, but instead only to report on his per-
sonal observations of those species that he saw and stud-
ied in the surroundings of his home area of Coburg
(Franken). What are mainly absent are the songbirds and
some game-birds, and he mentioned only a few water and
wetland species in his writings. Pernau too saw manifes-
tations of God’s work in nature’s creatures. He developed
individualistic and astonishingly modern ornithological
hypotheses and arrived at his own independent judge-
ments.

J.F.A. von Pernau was a senior civil servant at the roy-
al court of Sachsen-Coburg, but when his government
work allowed him some free time he used it for intensive
biological studies of birds. His intention was to stimulate
admiration for God’s creatures and to help the reader on-
to “the path to redemption out of frivolous pleasure-seek-
ing and dishonorable gluttony” (Stresemann 1951: 289).
There is no known portrait of this important figure. Per-
nau’s book of 1702 had the lengthy title Unterricht, Was
mit dem lieblichen Geschopff, denen Vogeln, auch auss-
er dem Fang, nur durch Ergriindung deren Eigenschafften
und Zahmmachung oder anderer Abrichtung man sich vor
Lust und Zeitvertreib machen konne [Lesson on what can
be done, for pleasure and amusement, with those delight-
ful creatures, birds, in addition to catching them, only by
inquiring into their qualities and taming or otherwise

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25

training them] (Fig. 13). A follow-up volume appeared in
1720 under the title Angenehme Landliist, deren man in
Staedten und auf dem Lande, ohne sonderliche Kosten, un-
schuldig geniessen kann [Agi^eeable country pleasures that
may be innocently enjoyed in town and country, without
exceptional expense]. These books greatly influenced or-
nithology in the 1 8th century, and most hunters, fowlers
and bird-keepers were familiar with them, which was less
the case among naturalists. They were published anony-
mously, reaching ten editions between 1702 and 1797.
However, the name of the author was known to several
of his writing contemporaries, including Wetzel, von
Gochhausen (see his entry on Chaffinch below), Wirsing,
and von DieBkau (see Schlenker 1982a). Later, Strese-
mann (1925) was the first to rediscover Pernau’s writings
and stress their importance in the history of ornithology
(see also Stresemann 1947, 1951, 1962). Pernau differen-
tiated bird species based on the manner in which they feed,
their preferred habitat, the degree of their sedentariness
(non-migratory, partially migratory, migratory), their
choice of nest site, their sociability during and following
the breeding season, their manner of bathing, or the way
in which they feed their young (from the crop or from the
bill), and it was Pernau who made the first explicit com-
ments on song learning in finches. He was both a bird-
keeper and field naturalist, discussing a few songbird
species, notably the Chaffinch. He speculated about the

SBog roit t)cmlieWiclien©efi5ot>fF/

ixnm

dug auflect>m|!|ang/

gtttr {»urcg v»w jErgrunhmg^rm

Sl6rt|ifung/

SDlan f?c& set £u!l un6 3cit'95{ttrei6 raa-

fpnnet

gtpfUr;

gii./gennWnSP. . . , . /

Anno 1703 .

Fig. 13. Title page of von Pernau (1702; from 1982 reprint).
Bonn zoological Bulletin - Supplementum 59: 1-116

function of territory as an exclusive source of food, sug-
gesting that the Nightingale “is forced for the sake of her
feeding requirements to chase her own equals away, for
if many were to stay together they could not possibly find
enough worms and would inevitably starve” (Fig. 14).

Pernau recognized territorial behavior in birds and its
biological significance, and postulated that a migratory
bird is not driven to depart by hunger or cold but “is driv-
en at the right time by a secret impulse which it obeys, in
contrast to humans, who often resist such impulses” (Stre-
semann 1947, 1951). Pernau used behavioral traits as an
aid to taxonomy (e.g., tail wagging in both wagtails and
pipits) and noted that unmated male songbirds sing more
than mated ones. He also interpreted the biological impor-
tance of bird calls, (e.g.) that the rain-call of the
Chaffinch has a warning, territorial-defense function. His
statements on how a bird learns its own song were con-
firmed by later workers. His ideas on the function of song,
how much is learned and how much is innate, the inher-
itance of “traditional” song dialects and their patterns, and
on imprinting were all of fundamental significance
(Thielcke 1988). Pernau’s books indicated the future di-
rection and aims of avian biological research; he and Em-
peror Friedrich II of Hohenstaufen were really the
founders of scientifically conducted biological field stud-
ies of birds. In the originality of his work he can be com-
pared with the founders of ethology Altum and Heinroth
in Germany, and Selous and Howard in England (Strese-
mann 1925: 612, 621; 1947, 1951).

J.M. Bechstein had also recognized the value of Pemau’s
contribution, editing the ninth and tenth editions of his
book. Bechstein wrote in his foreword that “there has not
been a bird book until now, excepting Zorn’s "Petinothe-
ologiej in which the practical natural history of these an-
imals has been presented more completely and with many
interesting observations than this one”. He was very sur-
prised, therefore, to find it so little referred to in writings
on ornithological systematics (see the anonymous review
of this edition in Allgemeine Literaturzeitung Vol. 1, no.
66, pp. 524-525, February 1799).

Oberjdgermeister Hermann Friedrich von Gochhausen
(1710, 1732). Von Gochhausen was baptized on 5 March
1663 and died in Weimar as sdchsisch-weimarischer
Oberjdgermeister in 1733 (Schlenker 1994). This natural-
ist was especially interested in the avian world and pub-
lished his findings on 1 04 species in a book with the im-
pressive title:

Notabilia venatoris,

Oder Jagd- und Weidwercks-Anmerckungen [...] aufge-
zeichnet von einem die Jdgerei liebenden Waidemann in
Weimar, welcher gerne in Waldern horete friihe der Vo-
gel Gesdnge [. . .] [Hunting and sporting remarks set down
by a lover of the noble sport in Weimar, whose pleasure

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Jurgen Haffer (t) et al.

Fig. 14. Frontispiece of von Pernau (1702; from 1982 reprint).

On Breeding Birds

A selection of the interesting comments by the author fol-
lows below, mostly in summarized form but some as
unabridged quotations. For each species Gochhausen gives
a short description of plumage, but that is omitted here.
Species order is as given by the author and therefore dif-
fers from that in online Appendix 2.

Auer-Hahn [Capercaillie]'. Eats the buds of beech trees
in winter; treads the female after display. She raises the
young without the help of the male. Also eats berries in
summer (pp. 70-74)

Trappen [Great Bustard]'. “In the summer this bird eats
the green unripe grain of cereals, later in the year only the
ripe grain, and in winter the seeds of cereals and root-
crops. What is notable is that if it has the opportunity, it
will also take larks and other small birds, or newly-hatched
chicks of domestic hens or quails, causing much harm.

The Great Bustard lays its eggs often in fields of oats
in spring, selecting those areas furthest from tracks and
paths, where it digs a depression in the bare earth to lay
its eggs in, never more than two. These are yellowish white
with a size between those of Turkey and goose eggs. What
is remarkable is that although the Great Bustard is a timid
bird it will sit fast on the eggs until forced from the nest.

it is to listen to the song of the woodland birds at dawn].

This work appeared for the first time in 1710 (Fig. 15) and
subsequently in ten further editions; we have made use of
that of 1732. For most species, especially the game-birds,
the huntsman gives details on breeding, food, and migra-
tion, as well as incubation period and clutch size, these
last not always being correct, most likely because he oc-
casionally relied on hearsay. Nevertheless, Gochhausen
made a considerable contribution to the dissemination of
ornithological knowledge in Germany (Stresemann 1925,
1926b). Of particular interest is his information on the
breeding of the Little Bustard in Sachsen- Weimar (see un-
der Brach- Vogel).

In his book, Gochhausen deals firstly with the mammals,
followed by “On breeding birds” (pp. 70-163), divided
into landbirds, waterbirds, and birds of prey. The next
chapter is on trees (pp. 164-251) and the last, “On hunt-
ing”, contains details of the equipment and authority of
the hunter plus remarks concerning the “properly designed
deer park” (pp. 279-2282). Several appendices deal with
instructions for hunting and forestry workers, lark-trap-
ping with nets, and with the known fish species in the re-
gions belonging to Saxony.

notabiliA

VENATORIS.

Hiiirdoojieiii

, lie cs icif(jero6e!)I)er^66li(f;cn

lijfgtltlfiil gcf^alfcil/ praclicabd obfC

imfrjd[c 3 bcJ flCilflltfi/

fhfileOukiKiitijifitiict iiii&
jjrtPiffi’n rvsrh'ii/

bKlcrlftj l)frcr SSf(j6iee Dili unisiuifbfr

III Slvilt uniim m finl'fti / Mt btitcl&cii

pclijfn unc ^11 ijfbriiud'cn/aiiijh nriCifreuiittTl'cbirtjiiriir/ riiifm

iu tEilTfn/ lUHfyiijc ejditji/
riid^c eKHigxT

TOO own*

SJeldfier geritc in SQdl&frn

Olbreft ®i6e Kr Mfiiicl I'eiiidflf.

iinll>m bn) .(fjrl ‘Jlnimhafan. 1710.

A* Alt

Fig. 15. Title page of von Gochhausen (1710).

Bonn zoological Bulletin - Supplementum 59: 1-116

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27

whereupon it does not leave the eggs in the same place
but carries them away (probably under its wings or with
its sinuous neck) to a distance of around 100 paces where
they are incubated again, as long as the bird is not further
disturbed, for a total of four weeks [= incubation period
21-26 (-28) days], then leads the young as soon as they
can walk to safety in the cereal crop. Otherwise it is con-
spicuous as a bird of autumn, since it remains here when
other wild birds leave, to return in the spring. The Great
Bustard moves around seeking the warmer fields, feed-
ing in winter time on the vegetation mentioned above.”

The courtship display of the Great Bustard “takes place
in Lent, and if the male has not just one hen but several,
like the Capercaillie, he spreads his feathers and tail like
a Turkey, but unlike the Capercaillie he makes no sound
during this display. At this time it is one of the most bad-
tempered of birds, often kicking and striking each other.”

“ when the male is old he has a finger-long beard of

two or three long, thin white feathers, which, when he is
angry, is spread out on each side of his head to give him-
self an impressive appearance” (pp. 74-77).

Birck-Wildpret [Black Grouse]: Displays on trees as well
as on the ground; raises his feathers like a Turkey and
jumps up from the ground. The hen is mostly nearby. Nest
is on the ground, 8-12 eggs. Food in winter is birch buds
and shoots, in summer berries and plants. Males and fe-
males are very differently colored (pp. 77-80).

Hasel-Hun [Hazel Grouse]: In old conifer forest where
hazel trees grow; eats catkins, buds, later berries and
plants. Male and female are similarly colored, but the for-
mer is larger with a black throat. Courtship takes place at
Lent; the display call is a whistle. Nests on the ground,
and incubates the eggs for three weeks [correct is 22-25
days], 6-8 young, stays in the breeding area during win-
ter and does not migrate (pp. 80-82).

Rebhun [ Grey Partridge]: Incubates eggs for three weeks
[correct is 24-25 days], has 16-18 young that are raised
without any help from the male; food is seeds and fruits,
in winter green seeds mixed with gravel and sand (pp.
82-83).

Wachtel [Common Quail]: The 7-10 young are reared by
the female only (p. 83).

Waldschnepfe [Woodcock]: “this bird can open and close
just the end [...] , about a thumb’s breadth, of its bill
(which is as long as a finger) just like a pair of pliers,
which is made possible by certain nerves in the bill sup-
plied by nature which no other bird possesses, and this can
be demonstrated on a dead Woodcock if the head is
squeezed”; 3-A young; migrates at night (pp. 84-85).

Bonn zoological Bulletin - Supplementum 59: 1-116

Brach-Vdgel [fallow -I and birds]: The most important fau-
nistic record made by Gochhausen is of the breeding of
Little Bustard (Tetrax tetrax) in Sachsen- Weimar (Kunz
1902; Stresemann 1926a: 691). Rewrites: “There are three
fallow-land birds. Firstly, the one called Keilhacken or
Fasten-Schlier [= Little Bustard], a bird around the same
size as a sickly (?) \yerkuhlet\ Turkey, and colored like a
Great Bustard, with long feet so that they run as fast as
an arrow, and sometimes they press themselves close to
the ground and are quite hidden. The proportion and form
of the bill and head are very like a Great Bustard and they
also fly as fast as an arrow, and make a high whistle call.
They migrate through our country in autumn and spring,
but do breed very rarely here, where they have been found
on fallow fields and between stones. They have two
young, and can only be obtained using a shotgun. They
feed on the smallest animals, especially earthworms.”

Secondly, the Saat-Hun [= European Golden Plover],
as big as a pigeon, with speckled greenish feathers, in
flocks on newly sown cereal fields in the autumn; passage
migrant.

Thirdly, the Ditgen [= Dotterel], smaller and with a
whiter belly than the Golden Plover, and also an autumn
passage migrant on newly sown fields, (pp 85-86).

Kibitz [Northern Lapwing]: Appears first in the spring; has
a bill like a pigeon; attacks people in the breeding season;
has a feather crest like a heron and eggs that are good for
eating; 3)-A young. Incubation period 14 days [correct is
27 days]; feeds on larvae, worms, etc. (p. 87).

Pingeltaube [Wood Pigeon]: Timid; nest made of few ma-
terials on oak or spruce tree and never has more than two
young (pp. 87-88).

Hohltaube [Stock Dove]: Has no white neck-ring, rather
bluish in color, nests in holes in trees (p. 88).

Turteltaube [Turtle Dove]: The smallest dove, with a white
band at the end of its tail; breeds in trees, male and fe-
male together (p. 89).

Tages-Schlaffe [Night]ar]: Short bill, at the head very
broad; shaped like a swift; active at twilight and lays 4
speckled eggs [error; never lays more than 2]; leaves us
to migrate in autumn (p.90).

This species was not included in the first edition of 1710.

Krammets- or Krannebet-Vogel [Fieldfare]: Feeds on ju-
niper berries; winter visitor, breeds in the east. Often ap-
pears with the Redwing and is caught in large numbers in
snares (p. 91).

Schnerr [Mistle Thrush]: Breeds in oaks and other trees
(like Blackbird and Song Thrush) and has 3-A young,

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Jurgen Haffer (t) et al.

feeds on mistle and juniper berries. Male and female very
similar in coloration (p. 92).

Meer-Amfiel [Ring Ouzel]'. “This species of bird does not
nest in our region and passes through in autumn and spring
like other birds. It is a black-gray bird, a little speckled,
and slightly larger than the Blackbird, with a white throat,
almost like a Dipper. It migrates on powerful wings, but
is a very stupid bird on the trapping ground, according to
fowlers; when it is caught by stalking [. . .] it allows itself
to be caught again” (pp. 92-93).

Stein-Amfiel [Rock Thrush]'. Lives like Northern Wheater
in rock clefts, cliffs, and vineyards but is as big as a Gold-
en Oriole, “ash-colored from the head to the rump, the bel-
ly brick-red, speckled slightly with white, tail red, and
chestnut brown on back and wings with the feathers slight-
ly dusted white”, feet brown as are bill and eye. Feeds on
worms, etc. and grapes; breeding similar to Wheatear and
departs in autumn. This species “is little noted in our re-
gion” hence is very rare (pp. 93-94).

Zipp-Drossel [Song Thrush]'. Common breeding bird on
trees, incubation period 14 days, 4-5 young; migratory
bird (pp. 94-95).

Schwartz-Amfiel [Blackbird]'. The male is blacker than the
female and with a wax-yellow bill and yellow eye-ring;
incubation period 14 days, 4-5 young. Some of the Black-
birds remain in winter (p. 95).

Wein-Drofiel [Redwing]'. Has red under the wings; feeds
on berries, snails, larvae, and worms; passage bird (p. 96).

Staar [Common Starling]'. Breeds in holes; migratory bird
(pp. 96-97).

Weyhrauch- or Kirsch-Vogel [Golden Oriole]'. Egg-yolk-
yellow with black wings; returns in late spring. Feeds on
cherries and worms, etc. “Forms its nest with wool and
bast [= fibers from the phloem of plants] gathered togeth-
er, on the fork of a branch, bound to the legs of the fork
with long strips of bast from lime trees and so cunningly
constructed that it hangs like a long pouch or like a church
collection bag, so that it is a marvel to see.” Incubation
period 14 days [correct is 15-16 days], 3>-A young (pp.
97-98).

Grienitz or Grienitz-Vogel [Common Crossbill]'. Breeds
in the winter (January-February) in conifer forests, where
the nest is in high spruce trees, 4-5 young. In their first
year they are gray and slightly greenish, in the second year
reddish, in the third yellow-green. Some have the upper
mandible crossed to the left, others crossed to the right.
People talk of larger and smaller Crossbills but that is not

true. They arrive when the conifer cone crop has been
good. “Regarding the breeding of Crossbills, the reason
why the young are hatched at the coldest time of year
could be that they do not arrive here only in the spring
but also in the autumn, and indeed come then in greater
numbers, most arriving when the fir cone crop is a good
one in our region because fir seeds are its favorite food.
And when these oil-rich seeds are sufficient to last into
the winter then the bird eats so many it puts him into the
mood for mating” (pp. 98-99).

Seiden-Schwantz [Bohemian Waxwing]'. Appears regular-
ly but is not a breeding bird; its food is Juniper berries (pp.

99- 100).

Kern Beisser [Hawfinch]'. Bill thick, breeding bird, 4
young, many Hawfinches spend the winter here (pp.

100 - 101 ).

Gumpel, Blut-Finck, Thum-Pfaffe or Rothschlegel
[Bullfinch]'. Eats rowan berries, 4 young, “which can eas-
ily learn to whistle whatever you want them to” (pp.

101 - 102 ).

Lerche [Skylark]'. Common; usually breeds twice in sum-
mer (pp. 102-103).

Heide-Lerche [Wood Lark]'. Tail shorter than in the last
species, breeds in clearings in the forest, 4-5 young (pp.
103-104).

Mantel-Krdhe or Blarack [European Roller]'. Blue, back
brownish, rare passage bird, does not breed here (p. 104).

Schwartz-Specht or Hohl-Krdhe [Black Woodpecker]'.
Breeds in holes it makes itself, ?)-A young, remains in win-
ter. Tail has stiff supporting feathers; 2 toes pointing for-
wards and 2 backwards (pp. 105-106).

Grun-Specht [Green Woodpecker]'. Stays here in winter
also (p. 106).

Bund- or Roth-Specht [Great, Middle(?), and Lesser Spot-
ted Woodpeckers] '. “There are three different kinds of three
different sizes”, black-and-white checkered, red on the
head and under the tail.

Perhaps alongside Great Spotted and Eesser Spotted,
Gochhausen here also distinguished Middle Spotted
Woodpecker (p. 107).

Blauspecht [Eurasian Nuthatch]'. Breeds in holes in trees
and remains here in winter (p. 107).

Baum-Reiter [treecreeper]'. With thin bill, curved at the
tip, breeds in hollow trees (p. 108).

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Wiedehopff [Hoopoe]-. Forages for its food like a snipe,
departs already in August and is the last to return (pp.
108-109).

Nachtigall [Common Nightingale]'. Breeds in thorny
hedges (pp. 109-110).

Wendehalfi [Wryneck]'. Migratory bird, its call is similar
to the Common Kestrel; breeds in hollow trees. It has a
long tongue with little barbed hooks like a woodpecker
(pp. 110-111). This species was not included in the first
edition of 1710.

Roth-Kehle {Robin\. A well-known common bird (pp.
111 - 112 ).

Roth-Schwantz [Common Redstart]'. Breeds in old trees
(pp. 112-113).

Fliegen-Schnepper [one of the Sylvia species]'. A little
gray bird, creeps around in hedges and bushes; breeds in
bushes or hedges (p. 113).

Grafi-Mucke [Whinchat]'. “This little bird is as big as a
Robin, but has a short tail and is speckled gray in color
with a yellowish belly, almost always found on meadows
where it searches for food in the grass and in bushes, un-
der which it lays its eggs and raises its young. Has usu-
ally 4 to 5 young, but what is remarkable is that, where
other birds have differently speckled eggs, this bird has
pure sl<y-blue eggs. Eats only flies, larvae, and worms and
because it is the first to leave on migration it returns very
late to the same place since it cannot stand the cold” (pp.
112-113).

Stein-Klatsche [Northern Wheatear]'. Related to the
Whinchat and with a similar shape, whiter on the belly,
but the Wheatear prefers bare barren land and stony places
to meadows; breeds in holes and lays 4 white slightly
speckled eggs, feeds on worms and larvae and is migra-
tory (pp. 114-115).

Finck [Chaffinch]'. “Herr Bernauer [=A. von Pernau] es-
pecially explains in his description how this bird can be
trained to utter a great variety of different songs, and he
himself has observed how this variety is produced” (pp.
115-117).

Buch-Finck, Quecker or Zdhrling [Br ambling]'. Colors
more attractive than in Chaffinch; winter visitor (p. 117).

Stieglitz and Hdnffling [Goldfinch and Linnet] (pp.
117-119).

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Schwanschel or Grilnling [Greenfinch]'. Food is seeds
(hemp seeds) (p. 119).

Zeifiig [Siskin]'. The nest is in high conifer trees; found
here in summer and winter (pp. 119-120).

Zitscherling [Common Redpoll]'. Winter visitor (p. 120).

Meifien [tits]'. Great Tit [PickmeiBe] is the biggest; oth-
ers are Crested Tit [Kupp-MeiBe]; Blue Tit; Coal Tit
[Schwartz-MeiBe]; Long-tailed Tit (pp. 121-122).

Bachstelzen [wagtails]'. (1) Gray and white, throat black
[= White Wagtail] and (2) underparts yellow, pale gray,
back mixed with yellow [= Yellow Wagtail] (pp. 122-123).

Aemmerling [Yellowhammer]'. Common (pp. 123-124).

Holtz-Muschel or Wilder Sperling [Tree Sparrow]'. Breeds
in hollow willows and remains in winter (pp. 124-125).

Zaunkonig [Wren]'. Sings loudly and builds a large nest
of moss with a small entrance (pp. 125-126).

Weiden-Zeisig [Chiffchaff] and Gold-Hahnichen [Gold-
crest/Firecrest] (p. 126).

Waterbirds

Schwaane [Mute Swan]'. Incubation period 4 weeks [cor-
rect is 35-38 days] (pp. 130-131).

Wilde Ganfi [Greylag Goose]'. Incubation period 4 weeks
[this is correct; 27-29 days] (p. 131).

Fisch-Reiher. The well-known gray-colored herons that
nest on high trees; in addition (1) large heron, light gray
on the back, white on the body speckled with black streaks,
bill long and red, feet yellow, with 2-3 long feathers on
the head [= Grey Heron], and (2) smaller heron, dark gray
on the back, black on the head, bill black, feet short and
blue; 2-3 long feathers on the head [= Night Heron] (pp.

132- 133).

Storch [White Stork]'. Widespread in the villages (pp.

133- 134).

Schwartzstorch [Black Stork]'. Bill and legs reddish; lives
in forests; incubation period 3 weeks [correct is 30 days]
(p. 134).

See-Rachen [Goosander]'. White, head black (p. 134).

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Jurgen Haffer (t) et al.

Gemeine grosse wilde Ente [Mallard]'. Breeds on willow
trees, even in Crow or Magpie nests near lakes or rivers,
“to which they later take their fledged young (which takes
3 weeks) from these willows or other trees, probably car-
rying them carefully in their bill, so that they are in safe-
ty”. Migratory, but many stay here in the winter (p. 135).

Schmal-Endte'. Similar to the large duck (above) but small-
er (p. 136).

Pfeiff- or Speck-Endte [Eurasian Wigeon]'. Head brown,
belly white, wings brown with white patches (pp.
136-137).

Horbel or Bldfilein [Coot] (pp. 137-138).

Taucher or Wasser-Huhn [Great Crested Grebe] (p. 139).

Ried-Schnepffe [Great Snipe]'. “This bird is as big as a
Woodcock and is as great a delicacy, and has also the same
color and form, but in flight it is much faster. It migrates
like other birds after it has raised its 4 to 5 young here in
the largest swamps and marshes, and nowhere else. Its
food is also the same as the Woodcock, namely the white
tender roots of plants they find in the marshes” (pp.
139-140).

Haarschnepffe [Jack Snipe]'. “This is rather smaller than
the previous bird, but however in color and nature is just
the same” (p. 140).

Pfuhlschnepffe or Himmelsziege [Common Snipe]'. “There
is no difference in color to the previous but one bird, on-
ly that species is larger and this one is different in that,
when it is disturbed, it climbs up high into the air then,
when it flies in an arc back towards the ground, it makes
such a loud quivering sound, which they say is made by
the wings, that one is amazed by it. When it sits on the
ground, by water and in marshes, it often makes a call
when the weather changes that has two different notes. It
breeds in swamps and on hillsides, without nesting ma-
terial, laying four eggs which are incubated in 14 days”
[correct is ca. 19 days] (pp. 140-141).

Sandlauffer or Griefihun [Little Ringed Plover]'. Speck-
led black with white breast; breeds on sandbanks; ring-
ing call (p. 141).

See-Schwalbe [terns] '. “There are two of these though they
are the same color, silvery white, but one of them is larg-
er than the other. The larger one swims now and then on
the water like a duck, which the smaller one never does
but instead flies up and down the whole day long like a
swallow, over the lakes and ponds, looking for its food,
which is small water animals. They breed in our region

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in reedbeds and have 3 to 4 young, but migrate like oth-
er birds” (p. 142).

Eifi-Vogel and Wasser-Amfiel [Common Kingfisher and
Dipper]'. The food of both is worms and snails and both
stay here during winter where there is open water (pp.
142-143).

Rohr-Dommel [Eurasian Bittern]'. Calls at twilight and at
night, and can be heard a long way away (pp. 144-145).

Rohr-Sperling [Reed Bunting]'. Breeds in reedbeds,
moves away in autumn (p. 145).

Birds of Prey

Males are generally smaller than females, many regurgi-
tate pellets. They never drink and survive on the liquid in
their food.

Steinadler [Golden Eagle]'. Takes roe deer and hares; nests
in the forest and has 2 young (pp. 146-147).

Schuhu [Eagle Owl]'. Nests on cliffs, takes hares. Remains
here in winter (pp. 147-148).

Blau-Fufi [Peregrine Falcon]'. “Although the Peregrine
Falcon is not much larger than the Goshawk, it has a great
advantage in its talons and bill since with these it easily
catches hares, ducks, and partridges. Unlike other birds
of prey, it does not seize them at the first strike but knocks
them to the ground with its feet, and only then does it grasp
them with its talons. It nests in our region in forests as well
as on old ruined towers and walls. Because of its skillful
way of killing it is much sought after by falconers, who
use it to hunt hares and ducks, and they catch them in nets
and by using a leather ‘saddle’ attached to pigeons. It
moves away in the autumn” (pp. 148-149).

Habicht [Goshawk, male]'. They leave in autumn and re-
turn early in spring. In the forest they look for their old
nests, where they are least disturbed, 2-A young, “which
hatch after 14 days” [correct is 35-38 days]. They are used
by men to catch partridges, quail, hares, and herons (pp.
149-150).

Sperber [Sparrowhawk, female]'. Slightly smaller than the
Goshawk; is used by men to hunt partridges, quail, and
songbirds (p. 150).

Baum-Falcke [Hobby]'. “Although there is no difference
in size, nest, or migration between the Sparrowhawk and
this falcon, there is a difference in color for this bird is
bluish on the back, yellowish on the throat and black in

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between. The young leave the nest very late, around St.
James’s Day [25 July]; they can be very useful in catch-
ing larks. [. . . ] This falcon has the habit that, when it sees
a hunter or anyone with dogs in a field, it stays close to
them and flies above them, so that if a lark or so is dis-
turbed and flies up then it can pursue if’ (pp. 150-1151).

Schmerl [Sparrowhawk, male]'. Nests rarely here. “This
is a small, delicate, barred-breasted bird, yellow feet, as
large as a Fieldfare” (pp. 151-152).

Rittel-Geyr [Common Kestrel]'. Its food is mice, voles and
young birds, nests on church towers; migratory bird (p.
152).

Horneule [Long-eared Owl] and Steineule [Barn Owl]'.
The former prefers to nest in hollow trees, the latter in old
buildings; both live on mice and voles and regurgitate pel-
lets; they do not move away in winter (pp. 152-153).

Mulane or Schwalben-Schwantz [Red Kite] (p. 153).

Mduse-Geyer [Common Buzzard]'. Feeds on mice, voles,
frogs, young hares, and also on carrion (pp. 153-154).

Fisch-Geyer [Osprey]'. Their food consists only of fish;
they migrate in autumn (p. 154).

Kautz [Tawny Owl] and kleiner Stein-Kautz [Little Owl]
(pp. 155-156).

Neun-Todter [shrikes]'. There are two species, (1) Wild-
er Kriick-Elster, rather larger and catches young birds
[Great Grey Shrike]', (2) a smaller species which feeds on
beetles and butterflies that it impales on the thorns of
hawthorn and blackthorn trees; nests in thorny bushes
[Red-backed Shrike/Lesser Grey Shrike?] (p. 156).

Kuckuck [Common Cuckoo]'. Feeds on caterpillars and
worms, does not have curved talons and has small feet like
a pigeon; in the autumn does not change itself into a Spar-
rowhawk. “.... Although they mate and produce young
they never lay their eggs in a nest they have made them-
selves, but lay them in the nests of other birds, like Whin-
chat, wagtail and such birds, and then let them feed their
young with worms, etc. and raise them” (pp. 157-158).

Nufi-Heher [Eurasian Jay]'. A common woodland bird (pp.
158-159).

Tannen-Heher [Nutcracker]'. Lives in fir forests, eats fir
and spruce seeds [no remarks are made on breeding oc-
currence] (pp. 159-160).

Bonn zoological Bulletin - Supplementum 59: 1-116

Golck-Rabe [ Common Raven]'. A harmful bird which does
not leave in winter (p. 160).

Kleiner Rabe [Carrion Crow]'. Smaller than the Raven,
nests in woodland and gardens and is shot while on the
nest. “But the crow that is gray on the body [= Hooded
Crow] does not nest in our region but comes here on mi-
gration and stays, since it has the same habits as the oth-
er crow, until spring” (p. 161).

Rucken [Rook]'. Colony breeder; feeds on seeds and lar-
vae, worms, etc. (pp. 161-162).

Dohle [Western Jackdaw]'. Nests in holes in walls; does
not leave in the winter (p. 162).

Aglester [Magpie]'. Cunning and harmful. The nest is
domed and has an entrance at the side (pp. 162-163).

The first book of folio plates of Central European
birds - Johann Leonhard Frisch (1733-1763). The sub-
rector and later rector of the famous grammar (or high)
school “Zum Grauen Kloster” in Berlin was a versatile
scholar, linguist, entomologist and ornithologist, inventor
of the famous “Berliner Blau” (a much-used blue dye), and
proponent of a silk industry based on local mulberry plan-
tations (Fig. 16). As a naturalist, he studied the life of an-
imals “to praise the eternal Creator”. He had gathered a
wealth of experience on long journeys in Austria, Hun-
gary, Italy, Switzerland, and Germany, before settling in
Berlin in 1698 (Schalow 1919). Here he collected data on
insects and the avian world specifically to support his
physico-theological studies. He published the first book
on the insects of Germany, Beschreybung von allerley In-
secten in Deutschland [Description of all kinds of insects
in Germany], in 1720-1738, and the first German bird
book in 1733-1763, Vorstellung der Vogel Deutschlandes
und beyldufig einiger Fremden [Presentation of the birds
of Germany, with some from other countries]'. Fig. 17.

The laher is a large folio work with 307 excellent col-
ored illustrations of birds, as far as possible life-sized, on
256 copperplates, which were mostly engraved by
Frisch’s son Ferdinand Helfreich, 28 in the final year al-
so by his grandson Johann Christoph, and hand-colored
by F.H. Frisch and his wife. The natural colors lend the
birds a very lifelike appearance. Where it was necessary
and possible, male and female of a species are illustrated
together, often one across the other. These plates became
models for later bird artists because of their excellent col-
or reproduction and positioning of the subjects. The orig-
inal drawings for the engravings are today in the Stadt-
bibliothek Mainz (Schlenker 2004, 2005); Figs 18a, b.

Schalow (1919) and Stresemann (1941b) gave biblio-
graphical details on the publication dates of the various
installments over 30 years, but neither informed modern

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Jurgen Haffer (t) et al.

Fig. 16. Portrait of Johann Leonhard Frisch (from Schalow
1919).

ornithologists about the book’s biological and historical
significance. Schalow (1919: 522) wrote, without further
explanation, “As far as the biological information set out
in Frisch’s work is concerned, I will not go into details
here. It is based on Frisch’s observations of birds in the
field as well as of his own captive birds”. Stresemann
(1941b: 4) said, “regarding the scientific importance [of
this work], I intend to return to the matter at a later date”,
which he never did.

Bastardization. Frisch was aware of the biological
species criterion (successful sexual reproduction) intro-
duced by John Ray, and already knew that certain close-
ly-related species that would never hybridize in the wild
could do so in captivity, but that the resulting crosses were
sterile. He made the following interesting observations on
hybrids and hybridization, under the title “On the Canary”
(1733, end of Class I):

“Firstly: All bastards are from those species of animals
that are closely related in many of their characters. There-
fore Goldfinch and Canary, and all other birds of this first
Class that produce bastards, must be closely related to each
other. It is simply not possible with others; the Nightin-

Bonn zoological Bulletin - Supplementum 59; 1-116

gale male can be as willing as he wishes, and the Canary
female mate with many different birds, one cannot pair
them or get them to mate because they are not very close-
ly related. So donkey and horse, or dog and wolf, can have
young together, but not horse and deer, or dog and bear.

Secondly: All bastards are born to animals that have
been forced to come together, that in the freedom of na-
ture would never mate with each other. [. . . ] A bird that
cannot find another of the same species in one place flies
to another part of the world to find one there.

Thirdly: All bastards resemble their fathers in the head
and tail. The mule has its head and tail like a donkey, the

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Fig. 17. Title page and frontispiece of Frisch (1733-1763),
with an engraving of a memorial bust of J.L. Frisch and portrait
of his son Ferdinand Helfreich Frisch by the naturalist’s grand-
son Johann Christoph Frisch, from a design by B. Rode.

rest of its body like a horse. The bastard of a Goldfinch
father shows the head and tail of its father and the body
of its mother, the Canary.

Fourthly: All bastards are sterile. As is well known, the
mule cannot reproduce, and this is also the case in birds.
No young can be got from a bastard. The main reason is
that the blessing at the Creation spoken over the animals.
Be fruitful and multiply, does not apply to them. They have
from their father and mother the blessing of life but not
of multiplication. The inventions and restrictions of Man
can create a third thing from two by mixing and the ef-
fect of nature, but it is not a new creature that can repro-
duce, only a temporary, deficient thing.

Fifthly: A special circumstance that is to be noted when
breeding birds is how far the blessings of Creation extend.
Birds are created to incubate their own eggs, an indispen-
sable part of reproduction. If an alien bird incubates an
egg then a half-bastard will hatch from it; that is, the bird
that emerges is like the parents but will never itself repro-
duce. They can lay eggs, good fertile eggs, but do not in-

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cubate them. At the time when the eggs must be incubat-
ed they are incapable of doing so, and only because the
eggs from which they hatched had been incubated by an
alien bird. Many householders have experienced to their
loss that ducks hatched from duck eggs incubated by a
chicken are of no use for breeding. A certain gentleman
wanted to have many pheasants on his land and his
‘clever’ managers had the pheasant eggs he had bought
incubated by chickens; the pheasants that hatched did not
procreate.”

The idea of the “half-bastard”, which seems so pecu-
liar to us, was later picked up by J.H. Zorn as a partial
explanation for the non- incubating Cuckoo (see below, and
Haffer 2006: 52).

Frisch also observed that snipe are able to open only the
tip of their long bill, while most of the bill itself remains
closed. Kingfishers, which breed on the steep banks of the
Oder river east of Berlin and which he kept in his aviaries,
regurgitated “pills” composed of undigested fish bones and
scales. By tying threads dyed with red watercolor to the
feet of swallows he proved that they do not hibernate in
the mud under water, as was then widely believed. When
the birds returned next spring the threads had not lost their

color. He concluded, “Most probably they fly to a coun-
try where they find food in wintertime”.

Frisch divided the birds into 12 Classes:

“F Small birds with short thick bills, for splitting hemp-
seeds

IF Small birds with thin bills, for eating flies and worms

III. Thrushes and blackbirds

IV. Woodpeckers and ‘tree hackers’

V. ‘Jays’ and magpies

VI. Ravens, crows, and jackdaws

VII. Diurnal birds of prey

VIII. Owls and night-birds

IX. Wild and domestic game-birds

X. Wild and domestic pigeons

XI. Wild and domestic geese and ducks and other swim-
ming birds

XII. Birds that like to be close to water or swampy places.”

Frisch’s species names are listed in online Appendix 2. He
had a good overview of the German avifauna, though un-

Sbr- r'fhrtf/d

Figs 18a, b. Two plates from Frisch (1733-1763). 18a: male and female Golden Oriole; 18b: Common Redstart and European
Robin (top), adult and first-year male Bluethroat.

Bonn zoological Bulletin - Supplementum 59: 1-116

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Jurgen Haffer (t) et al.

derstandably several reed warblers, Sylvia warblers, leaf
warblers, geese, ducks, raptors, gulls, and waders were
missing. In the following, we comment on several of
Frisch’s statements on a selection of species, and on some
of the birds illustrated on his color plates (cf also Haffer
2006: 49-51). We follow the numeration of his plates (see
also the review of the birds of J.L. Frisch in Bechstein’s
Handbuch [Vogel 2, pp. 583-741, 1793] and in Schalow
1919: 51-61). Frisch also illustrated some exotic birds,
which were in cages (parrots) or in the farmyard (Turkey,
pheasants), as well as various varieties of the Canary and
domestic pigeons and chickens, which we do not discuss
here.

Dompfaff or Blutflnk [Bullfinch], Plate 2: “Three differ-
ent species are found of different sizes. Whether this dif-
ference is the same everywhere, or varies from place to
place depending on better or poorer feeding for the birds,
must be further investigated.” Frisch was probably famil-
iar with the large race of the Bullfinch that appears in Ger-
many as a winter visitor from northeastern Europe. On the
other hand, Bechstein (p. 588) thought that the case here
was one of individual variation and that Bullfinches vary-
ing greatly in size could be found in the same nest.

Gelbkehliger Hdnfiing [Twite], Plate 10, Fig. 1: Frisch
writes of this bird, “This is the third species [of linnet-type
birds] known in our region, but only to the bird-trappers,
who strangle them along with other birds, otherwise it is
unknown. In the Mark Brandenburg it is called 'Quitter' .
They appear in small numbers and are trapped at finch de-
coys; has no song that one can hear when it is in a cage,
and leaves the area again in February.”

Bechstein (1793: 483, 593-594) had no personal expe-
rience of the Twite (“I have never seen it [Arctic Finch,
F. fiavirostris] in Thiiringen; but it has been observed in
winter in Oberhessen”) and believed that the bird illus-
trated here was a two-year-old Linnet, though he added,
“Where this bird gets its yellow bill from I do not know;
because as far as I am aware the Linnet in winter has a
whitish, or whitish-yellow bill, but I have never seen one
that was sulfur-yellow, as it is shown here.”

Kreutzschnabel, Grilnitz [Common Crossbill], Plate 11,
Fig. 2: The text refers to this species, but the pair illus-
trated are surely Parrot Crossbills, as is evident from their
large bills. Bechstein {Handbuch, 2nd edition 1807, p. 21,
footnote) recognized that this was the case, and wrote,“
When one looks at the plate in Frisch, because of their size
and the thickness of their bills, the birds shown there are
certainly Parrot Crossbills. The writer must have been able
to obtain this bird much more easily in the pine forests
than the Common Crossbill. But when one reads the ac-
companying description, it does fit much better to Com-
mon Crossbill, so it has probably been copied from per-

Bonn zoological Bulletin - Supplementum 59; 1-116

sonal or written sources.” In his text Frisch describes in
great detail the feeding behavior of crossbills on conifer
cones and their breeding in winter (see Haffer 2006: 50).
Brachlerche [Tawny Pipit], Plate 15, Fig. 2b: “When it
sings, which it does to attract a female, it sits on a slight-
ly higher post or tree, from which it sometimes flies high
into the air so that it can be heard even farther away, but
it soon returns to a perch again. [. . .] It can be told from
the Sky and Wood Larks by its wagging tail, while the Tree
Pipit [Gereut-Lerche^ is rather thinner and longer. But all
of them are almost identical in color.”

Wiesen-Lerche [Meadow Pipit], Plate 16, Fig. lb: “

climbs high into the air so that one cannot see it any more
. . . There are very few of these larks and many bird-trap-
pers do not know it. ... It feeds in grassy places and mead-
ows, and like the Sky Lark in young crops, and eats worms
and other kinds of creeping vermin ... It wags its tail just
like the Tree Pipit].”

Bechstein, who initially regarded Tawny and Meadow
Pipit as one species, wrote {Handbuch, 2nd edition, 1 805,
p. 302, footnote): “These birds, namely Tawny, Tree, and
Meadow Pipit, have been the source of many errors in nat-
ural history publications. I myself have also been led
astray. Therefore, since they differ markedly from the
larks, I have created a new genus for these birds which I
have cdXXQd Anthus ''

Schwarz-Kehlein [Common Redstart], male Plate 19, Fig.
la; female and Juveniles Plate 20, Figs, la, 2a, b: “This
bird is by some people divided into Common Redstart
[Garten-Rothling] and [Stadt-Rothling] depending on
where it nests and sings. The latter sings very loudly from
houses or roofs. [. . .] The song begins as early as March
so it is very pleasant to hear”. It is possible that Frisch
refers here to the Black Redstart, which colonized Cen-
tral Europe during the 1 8th century from the south.

Blaukehlein [Bluethroat] , Plate 19, Fig. 2a, b; Plate 20,
Fig. lb: Frisch described the female on Plate 20 as aRoth-
schwdnzlein with a half-red, half-black tail, clearly not rec-
ognizing the bird as a Bluethroat.

Two kinds of Nightingale [Common Nightingale and
Thrush Nightingale or Sprosser], Plate 21, Fig. 1: “Two
species have been separated, since one is slightly redder
than the other and is called Rothvogel [‘Redbird’], while
the other, called by some Sprofivogel or Sprosser, is slight-
ly bigger and has less red on the tail. This Sprosser sings
more at night than the other one. In the song itself there
is very little difference between the two. A musician could
perhaps describe the keys of this song, since it is not pos-
sible to even guess at the way it is done from the move-
ments in the neck and throat of these birds. If one could
measure the strength of the sound made by a Nightingale

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one would have to be amazed that such small lungs could
produce so much air.” Further there are detailed instruc-
tions how to keep and feed these birds in a cage.

Monch mit schwarzer und mit rdthlicher Platte [Black-
cap], Plate 23, Fig. la,b: Frisch, just as the later J.H. Zom
(1743), took male and female Blackcap for two different
species.

Weidenzeisig [Chijf chaff] , Plate 24, Fig. 1 : The bird illus-
trated is of this species; Bechstein (1793) thought that it
was a Willow Warbler.

Blaukdpfige rothe Amsel [Rock Thrush], Plate 32, Fig. 2:
“This bird is known only in a few states in Germany. . . .
[This illustration] was sent to me from Dresden by a good
friend, under the name of der grosse Roth-Wustlich. ... It
breeds or nests in the highest buildings in the scaffolding
holes”. Bechstein (1793: 615) commented on this pecu-
liar statement as follows: “What Frisch says about these
birds whose illustration he received from Dresden, that
they nest on the highest buildings in the scaffolding holes,
and that they have a song that is part high, pure whistling
and part very similar to the song of the grofie Grasmiicke,
sounds very much like the Wistling {Motacilla Erithacus
L. [= Black Redstart]), which also resembles these birds
in that it is bluish on the upperparts and has a rust-col-
ored tail.” Clearly only the illustration of this caged bird
came from Dresden, while the bird itself was possibly
trapped in the hills to the south of the city.

Rate [Common Raven], Plate 63: The text refers to the
Common Raven but the bird on the plate is a Carrion
Crow, which Frisch (in eastern Germany) would not know
as a breeding species. Bechstein (1793) too had his sus-
picions, commenting on the illustration, “If one did not
know that this is meant to be a Raven, one would think it
was a Carrion Crow (Corvus Corone Lin.)”.

Grau-weisser Geyer or Falck [Hen Harrier?], Plate 79:
“It is not common in our region, but it is native . . . also
pulls grasshoppers to pieces.” The bird shown on the plate
shows no black patches on the secondaries. Bechstein
(1791 : 256) described the Blauer Habicht (Falco cyaneus
= Hen Harrier) as a good species, but later, in 1793 (while
reviewing Frisch’s birds), he summed it up as being in-
separable from the Halbweye (= Montagu’s Harrier).

Weisser Geyer or Falck [Pallid Harrier?], Plate 80:
“Rather smaller than the previous species, the foot to the
knee not so long. The wing feathers are edged brownish
yellow in the previous species, in this one edged white.
The inner primaries in this species are pale gray, in the
previous one black.”

Rothes Italidnisches Rebhuhn [Red-legged Partridge],
Bonn zoological Bulletin - Supplementum 59: 1-116

Plate 116: But the illustrated species is, given the narrow
flank banding, black bib border, and well-defined breast
band, a Rock Partridge (Alectoris graeca).

Wilde Gans [Greylag Goose], Plate 155: “In reality this
is the well-known and everywhere common wild goose,
also called Grey Goose.” The bird in the illustration is,
however, a Bean Goose, shown by the orange-colored
patch on the upper mandible.

Moppelgans [Red-breasted Goose], Plate 157 SuppL:
“The actual India, or part of that country, in which this
goose is resident or nests, remains unknown to us.”

Nordische schwarze Ente [Velvet Scoter]: Plate 165 Sup-
pi.: “It actually inhabits the coasts of Norway, but often
crosses the Baltic, especially in winter, to the coast of Prus-
sia. The whole body is velvet black with silky soft feath-
ers; the male has a fleshy knob over the base of the bill,
which the female lacks. The stomach is often full of snail
shells.”

Grosse Halb-Ente [= Black-throated Diver/ Arctic Loon],
Plate 185, SuppL: “As big as a goose; it can weigh up to
32 pounds . . . When this bird lays eggs it screams as hor-
ribly as a man in fear of his life calling for help.” Because
of the powerful bill, the species illustrated is not a Black-
throated Diver but a Great Northern Diver/Common Loon.

Bienenfresser [European Bee-eater], Plate 222: The ori-
gin of the bird in the plate is not the Mark Brandenburg
but East Prussia, since J.T. Klein (1760, p. 113) in Danzig
wrote, “I have sent a nice illustration in lifelike colors to
Frisch in Berlin, which he used to produce the bird on
Plate 222, and from which one can best see the beautiful
colors”.

The physico-theologian Johann Heinrich Zorn and his
handbook of ornithology (1742-1743). The Protestant
pastor Zorn (1698-1748) wrote a two-volume handbook
on the general and specific ornithology of southern Ger-
many with the title Petino-Theologie^ (1742, 1743), with
which, within the framework of natural theology, he de-
sired to make 'An attempt to encourage men, through clos-
er observation of birds, in the admiration, love and rev-
erence of their most powerful, wise, and benevolent Cre-
ator, thus the subtitle of his work (see online Appendix
2). The foreword to the second volume contains the guid-
ing principle of physico-theology: “God’s existence and
qualities can be deduced from the nature of every crea-
ture” (p. 5). In the first volume, which deals with gener-
al ornithology, the author discusses the functional anato-
my of birds as well as molt, behavior, reproduction, move-
ments and migration (see Haffer 2006: 1 8-20). In the sec-
ond, in over 250 pages he reviews the ca. 130 species oc-

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curring in the mid-Franken region where he made his ob-
servations. Given the lack of large water bodies there,
many wetland and water species are missing. It was not
Zorn’s intention to compile an exhaustive avifauna of
southern Germany, but as a physico-theologian it was
rather his aim to describe the wonderful adaptations of
birds to “encourage men in the reverence of their Creator”.
For every species Zorn described (in varying detail)
plumage color, nest site, breeding, the rearing of the
young, feeding, behavior, song, occurrence, and migration.
He possessed considerable field experience gained over
many years of observing in the wild. The Petino-Theolo-
gie is not illustrated. Zorn had hoped to publish color
plates in several supplementary installments but died in
1748 before he could complete the work.

He classified the birds as follows:

I. Landbirds

Birds of prey, some of which hunt during the day and some
at night

Ravens, crows, jackdaws, and magpies
‘Jays’

Woodpeckers
Game-birds and larks
Pigeons and doves
Landbirds

Thrushes, blackbirds, and Starling

Seed-eating birds

Worm- and insect-eating birds

IT Waterbirds, some of which live on the water, and are
called swimming birds; some of which live by the water,
or in wet and marshy places, and are called marsh- or bog-
birds

In Volume 2 of his Petino-Theologie (1743), Zorn makes
brief pertinent comments on every species regarding ap-
pearance, behavior, and occurrence. We quote here, in the
order in which the species are listed, selected statements
that are of interest for a variety of reasons:

Gelbe Bachstelze [Grey Wagtail], p. 94: “Likes to build
its nest in water ditches, by watermills, and in old walls.”

Guguck [Common Cuckoo], p. 245: “Since I am of the
opinion that this bird sucks the eggs of other birds and
leaves the empty shells in their nests, I have put it in this
class [birds of prey].” Zorn knew a great deal about the
brood parasitism of the Cuckoo, and postulated that the
young Cuckoo forces the host nestlings out of the nest
(Haffer 2006: 51-53).

Der mittlere Neuntodter [Woodchat Shrike], p. 251 : “The
whole back of its head is colored light brown or brick-red,

Bonn zoological Bulletin - Supplementum 59; 1-116

which continues over the short neck down to the back.
This meets a black mantle bordered reddish; on both sides
are white patches against the black.”

Birkheher [European Roller], p. 268: The bird is now and
then seen here in pairs “which nest in holes in hollow oak
trees and have three, sometimes four young”.

Blau-Specht or Kleiber [Eurasian Nuthatch], p. 274:
“When the hole in which it wants to nest is too wide, it
plasters it with clay so it is then just wide enough for it
to creep in and out.” Stays here in winter.

Feldlerche [Sky Lark], pp. 291-292: “The extended sec-
ondary feathers near the body cover the wings proper, so
keeping them dry when in wet grass, as well as protect-
ing them from wear.”

Gereuth-Lerche [Tree Pipit], p. 297: Pernau “wants this
[species] in the wagtails rather than in the larks, because
it always wags its tail; but the colors, feet, occurrence, the
song, and the way it is executed, all combine to prevent
this bird being considered as anything but a lark.”

Brach- or Koth-Lerche [Tawny Pipit], p. 298: “The au-
thor of Agreeable country pleasures [= Pernau], p. 99,
wants to count this species too among the wagtails and,
to distinguish it from the others, give it the name Field
Wagtail; but the characteristics already mentioned should
suffice to make it clear that the bird belongs to the larks.”

Pip-Lerche [Meadow Pipit], p. 299: “The much praised
Herr Frisch writes of this bird: It breeds in bare places,
especially in meadows where people do not go much, and
makes its nest sometimes under a clod of turf or earth;
from the way it makes its nest it is called the clay bird.”

Amsel [Blackbird], pp. 3 1 8-319: “ It likes to be near wa-
ter, wells, and rivers where it finds food. In spring it has
its young before any other woodland birds, in early
March.”

Gold-Amsel [Golden Oriole], p. 322: “Its nest is hung like
a basket on the fork of narrow branches.”

Gemeiner Fink [Chaffinch], pp. 331-332: The spring
plumage of the male is not created by molt. “The old feath-
ers do not fall out, as stated in Country pleasures [= Per-
nau]; they just change color.” “As well as [grain and seeds]
it eats all kinds of flying and crawling insects, with which
it also feeds its young from the bill.”

F or Stieglitz [ Goldfinch ] and Grunfink [ Greenfinch ] Zom
stresses that the nestlings are fed from the crop.

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Kreuzschnabel [Common Crossbill], p. 346: When the fir
crop is good this visitor breeds in winter, “as Herr Frisch
has also remarked on”.

Goldammer, Emmerling [Yellow hammer], p. 347: The bill
“has something unusual: it is short but pointed towards
the tip, the lower mandible has an indentation on each side,
while the upper has a bulge on each side which fits into
the indentation, so that the bird is able to squeeze the
grains of barley and other cereals it feeds on out of their
husks, and other seeds are peeled.” The nestlings are fed
with “all sorts of larvae and worms and flying vermin”.

Platten-Meise, Nonn-Meise [Marsh Tit], p. 364: “They
hide hemp seeds in the bark of trees, or between the
branches, so they can live off them at a later time.”

Wiedhopf [Hoopoe] , p. 368: “The nestlings stink, which
comes from the excrement that the parents do not remove
from the nest.”

Monch mit schwarzer Platte [Blackcap] , p. 373: “I have
long been in doubt about whether the female of this bird
also has a cap or not; but I had some shot in the spring of
1741 and there were ones with black caps that had testes
and some that had ovaries, and the only difference I could
see was that the female cap was not so black as the male’s;
they are also slimmer than the males, which are fatter and
of a darker color.”

Monch mit der rdthlich- oder braunen Platte [Blackcap],
p. 374: “That this is the appearance of the male of this
[species] of warbler cannot be doubted by anyone who
knows about birds. Its clear and loud song, which one can
watch and hear it deliver in the spring in places where trees
have been felled, settles the question. However, what the
female looks like I have been unable to learn from any-
one who has written about these birds. Even the sharp-
eyed Herr Frisch is uncertain on the matter. He writes:
Some say the red-capped one is the juvenile of the black-
capped; but it is a male, his female is exactly the same
color and has no trace of red”. Both Frisch and Zorn, de-
spite intensive researches, thought that there were male
and female black-capped warblers and male and female
red-capped warblers.

Dornreich with song-flight /Com/wow Whitethroat] , p. 377

Kleiner Steinschmdtzer [Whinchat], p. 381: Frisch includ-
ed this species in the Grasmiicken [Sylvia warblers], Per-
nau in the redstarts’: “ But I do not see on what grounds.
For it is very different from the redstarts. It is speckled,
has no red tail, breeds on the ground, in hedges, under
rocks and clumps of earth, while the redstart nests in holes
in trees or in walls. I include it therefore in the wheatears.

because it has almost the same call and song as the large
Wheatear, prefers to be among hills and rocks, and nests
under or against stones.”

Haufi-Schmdtzer [Spotted Flycatcher] , p. 382: “Moves tail
and wings at the same time when it is standing, letting the
wings hang. Has no song, male and female look very sim-
ilar.”

Schwarz und weifi-scheckigter schmatzender Fliegen-Vo-
gel [Collared Flycatcher], p. 383: “The bill of the male
is coal-black; above it he has a snow-white patch, then
above that over the head down to the nape is coal-black,
extending to the sides of the face. Below the nape there
is a white ring around the neck. The bird is coal-black
again on the back, except the rump, which has a white
patch adjoining the black tail-feathers.” In his home re-
gion, Zorn possibly only saw Collared Flycatcher, with its
“white ring around the neck”.

Haufi-Rothling [Black Redstart] , p. 386: This species was
already widespread in mid-Franken in Zorn’s time.

Hauf -Schwalbe [Barn Swallow], pp. 398-399: The Barn
Swallow, with long tail feathers, builds a flat open nest
and can sit on it without damaging these feathers. House
and Sand Martins breed in holes and have no long forked
tail! “A special providence of the Great Lord.”

Wilder Schwan [Whooper Swan], pp. 403^04: In the cold
winter of 1740 around 30 individuals “made a loud din
calling in flight that one could hear at a great distance. The
front of the bill is coal-black, then yellow towards the
head, the yellow continuing onto the skin of the front of
the head”; without a knob at the forehead.

Mergente [Smew], p. 412: Head brown; the female.

(4.) Standstill in research (1750-1788)

From around 1750 until the end of the 1780s no impor-
tant works appeared containing new findings in avian bi-
ology in Central Europe. The books by J.T. Klein (1750,
1760) simply contain lists of the bird species in Europe
and elsewhere, as far as they were known (see below). Two
loose-leaf collections of birds (B. Dietzsch, Collection of
mostly German birds, 1772-1782) and of their nests and
eggs (F.C. Gunther, Collection of nests and eggs of dif-
ferent birds, 1771-1786) appeared, containing respective-
ly 101 and 108 colored plates. The bird plates are consid-
erably poorer than those in the earlier excellent work of
Frisch (1733-1763), and only about half of the plates in
both works are accompanied by short texts (see Haffer
2007a for further details and later literature).

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The field ornithologist W.H. Kramer (1756) distin-
guished Common Nightingale and Thrush Nightingale or
Sprosser, and reported on other species in his avifauna of
Lower Austria. The Benedictine friar Leopold Vogl (1785;
see Feldner 2002) made interesting observations on nest
building, breeding, and behavior of the Siskin (Carduelis
spinus). The translation by B.C. Otto of Buffon’s Natu-
ral History of Birds which appeared in 35 volumes be-
tween 1772 and 1809 - the first six volumes had been
translated by F.H.W. Martini - was important in as much
as Otto inserted his own observations on the occurrence
and distribution of many species in northern Germany
(Wenzel 1909). Now and again he referred to the “old ex-
perienced Zorn”. These volumes of Buffon’s work found
a large public in Central Europe.

However, in the last two decades of this period of time
those researchers whose publications from 1789 onwards
would usher in the flowering of Central European field
ornithology began to be active: Johann Andreas Naumann,
followed by his son Johann Friedrich, and Johann
Matthaus Bechstein.

A compiler - Jacob Theodor Klein (1750, 1760). This
first ornithologist in East Prussia was the town clerk of
Danzig and lived from 1685 to 1759. He possessed a fa-
mous natural history collection, containing plants, animals,
minerals, and ores, which in 1740 was transferred to
Bayreuth and subsequently to the Zoological Institute of
the University of Erlangen. Following Carl Einnaeus, he
set out to classify several animal groups solely on the ba-
sis of their external characters (Geus 1970), including the
birds (Klein 1750, 1760). These classification attempts are
placed ahead of the species lists in his books. Although
Klein’s works are often cited in the literature, they are lit-
tle more than compilations (including some extra-Euro-
pean species) containing no fresh data. They are lists of
names with brief inadequate descriptions and hardly any
biological information on the individual species. Since for
many Central European species Klein gave the number of
the color plate that depicts them in Frisch (1733-1763),
we were able to compare his German names with their
present-day equivalents. A collation of Klein’s list with the
Einnaean names in Beckmann-Reyger (1774) resulted in
complete agreement. Beckmann’s list however is far from
comprehensive, leaving the identity of many of Klein’s
bird names uncertain. Klein dealt with a total of 213 Cen-
tral European bird species, a similar number therefore as
Frisch, though less than Bechstein or Naumann because
he had omitted some raptors, wetland species, gulls. Mid-
dle Spotted Woodpecker, some pipits, reed warblers, and
leaf warblers (online Appendix 2). Some of his wetland
birds, ducks, and gulls cannot be identified. The notes to
some species are quoted here verbatim because of their
historic interest:

Bonn zoological Bulletin - Supplementum 59: 1-116

Trieltrappe [Little Bustard]'. “In the year 1737 a hen was
shot and brought to me, and being so beautiful I had her
drawn [. . . ] she was close to laying and had 2 eggs inside
her.”

Gehorntes Kdutzlein [Long-eared Owl]'. Klein says “In
size the same as a wild pigeon”, i.e. ca. 32-37 cm; Scops
Owl measures only 19-20 cm.

Blaukopfige rothe Drossel and Blaukehlein mit rother
Brust [Rock Thrush]'. “The neck, the throat, and head are
cornflower-colored, the back shades into black, [. . .] bel-
ly and tail are reddish. It lives among rocks and is not so
common in our area; but in 1756 I received a pair from
Oliva, and 3 birds were sent to me from Bordeaux.”

Parisvogel [Pine Grossbeak]'. “It also comes to us in
PreuBen: They are not seen in our region every year, on-
ly sometimes.”

Klein’s natural history cabinet also held a collection of
good color plates of European and exotic birds made by
various male and female artists during 1655 to 1737,
which can still be seen in Erlangen (Braun 1906, 1908;
Gengler 1912-1913). See Schondorf (2008) for an account
of “the first German bird collection in the wesf ’, that of
Jean Holandre in Schloss Karlsberg in the Pflalz, and al-
so Holandre (1785).

(5.) The flourishing of field ornithology around 1800

The widespread practices of fowling or bird-trapping, at
a “fowling floor” ( Vogelherd) or decoy, and keeping birds
in cages and aviaries led J.A. Naumann (1789), J.M. Bech-
stein (1791-1795, 1805-1809), and J.A. & J.F. Naumann
(1795-1803, with supplements 1804-1817) to collect the
knowledge they had gained through practical experience
and observations of individual species in the wild in good
ornithological handbooks. In so doing they initiated the
flourishing of field ornithology in Central Europe.

Johann Andreas Naumann (1744-1826). “Old ” Nau-
mann (Fig. 19) was the owner of an estate in Ziebigk near
Kothen, in today’s Bundesland of Sachsen- Anhalt. He, and
his eldest son Johann Friedrich (1780-1857) after him,
were well-placed economically. They could easily afford
the purchase of equipment, guns, and books and, impor-
tantly, also had the necessary free time for their ornitho-
logical activities (Wenzel 1988). Around 1774, when he
was about 30 years old, Naumann senior wrote the man-
uscript for his first book Der Vogelsteller [The bird-trap-
per] (1789), but he only had it published, with additions,
15 years later (Baege 1980: 10). This was followed by his

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little book Der philosophische Bauer [The philosophical
farmer^ (1791), and soon after that the printing of his hand-
book Beschreibung aller Wald-, Feld- und Wasservdgel
[Description of all forest, field, and water birds] (1 795 ff)
began. Here we wish to discuss the ornithological content
of Der Vogelsteller in some detail because this work has
been neglected until now, although it heralded the flow-
ering of field ornithology in Central Europe and hence is
of particular historical significance; see Fig. 20.

Der Vogelsteller

Oder die Kunst allerley Arten von Vogeln sowohl ohne als
auch auf dem Vogelheerd bequem und in Menge zu fan-
gen nebst den dahin gehorigen Kupfern und einer Natur-
geschichte der bekannten und neuentdeckten Vogel.
Leipzig, 1789

[The Bird-Trapper

or the art of trapping all kinds of birds, without as well
as on a trapping ground, easily and in large numbers,
with accompanying copperplates and a natural history of
the known and newly discovered birds.
Leipzig, 1789].

In this little book (18 x 10.5 cm) Naumann brought to-
gether the practical knowledge and ornithological expe-
rience he had gathered over several decades as an enthu-

Fig. 19. Portrait of Johann Andreas Naumann (from Thomsen
& Stresemann 1957).

Bonn zoological Bulletin - Supplementum 59: 1-116

35 0 g c I ji e 1 1 e t

oDtr He

^ u n it

flUfrlcD Slrten wn
al« ouctl auf bem SSug«l|-(eTb Nquem
)Uib in ^enge fangtn

titbit

ben ge^irigcti ^iipftrn

u n b

tintt Iflaturgtffliiclitt NFannttn
imb rieutntbtcfttn

xn

Stn&Ka# 0'laumiuttt.

Fig. 20. Title page of J.A. Naumann (1789).

siastic bird-catcher and watcher. The first edition appeared
in 1789, it was later reprinted (1806) and then sold with
differing title pages (Schlenker 1984). Although the book
was republished with an epilogue by Baege in 1980, un-
til now its ornithological content has hardly been given
due attention. See online Appendix 3.

In the 1 980s, in the Naumann Museum in Kothen, Lud-
wig Baege discovered a manuscript written by J.A. Nau-
mann in his old age, and already known about from re-
marks made by his son Johann Friedrich. This manuscript,
which was basically a continuation of Der Vogelsteller,
was published in printed form (1989) by J. Neumann af-
ter Baege’s death. In this booklet, Naumann discusses ad-
ditional trapping methods and the experiences he had
made with them after 1790, or had heard from others, but
it contains no new ornithological findings.

The content of Der Vogelsteller is little known due to
the rarity of the original, and also because bird-trapping
for private or commercial purposes fell out of favor in the
19th century. Only Sunkel (1927), in his book on bird-
catching, provides some extracts from Naumann’s work.
When a copy of the first edition of Der Vogelsteller was
presented to one of us (J.H.) by Frau Amelie Koehler
(Freiburg) - who in turn had received it as a gift from the
library of Frau Vesta Stresemann - we were surprised to

read at the end of the long title “ und einer

Naturgeschichte der bekannten und neuentdeckten VogeF
[ and a natural history of the known and newly dis-

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covered birds]. Never have Naumann’s observations of in-
dividual bird species in Der Vogelsteller been described
by a later author, let alone commented on or analyzed. We
wish to make up for that omission in the following pages,
since Naumann’s are the first, and extremely interesting
remarks on the natural history of Central European birds
since the appearance of J.L. Frisch’s folio work
(1733-1763) and the two-volume Petinotheologie of J.H.
Zorn (1742, 1743), two publications Naumann did not
mention and almost certainly did not know. He probably
did not possess B.C. Otto’s German translation of Buffon’s
great work either, in which Otto had inserted many com-
ments on north-German birds. Both Frisch’s book and the
Buflfon translation found their way into Naumann’s library
later, and father and son quoted extensively from them,
as well as from other ornithological literature of their time;
see Hildebrandt (2007: 10).

At the end of Der Vogelsteller, Naumann introduces
himself to the reader in a comic dialogue between a House
Sparrow and a Tree Sparrow, the former saying of itself,
“my landlord is a good man, he is a bird lover and allows
me much freedom; while he sometimes punishes my too
great cheekiness he does so with moderation and within
the rules; my landlord has more pleasant things to do [than
pepper me with his gun],

Denn er hat Acker, Drescherklang.

Garten, Busch und Vogelfang,

Wiesen, Jagd und Fischerey.

Da lebt er vergniigt dabey,

Schlagt oft frohlich in die Hande,

Und wunscht sich dieB bis an sein Ende.”^

[For he has fields, the sound of threshing, garden, woods,
and bird-catching, meadows, hunting and angling. With
all this he lives happily, claps his hands for joy, and hopes
it will never end until his dying day.]

The text of Naumann’s little book, following a foreword
(pp. Ill- VI) and an introduction (pp. 1-6), is subdivided
ornithologically and not on the basis of the various trap-
ping methods, like nets, decoys, etc.:

I. On the trapping ground and the birds that can be caught
there (pp. 7-132)

II. On other woodland birds, their food, feeding, and trap-
ping (pp. 133-161)

III. On birds of prey (pp. 162-173)

IV. On field birds (pp. 174-184)

V. On waterbirds (pp. 185-199)

VI. On tame birds (pp. 200-206)

Two copperplate engravings at the end of the book illus-
trate a trapping ground or fowling floor, as well as snares,
nooses, and other bird traps.

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Foreword. In his “Foreword to the bird-trapper” the au-
thor stresses that during the migration period one must
look after the trapping ground carefully and daily, even
when one has caught no birds for days. Wind and weath-
er can suddenly turn favorable for migration and a large
number of birds could arrive at any moment. The trapper
should be able to recognize the different species and know
their calls and way of flying: “the sweet joy that one ex-
periences from the knowledge gained will never allow its
fire to be extinguished” (p. V). Most of the fowlers at that
time judged the birds they had caught solely on the taste
of their meat or on their value on the market. By contrast,
Naumann’s interest in the differences between the vari-
ous species and in their lives, their “natural history”, meant
a considerable step in the flourishing of ornithology in
Central Europe (Baege 1980: 11).

Introduction. Hunting is a favorite pastime for very many
people: “But if everyone who loves the hunt and sporting
pursuits were allowed to enjoy them it would not be long
until all game and birds were completely extirpated, there-
fore it is only natural that the upper classes have reserved
the right to these excellent pastimes to themselves and that
certain restrictions are imposed on their subjects” (p. 2).
However, Naumann continues, in most places bird-trap-
ping is a free activity, or at least can be carried out for a
small fee. He wrote his little book on bird-catching to pass
on the things he had learned to those who were interest-
ed. He had been a lover of this craft since his early youth,
and came from a long line of fowlers. He introduced new
techniques and discoveries into the subject, was grateful
to learn new ideas from other trappers, and soon was very
proficient in the art. In order that others might profit from
his knowledge he put pen to paper, and everything he
wrote about was based on his own experience.

Main text. Here Naumann discusses the individual bird
species, with details of their colors and behavior, and has
arranged them in an “ecological” order of his own (wood-
land birds, birds of prey, field birds, and waterbirds), com-
municating also much biological information. Many
fowlers of the time knew a tremendous amount about the
birds they caught and probably had no trouble following
Naumann’s explanations, even without illustrations. What
interests us today is not only the wealth of detailed knowl-
edge Naumann had of individual bird species but also the
quality of his excursions into general ornithological sub-
jects, such as territory occupation, sexual dimorphism, mi-
gration, song-supression - the holding-back or stopping
of song in decoy birds until autumn. In those early days
of field ornithology in Central Europe some bird groups
still presented difficulties when it came to distinguishing
between and identifying species. These broad topics are
dealt with first, before the discussion of single species. Our
additions to Naumann’s text are in square brackets.

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General ornithological themes

Territory occnpation in birds. “As soon as the Chaffinch
[in the spring] sings loudly he has staked his claim, and
tolerates no other finch in his territory” (p. 58). ForNau-
mann, the territoriality of numerous species was self-ev-
ident. Since Aristotle described eagles holding territories,
the occupation of territory has been rediscovered inde-
pendently by many ornithologists (Pernau and Altum in
Germany, Howard and Lack in England; see Birkhead
2008).

Sexual dimorphism. “It must be additionally noted that
the male of many birds looks much more beautiful than
the female, while in others the sexes can be differentiat-
ed only by being a little lighter or darker, which can hard-
ly be seen even by an expert, and in others again the col-
ors are very different, and where this is the case I have
described the colors of both sexes” (p. 59).

Bird migration and winter quarters of Central Euro-
pean species. Migration is a well-known phenomenon,
writes Naumann, and each group of birds seeks its way
after its own fashion: “As a sailor on the sea finds his route
with the help of a compass, so a bird finds its route again
in the air by following its nature from one year to anoth-
er” (p. 11). Naumann thinks that migrants leave their
breeding grounds mainly because of the scarcity of their
food in the autumn, not because of an innate instinct. In
some places unusual weather conditions can bring alien
birds, which are regarded as a sign of bad luck (p. 16).

The Nightjar “is a summer visitor, and as soon as it finds
no more flies it moves elsewhere. All these birds, like the
Cuckoo, Golden Oriole, Hoopoe, Dunnock, Nightingale,
reed warbler. Barred Warbler \and/or\ Lesser Whitethroat,
Icterine Warbler, Willow Warbler, Robin, Common Red-
start, Goldfinch, Whinchat, Tree Pipit, wagtail, and Wren
move at night when it is still, partly because they are safe
then from raptors and partly because the wind usually does
not blow strongly at night, since these birds are not strong
fliers” (p. 155). “All the birds that cannot find their food
in the winter in our region migrate to warmer countries,
since a bird that feeds on insects will find none in the win-
ter and so must naturally move to a country where it can
find such food. For this it does not require a long period
of time for a bird can comfortably fly four or five miles
in an hour. So if the wise Creator has made this part of
the nature of all birds, I don’t know why He should not
have given this ability to the Swallow instead of, as many
claim, changing them into something else, or causing them
to bury themselves in swamps or hollow trees? Who will
extract them from these places, and dry them off so that
they can return to their life?” (p. 156). Naumann rejects
the widespread belief that swallows overwinter in the mud
of lakes.

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Song-suppression. Naumann mentions the suppression of
song several times (“Gesang-Verhaltung”; Aitinger 1626,
1653: 167) when writing about the decoy birds or lures
on the trapping ground, without explaining this practice,
which once was widespread and known to fowlers for cen-
turies. “Suppressed birds are those that sing at a time of
year that is different than the normal time for singing, and
which are very useful in the autumn for attracting and
catching wild birds” (Riedel 1833: 70, footnote).

“Many bird-catchers keep their Chaffinches all summer
in a dark place so that they will sing when desired in au-
tumn or winter, since as soon as they come into the light
again they start to sing but do not sing in the dark, and
they are used to lure other birds into the net” (Gessner
1555; see also 1669: 120-122).

“As soon as the bird which one wishes to suppress starts
to sing, or just before this happens, it is put in a cage and,
when it has found its food and water containers, the place
where the cage is kept, perhaps in a box, is gradually made
darker and darker until it is completely blacked out. The
bird should be kept cool but comfortable and clean. Some
keepers pluck out feathers at this time so that an artificial
molt will prevent singing. Once the bird is in complete
darkness, and knows where its food is, it should remain
there until it is needed. However, at least four weeks be-
fore it is to be used it should gradually be reintroduced to
the light and fed well, then it will let its song be heard”
(Riedel 1833: 70-71).

“Regarding Chaffinches, Bulliard [1778] advises catch-
ing birds of the year in September or October to keep them
as decoy birds for the start of the following spring. Then
those that are best at luring wild birds are selected. So that
they will sing in the coming autumn, when they will be
most needed as decoys, one must deceive them as to which
season it is, which the bird-trappers call suppressing or
stopping (Verbal ten). For this purpose, at the beginning
of June the decoy birds are put into a cool room or cel-
lar, and their cages made gradually darker and darker un-
til they are in total darkness. A cage must then not be
moved because the birds will be accustomed to finding
their food and water at a particular place. An even better
procedure was to let them feed every day in artificial light,
and to pluck out some tail or wing feathers to create a
‘forced molf . At the end of September they were slowly
accustomed to daylight, and then would fulfill their role
as singing decoy birds in October” (Sunkel 1927: 234).

Therefore the fowler accelerated the annual cycle of the
caged decoy birds by bringing them relatively rapidly
through an artificial autumn, with molt, in early summer,
from which they emerged between August and October
singing their spring song. As stated above, fowlers had
been using this technique of song-suppression for cen-
turies. Birkhead (2008) and Birkhead & van Balen (2008)
mention early reports on this by Conrad Gessner (1555),
Manzini (1575), Aldrovandi (1599), and Aitinger (1626),

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and discuss the connection between suppression (or stop-
ping) of song in cage birds and the avian annual cycle.

J.A. Naumann’s remarks on song-suppression in decoy
birds and the advantages of suppressed decoys during au-
tumn bird-trapping are as follows:

“The song must do most of the work, because for them
[the migrating conspecifics] it is something unusual at this
time of year so they enjoy listening to it and swoop down
eagerly to the trapping ground, hence a bird-catcher must
above all ensure that he gets excellent singers” (p. 59).
“When one [in the spring] has about eight of them
[Chaffinches in cages] then they are hung outside at a win-
dow, and after they have sung loudly for a few days, and
no longer stammer, they are brought in and put in a dark
closet or box until St. Bartholomew’s Day [= August 24th],
when they are taken out and hung up again outside. Those
birds that start to sing again inside a few weeks are good
ones, and those that do not sing can be rejected and re-
placed by others after their time outside” (p. 60). “A good
Chaffinch will start his song in March or April, but those
that start singing at the end of May, or even around St.
John’s Day [= June 24th] are of no use because they will
very rarely pick up their song again in autumn” (p. 63).
According to Naumann, blinded birds sing much better
than sighted ones because nothing disturbs them. “If one
wishes, one can put a good male [Yellowhammer] [...],
as with the Chaffinches, in a dark closet, so that he keeps
his song until autumn, then one will attract far more wild
birds at the trapping ground” (p. 71). “The decoy birds [of
the Ortolan] are kept in darkness, just like the Chaffinch-
es, in order that their song is retained because without good
song nothing much will be achieved [in the autumn]” (pp.
73-74).

Species knowledge. The list of species dealt with is re-
stricted to the birds which Naumann observed in the im-
mediate vicinity of his place of residence in northern cen-
tral Germany. This is the reason why there are gaps in the
wetland species (swans, geese, herons, ducks, gulls,
waders) and in some of the songbird groups (reed war-
blers, leaf warblers, Regulus sp., treecreepers). However
he filled many of these gaps in later years.

Trapping methods

Trapping devices like the fowling floor and its lures, nets,
nooses, and cages are discussed at length in Part 1 of Der
Vogelsteller. A trapping ground must be erected in a fa-
vorable place, with singing birds and decoys. When set-
ting up a trapping ground one must “know the thoughts
of the birds, if I may so express myself, so that one erects
everything just the way the birds like to have h” (p. 10),
that is, the fowler must know their habitat schema. The
main point is to “ abolish everything of artful appearance

Bonn zoological Bulletin - Supplementum 59; 1-116

that to our eyes appears beautiful, and to arrange the trap-
ping ground so that in the eyes of the birds it seems nat-
ural and attractive” (pp. 42^3). One catches few birds in
strong winds or in mist; the best kind of weather is over-
cast and still. Advice on the keeping of decoy birds fol-
lows, and on the best type, size, and fittings for a bird cage,
as well as on the various kinds of food. Naumann also dis-
cusses nets and traps for different bird species (pp. 48-57).

On the different kinds of trapping grounds or fowling
floors (pp. 91-111): An area for catching Chaffinches
(Finkenherd) must have a covering of thick grass as well
as trees around the trapping ground, for the wild birds to
alight on, and from which to hang the decoy-bird cages.
Naumann goes on to describe many additional con-
trivances on the trapping ground for other decoy birds,
such as the “runners” (Ldufer), which are kept on the
ground by having their wings tied to their bodies. On a
“shrubby trapping ground” (Strauchherd) the grass in an
open area with shrubs is cut evenly using a sickle, the grass
below the shrubs is removed, and the soil swept clean.
Twigs of rowan berries are stuck in the ground beneath
these shrubs as well as attached to them, the more the bet-
ter. The bird-trapper transports his nets and cages only on
long poles. He watches the trapping ground from a hut and
observes how many birds arrive. From the courtship calls
of his lures he must be able to recognize whether the wild
birds are near or far, and to distinguish courtship calls from
warning calls. He must be especially on the look-out for
weasels and birds of prey, which can easily kill the decoy
birds.

On snares (springes) and nooses (pp. 1 1 1-122): A wide
range of trapping devices, many of which Naumann him-
self improved, are decribed in great detail and with illus-
trations.

I. On the trapping gronnd and the birds that can be
canght there (pp. 7-132)

In each species section, before his general and biological
remarks, Naumann provided a brief description of
plumage, which we have omitted here.

Fink [Chaffinch] (pp. 58-64): The bird-trappers distin-
guish between different classes according to their song
{Schitzkebier, Deutschebier, Reitzu, Nutschkebier and
small Schitzkebier) [these fantasy names are onomatopoe-
ically based on the various songs], and then there is the
Waldgesang [woodland song], which cannot be compared
with them. “Bird-catchers are of differing opinions con-
cerning the song. Some say that the small Schitzkebier is
most suitable for attracting wild birds, others praise the
Reitzu, personally I believe that the best song is the one
that is the commonest among the Chaffinches” (p. 64).

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Qudcker [Brambling] (pp. 64—66): After the Chaffinch this
is the most important bird on the trapping ground. “Where
it nests I cannot say, because I have never seen a Bram-
bling in the breeding season in our region” (p. 64). Young
cocks make the best decoy birds.

Schwunsch [Greenfinch] (pp. 66-68): Migrates with the
Chaffinches into December, but singly. It particularly likes
hemp seeds.

Zeisig [Siskin] (p. 68): “The smallest bird among those
that are caught on the trapping ground, but it tastes the
best. [. . . ] They hang continually on the fruits of alders and
eat their seeds.”

Schdttgen or Schittscherlingen [Redpoll] (p. 69): “The
character and qualities of the Redpoll are exactly the same
as those of the Siskin. They are always together with
them.” They are rare, though singles are seen almost every
year, and in some years they are very common.

Stieglitz or Distelfink [Goldfinch] (pp. 69-70): A well-
known bird, which however is only caught as singletons.

Hdnfling and Steinhdnfling [Linnet and Twite] (p. 70):
There are (a) the large red-breasted Linnets and (b) the
small Twites, and both of them are only caught singly and
with some trouble. They are frequently seen in quite large
flocks on the stubble fields in winter.

Griinschling or Gehling [Yellowhammer] (p. 71): Anoth-
er well-known bird that is caught as singletons.

Strumpfweber [Corn Bunting] (pp. 71-72): “They get their
name from their song, which sounds like a stocking-weav-
er working at his loom.” They move away in September
but many remain here all winter.

Schiebge or Rohrammer [Reed Bunting] (pp. 72-73):
Lives in bushy places where it is marshy and where there
is plenty reed, whose seeds and those of other grasses it
feeds on. “It does not breed in our region.” Their main
movement is in October. Reed Bunting, Yellowhammer,
Corn Bunting and Ortolan are closely related to each oth-
er.

Ortolan [Ortolan Bunting] (pp. 73-74): “Where this valu-
able bird breeds I have never been able to discover” (p.
74). These ‘Ortolans’ are in fact young Reed Buntings (see
Hildebrandt 2007: 162, footnote).

Heidelerche [Wood Lark] (pp. 74-76): “In the breeding
season they stay near trees and sing from them, but when
they are migrating they no longer perch in trees. They dis-
like being in open fields” (p. 75).

Bonn zoological Bulletin - Supplementum 59; 1-116

Krammetsvdgel [‘juniper birds] (p. 77): This name is
used in our region for thrushes. Blackbird, Fieldfare, Mis-
tle Thrush and Waxwing “because they all have the same
character and all like to eat juniper berries.” They are de-
scribed as follows:

Drosseln [thrushes] (pp. 77-82): The Zippdrossel [Song
Thrush] is the first to arrive in spring. They migrate most-
ly at night, as do Robin and other small birds. By day, for
their food they search for berries in the hedgerows as well
as for insects and worms, etc. in meadows. The Wein-
drossel [Redwing] is slightly smaller than the Song Thrush
and is dark red on the flanks and under the wings.

Amsel [Blackbird] (pp. 82-84): There are three kinds: (a)
Ringamsel or Schildamsel [= Ring Ouzel], (b) gray Black-
bird [= female] and (c) black Blackbird [= male]. They
migrate at night.

Schnarre or Schnarrziemer [Mistle Thrush] (pp. 84-85):
This is the biggest of the ‘juniper birds’ and likes to be in
large woodlands with grassy clearings. Their contact call
is a rattling sound.

Ziemer or Krammetsvogel [Fieldfare] (pp. 85-88): The
Fieldfare probably breeds in Poland, Russia, etc. Irregu-
lar movements in winter.

Seidenschwanz [Waxwing] (pp. 88-89): Eats rowan
berries and is a winter visitor which does not appear every
5 or 7 years but occurs irregularly, whenever it likes, just
like all other birds that do not appear regularly or at def-
inite times. It probably comes to us from colder countries
since it is one of the largest winter visitors and appears
only in the hardest winters. “They come in large flocks
and readily come down [to the trapping ground], so that
one can expect to catch even the last one” (p. 89).

Schmiel, Dompfaff or Gimpel [Bullfinch] (pp. 89-91): A
well-known bird. “If one can obtain them young then they
will easily learn to whistle all kinds of songs, like the Lin-
net. Where they nest I cannot say because I have never
seen a Bullfinch, let alone a nest, in this area in the breed-
ing season {footnote, they breed in Bohemia, Thtiringen,
Hessen, etc.). They do not come to us every year but usu-
ally every second year.”

Staar [Starling] (pp. 1 22-1 24): A well-known bird, which
(if trained when young) will learn to whistle and speak.
“As soon as they have raised their young they gather in
large flocks and fly to damp meadows with wet places
where reed grows, where they roost at night. Whoever
wants to make a decoy to catch them should not live too
far from such areas where the birds gather, because while
the birds are not actually migrating they simply go where

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food is to be found, so it can be assumed that they will
not move far from their usual haunts. The time to catch
them starts at Whitsun and lasts until harvest time. Then
the Starlings go to the pastures where cattle graze and for-
age for food there until autumn, when they move away”

(p. 122).

Raubvdgel [birds of prey] (pp. 124-126): Some species
are dealt with here which are often responsible for killing
the bird-trapper’s decoy birds. The Habicht and Sperber
[ Goshawk and Sparrowhawk] (in both of which there is
a larger [= female] and a smaller kind [= male]). They pur-
sue small birds and are the worst culprits at the trapping
ground. They are “the sparrow’s greatest enemy, which
they even attack, without the slightest fear, in the farm-
yard where they feel safe. They do not even hesitate to
chase them right up to open stable doors or windows” (p.

125) . The Lerchenfalk [Hobby] is as big as a small Spar-
rowhawk and just as pernicious; “even if it mostly takes
larks in the fields a few times, it soon gets an inclination
to eat woodland birds too, but one has to watch for it es-
pecially at a trapping ground set up for Wood Larks” (p.

126) . “The remaining birds of prey do no damage at the
decoy, except in winter when they become hungry. So it
has often happened to me that the Grofier Mausehabicht
[Common Buzzard] has attacked my Fieldfare cages, and
at that time of year one cannot trust any bird” (p. 130).

Kriickelstern and Neuntodter (pp. 126-128): There is the
large [Great Grey Shrike^ and the small Kriickelster
[Woodchat Shrike^ and the Neuntodter [Red-backed
Shrike]. Many of the Great Grey Shrikes stay here in win-
ter. They “feed on birds, beetles, and small frogs, and like
to make their nests in wild pear trees which stand isolat-
ed in the fields” (p. 126). “The Woodchat Shrike is a good
bit smaller with a red cap on its head and also nests in
trees.” Red-backed Shrikes nest in thorn hedges. This
species and the Woodchat Shrike are summer birds, “and
before one has begun to catch Chaffinches they are already
gone”. All of them can be very harmful at the trapping
ground because they can attack the decoy birds through
the bars of the cages. “It is said that the Red-backed Shrike
must kill nine kinds of worm, larvae, etc., and when it is
sated sticks the remainder onto thorns; although this
sounds ridiculous I have actually found it to be true that
it spears worms, etc. on thorns when it has eaten enough,
in order to eat them another day. And it can often happen
that it does not find them again, or has been chased away
from the area; for example I once found a little frog, neat-
ly impaled through the mouth on a pointed dry twig and
completely dried out. When these shrikes pull their prey
apart they do not hold it under their feet, so they have to
stick it on a thorn or pointed twig to be able to pull pieces
off according to their appetite” (pp. 127-128).

Bonn zoological Bulletin - Supplementum 59: 1-116

He her and Holzschreyer (pp. 128-130): There are two
species, one of which is called Nufiheher or Holzschrey-
er [Eurasian Jay]. It is very common in the forests. “The
other species is not so common and is seen in our region
only very rarely, and I have encountered it only twice in
22 years. They are the same size as the former and look
completely black with white spots like the Starling and
white bands on the tail. Some call it the Tannenheher [Nut-
cracker] and others the Nufiheher. NuBheher is a good
name for it because it likes to eat nuts, which I have nev-
er seen the Jay do. Their migration is at the same time as
that of the Song Thrush, and they sometimes appear in
large flocks. They eat all kinds of vermin and tree fruits”
(p. 129).

Kreutzvogel or Krinitz [ Common Crossbill] (pp. 130-1 32):
“Around the year 1760 in the month of August 2 of these
birds were seen in my copse, sitting in the rowan trees and
splitting the berries apart to eat the seeds.

I immediately made a snare and hung it among the
berries, and when I returned I had caught the female but
the male was gone. This female looked yellow and had
black wings and a black tail. I fed her with hemp seed and
oats which seemed to do her good; she had a quiet and
charming song, this being an unusual thing among other
birds. The cock was red on the sides and front of the breast,
almost like a Chaffinch. In the spring of 1786, in the breed-
ing season, another pair appeared, the cock of which
looked Just like the previous one and had a strong and
beautiful song that could be heard at a distance.

However the female was ashy gray with brown cheeks,
and below her tail were several brown feathers. They built
their nest in a loose hedge, about 6 feet [2 Ellen] above
the ground, of dry stalks, like a warbler’s nest, lined with
horsehair. The eggs were like those of a Red-backed
Shrike in size and appearance, though spotted with light
red not dark red like the shrike’s eggs. They had 4 eggs.

Apart from these I cannot recall having seen Crossbills
in our area. Which is why I cannot include them in the
migratory birds; their proper home is supposed to be in
spruce forests, where they feed on the seeds from the
spruce cones; they hold the cone with their strong and
sharp claws while they extract the seeds with their hooked
bill, like the Siskin does with alder fruits.

[. . . ] Otherwise I have heard it said that these birds [. . . ]
do not breed in spring like the others but in January. Al-
though this sounds foolish, anyone who has no experience
of the matter should believe it until more definite infor-
mation is available, and this is why I have described my
own experience of these birds at such length, from which
the reader can see how tireless my efforts have always
been to discover the real nature of bird-trapping” (pp.
130-132).

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II. On other woodland birds, their food, feeding and
trapping (pp. 133-161)

Guckguck or Kuckuk [Cuckoo] (pp. 133-135): It is not a
bird of prey although it looks like the small Sparrowhawk;
however, bill and claws are more like those of the Black-
bird. “When it sits on a twig it grasps it with two toes be-
hind and two in front, which no other birds do but the
woodpeckers. [. . . ] Its food mostly consists of caterpillars,
beetles, butterflies, and all kinds of worms; hence they of-
ten swoop to the ground to take earthworms” (p. 133).

“Because the Cuckoo does not raise its own young, the
Creator has made it part of its nature to lay its eggs in the
nest of another bird which incubates them and rears the
young as its own. It never lays more than one egg [in a
nest], and always chooses the nest of a bird that feeds its
young with food that agrees with the Cuckoo’s nature. It
throws the bird’s own eggs out of the nest with its bill,
and since this often results in most of the eggs being bro-
ken, some have thought that it sucks the eggs, but this is
false because I have found both whole and broken eggs
below nests.

It often lays its egg in the nest of a wagtail or a war-
bler, which is proof that it cannot be a bird of prey be-
cause if it were it would lay its eggs in their nests since
they feed their young with all kinds of meat. Some say
that when the young Cuckoo has been raised then as a sign
of gratitude it eats its foster parents, and even claim to have
seen how it snaps at them, but all this has as little basis
in fact as what has been formerly said. If it snaps at them
then either from hunger or from anger at having been
brought too little food, for the wagtails have their work
cut out just to satisfy such a large bird. I have watched in
my own garden how a pair of wagtails fed a young Cuck-
oo, and they stayed in the same place long after the Cuck-
oo had left the nest and could feed itself.

After this I raised young Cuckoos and shot adults, and
having dissected them and examined their nature I could
find not the slightest evidence that they were raptors. If
we just look at the Red-backed Shrike, although it is the
smallest bird of prey it still has a hooked bill and power-
ful claws, both of which the Cuckoo does not have. It ceas-
es to call around St. James’s Day [= 25th July] and then
forages for its food in silence, and departs before St.
Bartholomew’s Day [= 24th August]. Because it stops call-
ing just when the fields are being harvested, when the
Hobby comes to forage, and since the Hobby at a distance
can resemble the Cuckoo, then ignorant people say that
the Cuckoo has changed into a bird of prey. The songbirds
mob it, and when they see it they pursue it with biting and
nipping. The Cuckoo defends itself by biting back, but I
have never seen that it would chase one or kill it, although
I have sat for many hours in the woods watching for this.
Whether this mobbing by the small birds comes from the
Cuckoo’s looking like a raptor or from its sometimes be-

Bonn zoological Bulletin - Supplementum 59: 1-116

ing caught at the nests one cannot know for certain, though
the former seems more likely to me than the latter. The
Cuckoo is on the whole a useful bird in the garden because
it feeds mostly on caterpillars, and in the spring it is one
of the most agreeable of the messengers that announce
pleasant summertime to us” (pp. 134-135).

Wiedehopp [Hoopoe] (pp. 136-137): “Its song consists of
a hollow note that sounds like hop, hop, hop. It feeds on
worms in the earth and other kinds of larvae, etc. It makes
its nest in hollow trees and because it smells so horribly
in the cavity some people say that it uses human excre-
ment in its nest, but this is wrong since the smell comes
from the bird’s own droppings. [. . .] It arrives not earlier
than April, when it becomes properly warm, and leaves,
like all summer visitors, during the month of August.”

Spechte [woodpeckers] (pp. 137-139): There are 6
species: Griinspecht [Green Woodpecker], Schwarzspecht
[Black Woodpecker] , Grofier Buntspecht [Great Spotted
Woodpecker] (in spring this one seeks “a dry broken end
of a branch and drums on it with its bill [. . . ] instead of a
song”), Kleiner Buntspecht [Lesser Spotted Woodpecker] ,
Blauspecht [Nuthatch], and Grauspecht or Wendehals
[Wryneck]. “The little Baumldufer [treecreepers] are al-
so included in the woodpeckers; they are as big as a Siskin
and have a long curved bill to dig out tiny verminous an-
imals from the cracks in trees.”

Meisen [tits/chickadees] (pp. 139-142): We have 5
species: Kohlmeise [Great Tit], Blau- or Pumpelmeise
[Blue Tit], Plattmeise [Marsh Tit], Tannenmeise [Coal
Tit], and Spiegelmeise [Long-tailed Tit]. An exact descrip-
tion of the hit-dance’ follows, used to catch these species,
in which a bird with broken feet flutters around.

Kernheifier [Hawfinch] (p. 142): The Hawfinch or
Kirschfmk [Cherry Finch] bites a cherry stone as if it had
been split by a knife; they also like to eat hemp seeds. They
are only caught singly on the trapping ground.

Spanier [Dunnock] (p. 143): Migrates singly and mostly
at night; they are only caught as singletons on the decoy.
“Where they breed I have not been able to discover.”

Nachtigall [Nightingale ] (pp. 1 43-144) : This bird is well
known, it nests close to the ground. “It mostly returns to
the same place to build its nest. It departs during the month
of August.” They can be caught on the trapping ground
for Chaffinches.

Pfingstvogel [Golden Oriole] (pp. 144-145): “It builds its
nest in trees, and it always looks for a narrow branch that
does not grow straight upwards but has a fork at the end.
In this fork it makes its nest, which hangs in the air, not

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supported on a branch like that of other birds. [...] It is
the largest summer visitor, leaving in August and return-
ing to us only in May. It eats especially cherries and el-
derberries.”

Weifikehlen /^Sylvia warblers] (pp. 145-146): There are
4 species: (1) Grasemucke with white \hrodii [Garden War-
bler], (2) Kleine Weifikehle [Lesser Whitethroat] is
brown on the wings, (3) Fliegenschndpper [Common
Whitethroat] is smaller still and has an ash-gray head and
snow-white throat, and (4) Plattmonch or Murrmeise
[Blackcap]-. “The cock has a black cap on his head and
the female a brown one. This bird has a beautiful song that
almost sounds musical, and when the other birds stop
singing around St. John’s Day [= 24th June] this one is
just getting started and sings until St. James’s Day” [= 25th
July]. In the autumn it feeds on berries like the Fieldfare,
and departs in September and October. The 3 other species
migrate in August and September, return to us again in
April and nest in hedges.

Schackruthge [Icterine Warbler] (p. 146): “It arrives with
the Golden Oriole in the month of May, and departs with
it in August. Its song is exquisite and powerful.”

Fitis [Willow Warbler] (p. 147): Looks similar to the Icter-
ine Warbler but is smaller and has a pale stripe over the
eye, and also stays longer.

Distelfink or Fliegenschnapper (pp. 147-148): There are
2 species of Distel-, Lohr- or Spiefifink, namely large
[Spotted Flycatcher] and small [Pied Flycatcher, female
and brown male]. The small ones look similar to female
Chaffinches, but eat mainly insects and live in low thorny
and thistly hedges. “They are as big as Lesser Whitethroat,
and nest in hollow trees.” The large one is light gray and
appears slightly scaly. It makes its nest of moss, like a
Chaffinch’s, on the old branches of trees and on pollard-
ed willows.

Rotschwdnze [redstarts] (p. 148): There are 2 species: the
common Rotschwanz [Common Redstart] nests in hollow
trees as well as in crevices and holes in walls. The Schild-
nachtigall [Bluethroat] “searches for its food almost al-
ways on the ground and is often found in ornamental gar-
dens as well as those where cabbage, lettuce, beans and
such plants are grown. It has a beautiful song and sings
at night like a Nightingale. [. . . ] In the months of August
and September it migrates.

Rothkehlgen [Robin] (p. 149): A well-known bird which
departs in September and returns in March. The young are
scaly brown, without a red throat.

Zaunkonig and Goldammerchen [Wren and Gold-
/Firecrest] (pp. 149-150): The Wren has a very strong
voice for the size of its body. The Gold-/Firecrest is slight-
ly smaller, has a yellow stripe on its head and always
moves around in the company of tits. [Firecrest and Gold-
crest were only distinguished in the 19th century.]

Rohrschliefer or Rohrsperlinge (pp. 150-151): There are
3 species: (1) the Grofie Rohrsperling [Great Reed War-
bler] lives on lakes and marshes with reedbeds, builds its
nest over the water and sings day and night, (2) the oth-
er species [Eurasian Reed Warbler] is as brown as the sec-
ond Weifikehle [Lesser Whitethroat], also lives in reeds
on lakes but its song is not so loud. (3) “The third species
is slightly smaller again, looks rather like a Dunnock and
has a light-brown stripe mixed with yellow on its head
[Aquatic Warbler?]. Its song resembles the chirruping of
a cricket or grasshopper, which is given in one long breath
[?]. They all nest above water in sedge and reeds, or in
shrubs among willows. They feed on all sorts of worms,
etc. from the water and depart with the summer visitors.”

Spiefilerche [Tree Pipit] (p. 151): “Breeds in the woods
and builds its nest in grass on the ground. It has a rather
attractive song, and when it sings it flies high above the
trees, letting itself down onto a tree, singing all the while.
It migrates in August and September.”

Krautlerche [Whinchat] (pp. 152-153): “Its shape is al-
together very like that of the Wheatear, only it is small-
er. In spring it remains in young tree plantations and field
hedges, where it nests; it likes to build its nest in mead-
ows, in the grass. It has a beautiful song; in the middle of
August it starts to migrate and then is only found in the
fields, especially among cabbages.

Bachstelzen [wagtails] (pp. 153-154): The blue wagtail
or Ackermann [White Wagtail] is one of the earliest sum-
mer visitors to arrive in spring. It nests in hollow trees,
thatched roofs and on river banks, etc. The Gelbe Bach-
stelze [Yellow Wagtail] belongs to the field birds and fre-
quents meadows and fields. It arrives later than the White
Wagtail in spring and is a welcome messenger for the
farmer because it heralds the summer.

Tageschldfer [Nightjar] (pp. 154-159): A nocturnal bird
with a thin bill. “But when it opens the bill its whole head
splits in half, like a pair of pincers. During the day it sits
and sleeps below hedgerows on roots and stumps, as well
as in the middle of forest rides, and sleeps, so that it can
often be mistaken for a piece of tree bark. At night, on the
other hand, it flies around feeding on gnats, beetles, and
all kinds of small animals, which it skillfully catches in
the air. It is a summer visitor and as soon as it finds no
more flies it moves elsewhere.”

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Waldschnepfe, Becasse [Woodcock] (pp. 159-1 60): The
Woodcock has a soft bill, its food is earthworms, slugs and
larvae in the soil. “During the day it forages for food un-
der hedgerows, and in the evening when twilight begins
it flies to the meadows and arable fields and looks for food
there; they make their nest in grass on the ground, but that
is something rare in our region. They migrate in October
and return in March and April; they migrate in the night;
when they start off at twilight they call loudly, making a
sound almost like the chirruping of young chickens; they
only do this in spring though, not in autumn.”

Wilde Tauben [wild pigeons] (pp. 160-161): The biggest
species is the Ringeltaube [Wood Pigeon] \ the common
Holztaube [Stock Dove] is rather smaller than the domes-
tic pigeon. The Turteltaube [Turtle Dove] is smaller, light
brown with a black-gray tail. Wild pigeons nest in trees,
and depart in autumn to return in spring.

III. On birds of prey (pp. 162-173)

Krdhen [crows] (pp. 162-164): There are 6 species: (1)
Rabe [Raven] with its gruff, staccato voice and wedge-
shaped or round tail (straight as if cut off in the crows);
not as common as the crows; (2) Schwarze Krdhe [Car-
rion Crow], similar to the Raven but smaller; (3) Schild-
krdhe [Hooded Crow] is just as big as the previous one,
black but with ash-gray belly and back. These last two
crows have “a single nature and characteristics, so much
so that they sometimes mate with each other”. (4) The
black Feldkrahe [ Rook]', the bill is weaker than in the pre-
vious species, and the skin towards the head rough and
white. They damage root crops but this harm “is not by a
long way as great as the good they do because they eat
all the creatures harmful to crops like caterpillars, beetles,
grasshoppers, etc.” (p. 163). (5) Dohle or Schneekrdhe
[Jackdaw], black with an ash-gray neck. (6) Blaue- or
Mandelkrahe [European Roller]. They feed on beetles and
other vermin, do not take smaller birds, and do not asso-
ciate with the other crows. “They belong to the summer
visitors, but are not seen so frequently as the others.”

Elster [Magpie] (pp. 164-1165): A well-known and very
injurious raptor that searches for nests of Pheasant and Par-
tridge and hunts young Pheasants, Partridges, ducks and
hares. When they realize that someone is climbing their
nesting tree “they carry their young away to another nest,
and so it is probable that they have two nests”.

Eulen [owls] (pp. 165-166): There are 5 species: (1) The
Uhu or Schuhu [Eagle Owl] “is the largest, it lives in the
high hills and is seldom encountered in our area”. (2) The
Knappeule [Tawny Owl], brown mixed with ash-gray, “a
harmful visitor to dovecotes, which it frequently enters at

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night to pull out the young pigeons”. (3) Horn- or Ohreule
[Long-eared Owl], smaller than the previous one, dark
gray with two feather ‘ears’; (4) Kohleule [Short-eared
Owl], brown like a Tawny Owl; “it is often found in fields,
among cabbages, also in meadows and in long grass”. (5)
Kauz [Little Owl], the smallest species and pale gray, nests
in hollow trees, old walls, and buildings. “Since owls are
not seen during the day birds are not used to them, and
their mobbing behavior is sometimes shown even towards
the Nightjar, although it is not a bird of prey” (p. 166).

Falken [ ‘falcons ] (p. 1 66): “The Adler [White-tailed Ea-
gle] is the largest of them, but is seldom seen in our re-
gion. It is black-gray in color with white spots [?]. The
bill is one inch straight and then is hooked at the end,
which distinguishes it from other birds of prey. The
Steinadler [Golden Eagle] is slightly smaller”.

Mausehabichte [Rough-legged and Common Buzzards,
with color morphs] (pp. 166-168): There are 5 kinds: (1)
Plumage dark black-gray, speckled a little with white on
the breast and under the wings. (2) Similar to the first
though not colored blackish but brownish, and with more
white speckling on the breast and under the wings; this is
the commonest kind; (3) similar to the previous two, “ex-
cept that this kind has shaggy feet like an owl” [Rough-
legged Buzzard]; (4) also similar, but with a longer tail
and white rump; (5) with a longer tail and longer wings,
speckled with gray, brown and white.

Fischaar [Osprey] (p. 168): Hunts mainly fish, “then they
hover in the same spot in the air until they see a fish, when
they shoot down and under the water like a duck, seize
the fish, and fly off with it. It does not remain here in the
winter”. Its legs are covered with rough skin, like a wood-
file.

Weyhen [harriers] (pp. 169-170): There are 4 species: (1)
Gabelgeyer [Gabelweihe = Red Kite]', it flies slowly and
cannot catch any prey in the air; hunts young ducks and
geese; (2) Moosweyhe [Marsh Harrier] only found near
water and marshes; (3) Kleine Weyhe [Montagu ’s Harri-
er], also known as the Milane [kite], smaller than the pre-
vious one, pale gray with a white rump, yellow legs, and
white belly and breast streaked with black, “flies slowly
and low, always looking down at the ground”; (4) Weifie
Weyhe or Weifier Falke [Hen Harrier] is not as big as the
previous one, white shading to ash-gray, wing-tips black,
legs yellow. Shape and behavior as the previous harrier.
All harriers leave in the winter.

“The small Rudelgeyer [Common Kestrel] can also be
included in the harriers. [. . . ] It can hover for a long time
in the same spot in the air by continuously steering or flut-
tering with its wings, hence its name of Rudelgeyer [‘row-
ing raptor’]. It lives mostly on mice/voles” (p. 170).

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Blaufufi [Peregrine Falcon] (pp. 171-172): “In our region
we have two species of falcon. The large one is called
BlaufuB [‘blue foot’] and the other is the small Lerchen-
falke [Hobby]. The Peregrine Falcon is “the worst enemy
of Partridges and pigeons. As soon as it spies a flock of
pigeons it looks for the slowest one, mostly selecting the
white or colored ones. It shoots towards them like an ar-
row, but if the pigeon succeeds in reaching a hedge, tree,
or water (since a pigeon can dive in water like a duck,
which I have seen with my own eyes) then the falcon will
retire. But if the pigeon does not manage to reach such
safety it will fly as fast as possible, skillfully avoiding the
falcon’s attempts to seize it by flying higher, though the
raptor will try to climb above it, then the pigeon repeats
the procedure. If the pigeon succeeds again and again in
being higher than the falcon then the latter will finally tire
and turn away, allowing the pigeon to escape, which I have
often observed. Otherwise it is a shy bird and difficult to
shoot.”

IV. On field birds (pp. 174-184)

Trappe [Great Bustard] (pp. 174-175): The largest field
bird, nests on the ground in cereal fields, lays 2 eggs, eats
green crops and plants, does not migrate in autumn and
can only be caught with some cunning.

Krannich [Common Crane] (p. 175): Avery cautious pas-
sage bird.

Rebhuhner [partridges] (p. 176): There are (a) Reb- or
Buschhuhner and (b) Feldhuhner, which breed respective-
ly in copses or in fields. But it is not said whether they
differ in their plumages.

Wachtel [Common Quail] (p.l77): Returns in May and
leaves again in autumn. They do not live in groups like
the Partridge “but mostly fly around alone”.

Wachtel konig, Schnarker [Corn Crake] (pp. 177-178):
The Corn Crake “breeds in meadows and cereal crops and
migrates with the Quails; it does not lay many eggs which
is why they are not as numerous as Quail. [. . . ] Their song
sounds like the creaking of a door and is not exactly pleas-
ant to hear”.

Lerche [Sky Lark] (pp. 178-179): “It is widely known
what kind of bird the Sky Lark is, it arrives here in March
and nests in our fields. [. . .] Because the Sky Lark has a
beautiful song it is kept by many people in a cage and fed
with oats”.

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Hiister [Meadow Pipit] (p. 179): “The Meadow Pipit is
in shape and color very like the Tree Pipit, though a lit-
tle smaller and less yellow, but otherwise it is of exactly
the same character. It is a bird of the fields and is caught
along with the Sky Lark; they prefer to be in green crops.”

Schneelerche [Snow Bunting] (pp. 179-180): “A rare bird
and little known. It is only seen in winter, especially when
there is much snow on the ground, at the end of Decem-
ber and in January; they are only found singly and eat all
kinds of small seed-heads sticking through the snow; they
are smaller than Sky Larks, white in color with gray breast
and back, and have a peculiar call; it is supposed that these
birds breed in very cold countries, and only at certain
times, when the coldness there is very great and food
scarce, do they migrate to our region: they do not appear
every year; it is the same with the Waxwing, and we have
no other winter birds but these two species.”

Steinpicker or Steinklitsch [Northern Wheatear] (p. 181):
“It occurs in fields in the high hills, it builds its nest on
river banks or in rock crevices, also even in mole holes.
[...] According to its character it has much in common
with the Whinchat.”

Kiebitz [Northern Lapwing] (pp. 181-1 82): “Its habitat is
wet meadows and arable fields, it makes its nest on
{Kufenl^ and in clumps of rush. [. . . ] In the autumn they
gather in large flocks and get ready for migration.”

Brachvdgel [Curlew, etc.] (P. 182-184): Four (or five)
species - (1) Keulhaken [Eurasian Curlew], (2) Saatvo-
gel [European Golden Plover], (3) Dutgen [Dotterel] and
Kleiner Brachvogel [Whimbrel]. “I once saw another kind
of Brachvogel, which was no bigger than a lark and looked
almost like a White Wagtail.”

V. On waterbirds (pp. 185-199)

Schwan [Mute Swan] (p. 1 85): “It can be easily tamed and
is kept as an ornamental bird by the gentry.” It migrates
in small groups in spring and autumn, and feeds on plants.

Wilde Gdnse [wild geese] (pp. 185-187): The wild goose
is ash-gray with a white belly; it breeds in marshy places
and is a migratory bird, but some also stay here in win-
ter, when they feed on “green cereal crops and grass”.
“Their migration is wonderful to see; no other bird pays
more attention to discipline and caution than Cranes and
geese; they always fly in the form of a triangle, at whose
peak is always an old gander that is akin to the leader of
the whole army. Whoever wants to shoot them must use
all his cunning.

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Wilde Enten [wild ducks] (pp. 187-189): (1) Common
wild duck [Mallard]-, nests on old pollarded willows or
at the water’s edge; departs in autumn, though many stay
here; breeds with farmyard ducks; (2) Smaller, with a short
blue bill, head of male brown with whitish gray on the
wings [Common Pochard], (3) Grofie Kriickenente [Gar-
ganey], (4) Kleine Kriickenente [Eurasian Teal], (5) Grofie
Loffelente, tail long and pointed like that of a pheasant
[Pintail?]-, (6) Bill like that of Pintail [?] but tail short
[Shoveler]. (7) Grofie Tauchente, larger than Mallard, ash-
gray, head brown, bill and legs red, very rare [female
Goosander, Red-breasted Merganser] , (8) Kleine
Tauchente or Pfeifente [Eurasian Wigeon], “because it can
whistle just like a person does with his fingers. [. . . ] They
often dive below the surface and are skilled fish-catchers
[?].” (9) Weifie Tauchente, head and wing-tips black, very
rare [Goldeneye]. These diving ducks migrate in autumn
and return in spring.

Reiger [Grey Heron] (pp. 1 89-190): “It does not carry its
neck stretched out when flying, like the stork, but dou-
bles it over. [...] Its food is fish, frogs, and shellfish.”

Rohrdommel [Eurasian Bittern and Little Bittern] (pp.
190-191): Large and small bittern. “When the Bittern is
sitting and notices somebody approaching then it stretch-
es its bill, neck, and whole body straight upwards, with-
out making any other movement, so that it looks like a
pointed wooden post or old tree trunk. [. . . ] When it wants
to call then it puts its bill under the water, which increas-
es the volume so that it can be heard at a great distance.”
On p. 155 Naumann wrote that he had observed how the
Bittern puts its bill “under the water” when calling. “I
spent a lot of time stalking them before I was able to see
this, until once I was successful and, luckily, in bright
moonlight saw it because the bird went down to the wa-
ter.” — In size and character the Little Bittern is similar
to the larger species, and “when it sits in a tree it points
its bill and body straight upwards and so could be mis-
taken for a pointed branch”.

Mowen or Seekrdhen [gulls and terns] (p. 192): There are
three species: (1) The large Seekrahe [‘sea crow ’= Com-
mon Tern], as large as a pigeon, it is white and has a black
cap on its head, and a hooked and sharp bill; (2) Thrush-
sized with longer wings, ash-gray, and in behavior simi-
lar to the first species [Black Tern]-, (3) All white and not
so common as the previous two [Black-headed Gull, non-
breeding plumage].

Eisvogel [Common Kingfisher] (pp. 192-193): The ‘wa-
ter-woodpecker’ “always sits on the branches of trees that
hang over water and when it sees some worms in the wa-
ter it dives down into it, seizes a worm, and flies back up
to its perch on a twig or post.”

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Hurbel or Bldsse [Coot] (p. 193): The Coot is constant-
ly on the water and lives on small fish and all kinds of
water worms, etc.; they build their nests in the reeds on
the water, and move away in the autumn.

Wasserhuner [waterfowl] (pp. 193-194): 3 species with
long narrow toes: (1) The Grofie Wasserhenne [Moorhen]
with red shield above the bill, (2) very like that species
but no larger than a Quail [Little Crake/Water Rail?]. (3)
Black speckled with white, and with many white feath-
ers in the tail [Spotted Crake].

Taucher [grebes] (pp. 194-195): (1) Grofier Taucher
[Great Crested Grebe] is “more seldom seen than the
small one; they have a pointed bill; [...] they dive con-
stantly and live on fish”. (2) Kleiner Taucher [Little
Grebe] “is as big as the small Wasserhenne and they are
the commonest [grebe]”.

Wasserschnepfen [waders] (pp. 195-199): “The names are
very uncertain for in one place they are given one name,
in another place a different one. Everyone should call them
by the name that is used in their area.” [For curlews and
plovers see above under Field Birds.]

Grofie Wasserschnepfe [Black-tailed Godwit], call ‘kla-
rit’

Grofie Pfuhlschnepfe, legs olive-green [Greenshank]
Kleine Pfuhlschnepfe, coloring similar to the previous one
[Wood Sandpiper]

Similar to Dotterel but with a longer, finer bill [Common
Sandpiper]

Grofier Rotschenkel [Spotted Redshank], ash-gray and
speckled white, white rump, red legs, call ‘ta - it’
Kleiner Rotschenkel [Common Redshank], like the pre-
vious one but smaller.

Followed by various species of stints and sandpipers that
cannot be identified on the basis of the descriptions.

Bekassinen [snipe]-. (1) Ketschnepfe, Gras- or Ried-
schnepfe [Common Snipe], (2) slightly smaller [Jack
Snipe], does not fly so rapidly

VI. On tame birds

“Tame birds are what I call those that live in towns and
villages, and which prefer to be around people.”

Storch [White Stork] (p. 200): “It likes to build its nest
on buildings, especially on thatched roofs, as well as in
high trees. There is also another species of stork [Black
Stork], which is coal-black with a white belly and breast,
but it is found much less frequently.”

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Schwalben [hir undines] (pp. 200-201): The Haus- or
Stachel Schwalbe [Barn Swallow] nests in buildings and
stables. The Mehlschwalbe [House Martin] “hangs its nest
on the outside of buildings, closing it up except for a small
hole through which to enter”.

Haus- or Sallatlerche [Crested Lark] (p. 201): Slightly
larger than the Sky Lark, with a pointed crest on its head;
it has a very pleasant song, builds its nest in gardens un-
derneath the garden plants or on walls, most likely also
in thatched roofs. “It is not found everywhere, only in up-
land villages with few trees around the buildings and ad-
jacent to open fields.”

Sperlinge [sparrows] (pp. 201-202): Haussperling [House
Sparrow] and Holz- or Rohrsperling [Tree Sparrow], “the
latter is slightly smaller and has a brown head. At the end
of this book I will put both of these sparrow species on
the stage, where they will tell each other about how they
live.”

Two years after the appearance of Der Vogels teller, J. A.
Naumann published a further small book:

Der philosophische Bauer

Oder Anleitung, die Natur durch Beobachtung und Versu-
che zu erforschen (1791)

[The philosophical farmer

or instruction in the investigation of nature by observa-
tion and experiments^.

A new edition in a different format was published by
Paul Leverkiihn (1900), with an introduction and detailed
annotations by the editor.

Naumann speaks here on nature, weather forecasting,
anthropology (temperament and physiognomy), alchemy,
honeydew, ergot, mildew, on the immutability of matter
and its quantity, on chemical processes, as well as on bird
migration and birds as weather “prophets” (Siskin, Red-
poll, Fieldfare, Waxwing, Snow Bunting); to the latter ob-
servations Leverkiihn (1900) commented in detail, draw-
ing on later research.

This little book (19x11.5 cm) contains, as Baege (1980:
6) wrote, “all kinds of reflections on occurrences in na-
ture, but so peculiar were some of his explanations that
one scientist felt obliged to advise the farmer to stick to
his plow” (see Leverkiihn 1900: 63-64 and Allgemeine
Deutsche Bibliothek, Volume 110, Kiel 1792, p. 451).

Johann Matthans Bechstein (1757-1822). Bechstein
(Fig. 21) was initially a schoolteacher at the Philanthropin
in Schnepfenthal and later, while he was Director of the
Forstakademie [Academy of Forestry] in DreiBigacker
near Meiningen, the first biologist among the founders of

Bonn zoological Bulletin - Supplementum 59; 1-116

scientific forestry (Hildebrandt 1933; Mey 2003): “He was
the first man to give forestry studies a sound scientific
foundation, sending emissaries to spread his ideas rapid-
ly through the whole of Germany” (Ratzeburg 1872)f
Bechstein recognized the necessity of regarding the whole
of nature as a unified interdependent organism and saw
that: “Nature is in balance when left to herself” {Muste-

rungder Thiere 1792a, Introduction p. 4) [Review of

the animals ]. An initial propagation of the idea of

bird protection was introduced in his work. Bechstein left
an enormous life’s work in several fields at his death, but
“in all of natural history my favorite subject has always
been ornithology” (1791, Foreword, p. V). He was an ex-
perienced field ornithologist, who roamed through the
Thuringer Wald and its surroundings on numerous excur-
sions, and who constantly kept 30-60 birds of different
species in large aviaries at his home, observing their move-
ments and behavior. “I dare say that there is probably no
bird in Germany, at least not in Thiiringen, that I would
not be able to quickly identify at a good distance by its
voice and flight” (Foreword, pp. VIII-IX, footnote). Jo-
hann A. E. Goeze (1731-1793), a physico-theologian in
Quedlinburg who knew Bechstein well, wrote to him on
March 16th, 1792: “If only I could go hunting with you!
We wouldn’t want to shoot. Just observe” (L. Bechstein
19855: 32).

Just like the natural history research of his forerunners,
Bechstein’s work was also done in the light of physico-
theology. In the Foreword to Bechstein’s early work on
the Gemeinniitzige Naturgeschichte Deutschlands (Vol-
ume 1, 1789, pp. II-III) [Popular natural history of Ger-
many], C.G. Salzmann, Principal of the Philanthropin in
Schnepfenthal and a great influence on Bechstein, wrote:

“Nature is [. . . ] the book of God, which tells of the pow-
er, wisdom, and goodness of its author. [. . . ] When I watch
them [= the ‘useless’ insects] when they are on the mel-
ons and cucumbers, and I see how they powder themselves
with the pollen of the male flowers, how they fly with it
to the female flowers, and so in their fashion fertilize
them^ then I cannot help but to look up to the Almighty
with a feeling of gratitude that something unknown has
been made clear to me.

The more insights I gain in the context of the things I
see around me, then the more I come to know the Heav-
enly Father and the deeper becomes my reverence, my
love, my trust in Him.”

Bechstein himself asserts in the Foreword to his
Kurzgefafite Gemeinniitzige Naturgeschichte (1792b:
V-VI) [Shortened popular natural history ] :

“Nothing lets us better know the mighty, wise, and good
Creator [than natural history]. The works of God are - and
the wise Creator wished it so from the beginning - the
schoolmasters of mankind and we their pupils.”

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Jr 1

Fig. 21. Portrait of Johann Matthaus Bechstein after a pastel
drawing of 1803 by J.P. Bach (from Rudolstadter naturhistori-
sche Schriften 11 [2003], Original in possession of the Meinin-
ger Museen der Kulturstiftung Meiningen).

J.A. Naumann’s Vogelsteller (1789) had appeared two
years before Bechstein published the first volume of his
avian handbook in 1 791 , and the latter used passages from
Naumann’s book in several places: “Entire passages and
sentences from Naumann appear in condensed form in
Bechstein, as well as a re-engraved copy of a plate, and
all unacknowledged” (Baege 1980: 14).

Principal ornithological work. Bechstein’s Gemein-
niitzige Naturgeschichte Deutschlands nach alien drey Re-
ichen [Popular natural history of Germany covering all
three kingdoms^ appeared in four large volumes, numbers
2-4 of which dealt with birds (1791-1795) (Fig. 22; on-
line Appendix 3); a second expanded edition was pub-
lished in 1805-1809. Volume 1 is on the mammals. All
birds known to the author in the country are described in
great detail, and every species section is consistently di-
vided into field characters, description, characteristics, dis-
tribution and occurrence, food, breeding, predators, dis-
eases, usefulness, harmfulness, names, and varieties.
When Bechstein began writing his manuscript in 1786, or-
nithology in Central Europe was still in its infancy. No dis-
tinction was made between Ringed and Eittle Ringed
Plovers, Green and Grey-headed Woodpeckers, Great
Spotted and Middle Spotted Woodpeckers, or between

Goldcrest and Firecrest. Nevertheless, at the end of the
1 8th century Bechstein’s Naturgeschichte was the most
important and most complete handbook of birds, which
were presented in the modem Einnaean methodology, each
bird with a Eatin genus and species name. In Germany this
methodology had been employed in ornithological works
since the 1760s by PS. Pallas and J.R. Forster, and intro-
duced later in larger works dealing with the entire animal
kingdom by Statius Muller (from 1773) and Gmelin
(1788-1789).

Bechstein defined bird species in a purely morpholog-
ical-typological fashion: “If several such things [= indi-
viduals] have a great resemblance to each other in their
important characteristics and parts then they can be con-
sidered as the same species. If several species agree in cer-
tain principal characteristics then they constitute a genus”
(1792b, pp. 7-8). The species criterion of successful sex-
ual reproduction employed since the days of John Ray and
Georges-Eouis de Buffon and later by German zoologists
like Frisch, Blumenbach, Pallas, or Zimmermann, was
probably unknown to Bechstein, at any rate he neither used
nor discussed it. This meant that in many cases he de-
scribed individuals of one species but different plumages
under several new species names. For example, without
any further disccussion he classed the gray- and black-
backed Pied Flycatchers, which represent plumage
morphs within a variable population, as two separate
species. Similarly, he gave newly-devised Eatin names to
birds in winter, breeding, and juvenile plumages of sev-
eral species of waders, on a pure morphological-typolog-
ical basis without any comparative discussion.

In the Foreword to his Naturgeschichte, Vogel [Natur-
al history, birds]. Volume 1 (pp. VIII-IX), Bechstein
(1791) wrote:

ru hIh

•ffin

e f J Lnlili Fif hcitii!}
fuE Stijlmliiwr, QuiitnSrljntr (inS

Wf ( I y u p f f E
^ f i P I t 3-

tfk ei.jfriEP triflul.

I T 9 !■,

Fig. 22. Title page and frontispiece of Bechstein (1791-1795;
vol. 3, 1793).

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Jurgen Haffer (t) et al.

“In order to give my natural history of birds a certain truth
to the best of my ability, and to make my way as secure-
ly as possible, I have consistently and carefully studied,
in all seasons of the year, those birds that I have always
had around me (and I think I may count all the species in
Thuringen among them), have taught myself their natu-
ral history by observing them, and have described the
characters of every bird from life, taking exact account
of season, sex and age. ”

As an anonymous critical reviewer of the first bird vol-
ume of 1791 (perhaps Blasius Merrem?) remarked, the
above guideline was only partly correct, since the author
had naturally used many accounts of birds he had not per-
sonally seen from other books, “by which his own, often
excellent remarks are frequently stifled”. The same re-
viewer continued: “Everywhere apparent is a lack of crit-
ical awareness and theory of natural history. [. . . ] The au-
thor comes to grief completely in the Introduction [on
avian anatomy]; it is full of errors.” Because of the au-
thor’s verbosity and digressions “some of his own, often
important observations, and the many good aspects of this
book, are overshadowed and wasted” {Allg. Literatur-Zei-
tung, October 1792, Vol. IV, no. 265, cols. 53-56).

However, following the completion of the
Naturgeschichte the work was given a more favorable re-
ception, again anonymous, in the form of a review of the
third volume (1795). The reviewer “feels genuine grati-
tude for the immense efforts made by this ever attentive
observer. [...] The history of the birds of Germany is
brought to an end with this volume. In our judgement it
is the most important and richest ornithological work of
our time” {Allg. Literatur-Zeitung, July 1796, Vol. 3, no.
218, cols. 121-124).

Another anonymous reviewer of the first bird volume
praised “this most excellent and, when it is complete, cer-
tainly unique work of its kind”. The “wealth of the author’s
own fruitful remarks and observations often throw light
on aspects of ornithology all at once that until now had
lain in darkness”. He goes on to point out gaps in Bech-
stein’s work that he has noticed, (e.g.) there being no men-
tion of Barrington’s observations on birdsong, as well as
the fact that seabird feathers are water repellent, and “on
the whole we would wish to see laying and incubation
times given more carefully and more exactly” {Gothais-
che gelehrte Zeitungen, no. 5, pp. 41-48, 18 January
1792).

Bechstein made extensive use of earlier ornithological
literature, but recognized that “apart from Buffon’s his-
tory of birds, most excellently adapted by Professor Otto
[in 35 volumes], we in Germany possess no book that
deals with this branch of natural history in a comprehen-
sive manner” (1791, Foreword, p. X). The illustration of
the Bald Ibis, and the text, are most likely from Buffon’s
work, as Bechstein was certainly not acquainted with this

Bonn zoological Bulletin - Supplementum 59; 1-116

species. He praised J.L. Frisch as being “in the best pos-
sible sense my forerunner in the description of German
birds” (1791, Foreword, p. IX), and dealt most thorough-
ly with his birds using Linnaean methods in his Natural
History. In the second edition of the Natural History he
placed this reworking ahead of his own species accounts
so that “the lover of ornithology can experience, before
reading my description of German birds, the content and
manner of Frisch’s knowledge” and “because of the ex-
cellence of Frisch’s work I will always pay close atten-
tion to his representations in my accounts and cite them
often. As a result, it will also become clear where they
should be placed according to the system I use” (Beeh-
stein 1 805: 283, 284). Similarly, Bechstein was a great ad-
mirer of Freiherr von Pernau’s Angenehme Landlust
(1720), which he himself published in a new 10th edition*^,
and he also wanted to edit Pastor Zorn’s Petino-Theolo-
gie (1742^3) but this plan was never realized.

Illustrations. A great weakness of Bechstein’s Popular
natural history of the birds of Germany are the illustra-
tions. Around 100 small depictions show frequently stiff
and badly mounted birds, and they are equally unsatisfac-
tory in their eoloration. His Ornithologisches Taschenbuch
von und fur Deutschland (1802-1821) [Ornithological
pocketbook of and for Germany] contains only 39 illus-
trations and these “are miserable and not worth looking
at, apart from the fact that some are simply wrong” (B.
Meyer in litt. to R.H. Schinz; see Moller 2002). The lack
of good illustrations in Bechstein’s works was definitely
regarded as very annoying by his readers. In an attempt
to remedy this deficiency, Bechstein published a general
book of plates, not limited to birds, called Getreue Abbil-
dungen naturhistorischer Gegenstande in Hinsicht auf
Bechsteins kurzgefasste gemeinnutzige Naturgeschichte
des In- und Auslandes (1793-1809) [Faithful illustra-
tions of natural history objects relating to Bechstein’s
shortened popular natural history of Germany and
abroad]. But here too:

“the birds are very more badly drawn from badly mount-
ed specimens. Here the fault does not lie with the engraver
or the coloration [...] but solely with the drawings pre-
pared under the supervision of the author. [...] The illus-
trations supplied here are hardly in a position to deliver
what they are meant to, namely to compensate for the nec-
essary deficiences of a description” {Allg. Literatur-
Zeitung 1796, Vol. I, no. 46, col. 361-363). (Figs. 23a, b).

Species knowledge. Bechstein knew all the breeding birds
of Thuringen, and in his Naturgeschichte attempted to deal
also with those species which other authors had commu-
nicated from different regions, and thus to achieve com-
plete coverage of Germany. However he encountered dif-
ficulties with the wading birds, describing the Juvenile,

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breeding, and winter plumages of several species under
different species names.

New descriptions. Bechstein was a field ornithologist and
not a systematist, but he did occupy himself with prob-
lems of systematics, allocating and publishing many Lin-
naean species names (consisting of a genus name plus spe-
cific name) for birds which, as it seemed to him, had not
yet been given names using this methodology. To be able
to form an opinion and judge the value of Bechstein’s ac-
tivity in the field of taxonomy it is no longer sufficient to
simply “cherry-pick” those names that later had their va-
lidity confirmed (Tittel 2002; Mey 2003). Instead, in his-
torical works all names given by Bechstein should be in-
cluded. After all, in this sense Bechstein himself had al-
located and accepted names that are currently both valid
and invalid; he held them all to be equally “valid”. More-
over, we ought to differentiate between (a) names which
Bechstein gave to birds he knew from life or as skins he
was able to examine and compare, and (b) those names
he attached to a taxon already described in the literature
but named only in English, in other words birds that he
knew neither from life nor as skins. Bechstein described
a total of around 170 bird “species” as new. His assump-
tion, or “hope”, that the validity of these names would be

assured, was to be disappointed, for only very few of his
designations were valid and became established.

(\) European species and subspecies (see Mey 2003 and
Haffer 2006, Anhang [Appendix] 4): Of the almost 100
new “species” names of European birds given by Bech-
stein only 5(!) refer to genuinely new and at the time un-
known species that he had discovered in Thiiringen: Marsh
Warbler, Collared FlycatcheE, Red-breasted Flycatcher,
Wood Warbler, and Barred Warbler. The Rock Partridge,
Barnacle Goose, and Thrush Nightingale that he also
named had been known to J.E. Frisch in the pre-Einnaean
period, who depicted them in very good color plates. The
Marsh Sandpiper had been described by Naumann father
and son in 1799 under the non-Einnaean name Scolopax
Glottis minor\ also excellently illustrated in color. Hence
this species too was not really “new” when Bechstein gave
it a Einnaean name in 1803, depicting it in a poor color
plate following his collection of the bird as it passed
through on migration. The Naumanns had also already de-
scribed the North American Upland Sandpiper in 1 8 1 1 un-
der the name Tringa macroura (which did not become
valid), therefore one year before Bechstein’s naming of it
as Tringa longicauda, based on Eatham’s published Eng-
lish description. Bechstein described the White-backed
Woodpecker in 1 802 from a color illustration sent to him

a

b

%6xm{

Figs 23a, b. Two plates from Bechstein (1791-1795; vol. 4, 1795, from photocopies by J. Haffer). 23a; female Blackcap (top),
Garden Warbler; 23b; Eurasian Stonechat (top), Icterine Warbler.

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Jurgen Haffer (t) et al.

from Silesia. The “discovery” of the Parrot Crossbill was
occasionally attributed to J.M. Bechstein in his Ornithol-
ogisches Taschenbuch (vol. 1, p. 106, 1802), as Hartert
(1904: 122) noted. Yet here too, B.C. Otto (1778) had
clearly differentiated this species from Common Cross-
bill as “Tannenpapagei” [Fir Parrot], but he did not give
the species its own Linnaean name, which was done by
Borkhausen in 1793: Loxia Pytyopsittacus .

While almost all the scientific names given by Bech-
stein were applied to species that were unknown to him
at the time of their description, most of these birds had in
fact already been allocated Linnaean names by other au-
thors. Bechstein’ s names for them were “redundant” syn-
onyms, an unnecessary burden on nomenclature. A simi-
lar unfortunate balance between valid and invalid names
exists in the work of other naturalists of the time, who all
gave scientific (Linnaean) names (with the additional mi-
hi = “by me”) to birds and other animals that appeared new
to them, in the assumption or hope that the name would
later be validated, securing “immortality” for the author’s
name. But we must pardon these early authorities by re-
minding ourselves that (a) due to the difficulty of com-
munication and the small print runs of many books they
were often poorly informed of current research, and (b)
their inadequate knowledge of various plumages and molt
stages in some species meant that new names were often
given to differently plumaged males, females, or juveniles
of the same species. Despite this, the impression remains
that Bechstein, as well as some subsequent authors in the
19th century, distributed “new” scientific bird names with
a certain recklessness (or mihi addiction!). Considering his
great experience, Bechstein was clearly aware of the strik-
ing molt in some species and therefore understood the ne-
cessity of careful comparisons before it was reasonable
to allocate a new name. Possibly one of the reasons why
he chose to translate the works of Latham and Levaillant
in particular (see below) was that they were among the
authors who rejected the Linnaean nomenclature, thus af-
fording their translator the opportunity of giving scientif-
ic names to the species newly described in their books, in
which only common names were printed. It can be as-
sumed, though it is not apparent, that Bechstein knew the
differences between the songbirds genuinely discovered
by him, and properly described with song, etc., and the
many “species” to which he assigned his (invalid) Lin-
naean names after examining one or two skins, or on the
basis of descriptions by other researchers, without any fur-
ther knowledge of their status.

Bechstein’ s genus names for European birds that are still
valid are Cygnus (1803), Crex (1803), Anthus (1805), Cal-
carius (1802), and Saxicola (1802). Several of his remain-
ing genus names were rejected by later authorities (Haf-
fer 2006: Appendix 4). The situation with novel genus
names was unlike that of new species names in that he was
not dealing with previously unknown birds (unless a new

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species was placed at the same time in a new monotypic
genus), but simply with a new grouping of already famil-
iar species. But also here, with regard to the pipit genus
Anthus, it should be noted that father and son Naumann
(1798) had already separated the larks and pipits as groups,
though without giving the latter their own Linnaean genus
name.

(2) Non-European species and subspecies'. In the years
1793-1812 Bechstein translated the great work of the
British ornithologist John Latham, A General Synopsis of
Birds (1781-1785), into German, adding many comments
of his own. Latham was one of the ornithologists who
thought that the new nomenclature was superfluous and
consequently refused to give Linnaean names to any
known species or any species newly described by them-
selves, using English names only. Their discoveries were
later appropriated by other authors, who, by supplying
these new species with their own scientific names, were
“adorning themselves with borrowed plumes”. One of
these was Bechstein, who, during his translation activi-
ties, allotted scientific names to ca. 80 bird “species” that
were described for the first time in English or French in
works by Eatham, Eevaillant, and Vieillot. The majority
of Bechstein’s names were also (invalid) synonyms, but
some have survived as valid species or subspecies names
(see Peters’s Check-list of the Birds of the World, the valid
taxon is printed in bold in the following list):

Coragyps atratus (Bechstein, 1793)

Bartramia longicauda (Bechstein, 1812)

Numenius americanus (Bechstein, 1812)

Eos squamata riciniata (Bechstein, 1811)

Trichoglossus haematodus capistratus (Bechstein, 1811)
Psittacula krameri manillensis (Bechstein, 1811)

Ara ambiguus (Bechstein, 1811)

Ara tricolor (Bechstein, 1811)

Geoffroyus geoffroyi (Bechstein, 1811)

Upupa epops africana (Bechstein, 1811)

Dicrurus adsimilis (Bechstein, 1794)

Phylidonyris niger (Bechstein, 1811)

Sialia currucoides (Bechstein, 1798)

Spizella passerina (Bechstein, 1798)

Among the birds on this list, Bechstein described the par-
rot Psittacula krameri manillensis from a cage bird, per-
haps one he had seen in the collection of rare birds be-
longing to Herzog [Duke] Georg von Meiningen. He knew
the others only from descriptions mainly by Eatham, at-
taching his own new Einnaean names to them. Ornitho-
logically speaking, these names are then something com-
pletely different from those he gave to the five bird species
which he himself had discovered in Thilringen (see above).

Birds have been named in honor of Bechstein only
twice, but these names are no longer valid: (1) Turdus
bechsteinii J.F. Naumann, 1822 [= T. ruficollis Pallas,

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1776] and (2) Tringa stagnatilis bechsteini Zarudny &
Smirnov, 1923 [= T. stagnatilis, monotypic].

Reviews of the publications. Although Bechstein was one
of the most important Central European ornithologists of
the early period, and despite the fact that much has been
written on his life and his work as a pioneer of scientific
forestry, a thorough analysis of his contribution to ornithol-
ogy has yet to be undertaken, nor will it be attempted by
us here. Mauersberger (1990) has made some comments
in this regard indicating the type of approach that should
be adopted in this case (and in those of other early or-
nithologists): (1) determination of the degree to which
Bechstein recorded the number of species in Central Eu-
rope and clarified their taxonomy; (2) his sources in the
literature; (3) the influence of his work on future authors;
(4) a comparison of his knowledge of breeding biology
and ecology with that of earlier, contemporary, and later
researchers, and (5) his field ornithological methods, es-
pecially with respect to his Ornithologisches Taschenbuch.
Only after the application of such a comparative analysis
will it be possible to recognize which aspects of ornitho-
logical progress in Central Europe follow from Bechstein’s
work.

In his Naturgeschichte, Bechstein generally gave de-
scriptions of a large number of known, but also of then
unknown facts relating to European birds, but usually
without subjecting them to any kind of biological inter-
pretation, as before him Pemau, and Zorn in particular,
had so diligently done. In the general chapter on birds in
his Naturgeschichte there are the beginnings of such an
approach in his function-related terminology of types of
foot structure, and in the pointers to the relations between
wing length and breast muscle weight or intestine length
and diet, as well as species-specific nest materials and con-
struction (Mauersberger 1980, 1990), but the relations be-
tween structure, movement, and food choice are not yet
grasped. See also Mey (2010) for a good overview.

On his ornithological excursions in Thtiringen Bechstein
discovered five European bird species (see above). This
illustrates Just how sharp his fieldcraft and critical attitude
were. He had a wealth of knowledge on German birds,
which he brought together to great effect in his 3 -volume
Naturgeschichte, to the advantage of the progress of or-
nithology. However, of the total of 170 Einnaean species
names which he introduced into the literature, very few
attained permanent validity. As early as 1 805, the expe-
rienced Herr von Minckwitz wrote to J.F. Naumann: “In
my opinion, many birds in Bechstein’s Taschenbuch are
presented as true species which are nothing of the kind”
(Thomsen & Stresemann 1957: 56), which was confirmed
in the reply from J.A. Naumann: “After my experience
with Bechstein’s Orn. Taschenbuch I would just delete a
good few of the species” (see Thomsen 1930: 13). Of the
names which Bechstein gave the species discovered by

Bonn zoological Bulletin - Supplementum 59: 1-116

Eatham, and allocated only English names by that author,
some remain valid. But these are purely “administrative”
names, given within the framework of the Einnaean
method to species discovered by another ornithologist, and
therefore to be regarded in a completely different light
from the new species discovered by Bechstein himself in
the field.

In 1848, looking back over many decades, C.E. Brehm
published a harsh critique of Bechstein:

“Bechstein was a friend of my late father and used to live
just two hours away from my place of birth. I respected
him greatly, but when as a nine-year-old boy I read the
erroneous description of the Blackbird ’s nest he lost all
credibility for me. Many other things that 1 recognized as
false in my youth, e.g. the early breeding of the Magpie
and the Dipper (both of which were said to build in Feb-
ruary), the completely mistaken description of the King-
fisher ’s nest, and so on, put me off the writings of this great
man to such an extent that I simply couldn ’t read them any
longer” (Thomsen & Stresemann 1957: 147).

Stresemann (1951: 296) considered Bechstein’s Gemein-
nutzige Naturgeschichte to be “nothing but [....] a rigor-
ously classified collection of material, though unprece-
dentedly rich in detail”. When E. Mey (2003: 70) writes
that this judgement is “unjusf’ since it implies that Bech-
stein could be “at best regarded as mainly an industrious
compiler” then this results from a misunderstanding. An
“industrious compiler” is one who brings together already
familiar information scattered throughout the literature.
But this is not what Stresemann intended. He meant that
in its day the Naturgeschichte as a collection of facts was
unrivalled in its completeness, that is it dealt with the
known and many (until then) unknown facts regarding the
most diverse aspects of birds. But, continues Stresemann,
Bechstein - as opposed to Pernau and Zorn - was very
reluctant to draw conclusions from these facts and to in-
terpret some of them with regard to their biological sig-
nificance. See especially Mey (2003, 2010) for a very ex-
tensive appraisal of Bechstein.

Biologists (= physiologists) at that time saw themselves
(in today’s language) as “causation investigators”, con-
cerned with the immediate physical-chemical,
(proximate) causes or explanations of biological phenom-
ena (an enterprise that lacked even the fundamental con-
cepts in Bechstein’s day), while the physico- or natural the-
ologians emphasized the fitness for purpose (design) of
many structures, interpreting them teleologically, so
were concerned with their historical (ultimate) explana-
tions. As we now know, every biological phenomenon has
functional as well as historical explanations, so that an an-
tithesis between the “causation investigators” and the tele-
ologists, as referred to by Stresemann, did not actually ex-
ist in retrospect. Yet because of this supposed conflict

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Jurgen Haffer (t) et al.

many naturalists around 1 800 were reluctant to express
their interpretations either in the one direction (function-
al) or the other (historical). Regarding his explanations of
biological structures and avian phenomena, Bechstein
might be placed within this group of naturalists.

However, in 1795 the bird-keeper Bechstein suggested
in his Naturgeschichte der Stubenvdgel [English transla-
tion of 1 838: The Natural History of Cage Birds] that in-
dividual differences in the quality of song were important
in mate choice as he had noticed that female canaries pre-
ferred the males with the best voices (Birkhead & van
Balen 2008: 285).

Johann Andreas Nanmann and Johann Friedrich Nan-
mann (1795-1817). In a 4-volume handbook of the birds
of northern Germany, published by Johann Andreas Nau-
mann and his eldest son Johann Friedrich between 1795
and 1803 (with 8 supplements from 1804 to 1817), all
species treated are, for the first time, illustrated in a total
of 234 outstanding color plates, almost all of them by the
younger Naumann, initially in folio and after 1 804 in oc-
tavo format. The complete title of the work was at first

Ausfuhrliche Beschreibung alter Wald-, Feld- und
Wasser-Vdgel, welche sich in den Anhaltinischen Furs-
tenthumern und einigen umliegenden Gegenden aufhal-
ten und durchziehen

{Detailed description of all woodland, field, and water
birds which live in and migrate through the Principali-
ties of Anhalt and some surrounding regions].

With the fourth instalment of the first volume (1797) this
title was altered to:

Naturgeschichte der Land- und Wasser-Vdgel des nord-
lichen Deutschlands und angranzender Lander,
nach eignen Erfahrungen entworfen, und nach dem Le-
ben gezeichnet

{Natural history of land and water birds of northern Ger-
many and adjacent countries, from our own observations
and drawn from life].

AU^fubrltdii

e I'f ^ r c i bun ^

isfkc

raclttif fid'

ijT bm Sfn&£iUi|‘c5)cti Surfctitl^umfrti

llfl6

r

£(in&^un&a8a|fcv=Si>i){jcl

11 a ( i 3 H t n

cticwurfcti t tctcrt

l 9 tn

Qlit i

itipjis, I 79 f.

Utit fn toirmlfflsn frfij0riftiti(ti

QSierter

SBU

I So?-

3fuf fluften

un^ iH Scmmifjlpii bci 3.

J

Figs 24a, b. Title pages of J.A. and Johann Friedrich Naumann (1795 [vol. 1, part 1] & 1803 [vol. 4]). Note change of title after
vol 1, part 4.

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The physico-theological concept informing Johann An-
dreas Naumann’s study of nature, and of birds in partic-
ular, is expressed in the words with which he opened his
Naturgeschichte (1795-1817):

“From time immemorial, and especially in the present en-
lightened era, knowledge of fairest nature has been the
study that has preached to us of the mighty Creator of the
World in his greatness and omnipotence. From the worm
to Man, everything reveals to us His omnipotence and
greatness. Man would still be too weak, even if he had a
thousand tongues, to relate and praise the wonders that
surround him in Creation. All the elements praise the glo-
ry of the Almighty. The denizens of the air, the birds, are
the object with which I intend to entertain my readers and
to encourage them in praise of the Almighty” (beginning
of the Preface, 1795).

From their earliest youth, his three sons, Johann Friedrich,
Carl Andreas, and Gottfried Leberecht accompanied their
father on his excursions in the country around Ziebigk ne-
ar Kothen, learning from him - a “huntsman grown gray
in the hunting and trapping of birds” - an intense study
of nature and a religiously-founded wonder in all living
things. Immediately following the Naturgeschichte of
Bechstein, the Naturgeschichte der Land- und Wasser-Vd-
gel des ndrdlichen Deutschlands (1795-1817) appeared,
with 192 excellent color plates of German birds in folio
format, done in the style of the famous bird publications
of J.L. Frisch. Almost all plates were by Johann Friedrich
Naumann, who was then just 15 years old (he had in fact
drawn the first of the illustrations at the age of 14). (Figs.
25a-d). His artistic progress can be observed in the ear-
ly folio plates. In 1804 the Naumanns changed the plate
format from folio to the smaller octavo, so the son engra-
ved all the 1 92 folio plates again in the smaller format.
The species texts in the first two volumes of this Natur-
geschichte were kept very brief, but the treatment was con-
siderably expanded when Johann Friedrich began to co-
operate on their production from volume 3 onwards (as
he mentioned in a letter; see Thomsen 1930: 15), though
he is credited as co-author on the title page only from the
fourth instalment (1803) of volume 4. The texts of the
work are based solely on data collected in the field by the
Naumanns themselves, and by Naumann senior in his
aviary directly adjoining his sitting room. Using these cap-
tive birds he was able to calculate the lengths of their mi-
gration periods in autumn on the basis of their migratory
restlessness, or Zugunruhe (1795: 4, 8-9), deducing from
his observations that many of them must overwinter in tro-
pical Africa, which was completely unknown at the time
(Haffer 2006: 28).

“Adjoining my sitting room I have built a little room
on the east side, with a window facing south covered on-
ly with wire mesh so that it is open to the air in spring,

Bonn zoological Bulletin - Supplementum 59: 1-116

summer, and autumn, but in the winter a pane of glass is
fitted into it. There is a door into my sitting room made
only of wire mesh and paper, so that I can let heat into
that room when necessary and at the same time can hear
what sort of movements the birds are making, day and
night, since I have 30 or 40 birds of many different kinds.
I keep some to observe their habits, others to hear their
attractive song, and others again for use on the trapping
ground: in other words I have forest, field, and water birds
in there. I can note the exact time of their nocturnal move-
ments because as long as their instinct is active, at night
when the moon shines, they fly around in their room, al-
so when the stars are visible in a clear sky” (1795: 4).

“According to my calculation, they can [= summer vis-
itors in autumn] make this journey inside a month; how-
ever I have noticed that in their little room the flycatch-
ers and Golden Orioles ‘migrate’ until the middle of No-
vember, because they are restless until that time, but af-
ter then become completely still. That Golden Orioles and
flycatchers, as well as swallows and other birds, migrate
to Africa must be regarded as the truth, since these birds
live on insects alone and cannot abide the cold here”
(1795: 8-9).

“They [= Golden Orioles] became very restless when
the migration period started, flying around in the cham-
ber all night into November. From this one can deduce that
this bird migrates a great distance, probably as far as
Africa. In Febmary they started their molt and looked a
sorry sight. I had to take good care of them because in the
past some had died at this time. But as soon as they had
come through they grew very lively and started to sing,
but in March they were restless again at night” (1795:
196).

“I have often had one of these ‘decoy birds’ [= Pied Fly-
catcher] in the chamber and got it used to eating some all-
purpose food to keep it alive through the winter, in order
to note how long it migrates, and I have found that it is
restless until the middle of November, but then becomes
quiet again, from which one can conclude that they must
migrate just as far as the Golden Oriole” (1797: 203).

Naumann senior reported numerous observations on the
behavior of these birds, some of which are given here to
illustrate just what an able ornithologist he was.

He wrote on the nest-building of the Golden Oriole {Ori-
olus oriolus)'. “It is a real pleasure to watch an Oriole
building its nest, and I did this often and gladly. Male and
female perform this task jointly without being in each oth-
er’s way. When one of them arrives with a long strand of
wool or a dry blade of grass, it attaches first one end of
it on the twig with its saliva and then takes the other dan-
gling end and flies with it around the twig, fastening the
nest to it so thoroughly that one is unable to detach it with-
out either breaking the twig or tearing the nest apart” (vol.
1, 1797: 195-196).

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Figs 25a-d. Four plates from J.A. & J.F. Naumann (1795-1817). 25a: male and female Garganey; 25b: male and female Com-
mon Kestrel; 25c: Eurasian Treecreeper (top), Eurasian Nuthatch (center), Wryneck; 25d: Aquatic Warbler, Sedge Warbler (top),
Firecrest (center). Winter Wren, Goldcrest. Note also eggs.

A hand-raised Stone-curlew {Burhinus oedicnemus) as I am in the room. When I enter, the bird approaches

“has become so used to me that it sits at my feet as long me joyfully, lowers its bill towards the floor, spreads its

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wings and tail, and softly calls ‘dick, dick, dick’. When
it is standing in the sun or in some other comfortable place
and someone intereferes with it, then it makes a loud snor-
ing sound to show its displeasure. Its screaming voice is
heard only at dusk and dawn. It feeds just as happily at
its bowl at night, when a light or the moon is shining, as
during the day” (vol. 2, 1799: 74).

In order to discover whether individual Common Buz-
zards (Buteo buteo) - which have very variable plumages

- retain their own particular coloration pattern in succes-
sive years or molt to a different one, he invented the band-
ing experiment: “In my last years of collecting birds I tried
to get to the bottom of this difficult question, so I left alive
all those buzzards I caught and fastened a copper ring
round their tarsus, on which my name, place of residence,
date of capture, and a letter to indicate their plumage type
were engraved. Then I set them free hoping to catch them
again some time later and perhaps in a different plumage
type. Even though I did this with a large number of birds
I never again saw a single one of them. All the bands that
were sent to me from the neighborhood were from birds
I had banded Just a short time previously” (vol. 4, 1803:
212; see also Thomsen & Stresemann 1957: 36).

Johann Andreas Naumann rediscovered the fact that
long-billed waders are capable of opening only the tip of
the bill when necessary (vol. 3, p. XXI, 1799; see also
Frisch in his Woodcock account above), and was shocked
when this was doubted by a reviewer of his book (sup-
plement 2, 1 805: 57). His son was his co-author after vol-
ume 3 but was solely responsible for the supplement texts
because his father’s eyesight had become so bad.

Hybridization of Carrion and Hooded Crows, and the
breeding and molting periods of many species are de-
scribed in the text, as is the separation of pipits and larks

- the former dips the tail up and down like the wagtails
(1798: 47). The differences between several similar Acro-
cephalus species are outlined, illustrated by Johann
Friedrich’s magnificent color plates.

Hildebrandt (2007) has done a great service by collect-
ing many faunistically and biologically interesting pas-
sages on all the species in the work of the Naumanns, to-
gether with a wealth of useful notes and several of J.F.
Naumann’s color plates, thus making these important pub-
lications easily accessible once again.

Species knowledge. The work contains descriptions and
illustrations of 285 taxa recognized today as species or
subspecies. Those missing in the main text are Wood War-
bler, Grey Wagtail, Firecrest, Bearded Reedling, and Pen-
duline Tit, but all are dealt with in the supplementary vol-
umes together with other species. Black and Red Kites are
acknowledged as separate species, but Hen, Pallid, and
Montagu’s Harriers are lumped together under the species
name Halbweyhe [“Semi-Harrier”]. The color variants of
Common Buzzard are treated as such, and banding exper-

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iments are devised to test their constancy (see above). That
there is only one species of Blackbird, and not two, is set
down here, as well as that there is only one species of Grey
Heron. Juvenile Oidemia nigra [= Melanitta nigra] and
Colymbus septentrionalis [= Gavia stellata] are thought
to be species in their own right in the principal text, but
this too is corrected in the supplements. Today’s Acro-
cephalus, Sylvia, and Phylloscopus warblers are correct-
ly differentiated. The two Regidus species and Certhia
species are not yet distinguished. Shore Lark, Crag Mar-
tin, Water Pipit, Alpine Accentor, Savi’s Warbler, Bonel-
li’s Warbler, Collared Flycatcher, Red-breasted Flycatch-
er, Citril Finch, European Serin, and both choughs are al-
so absent. However the Naumanns were dealing here on-
ly with the birds of northern Germany and hence made
no great effort at completeness by including species from
the south of the country or the Alps. Problems with the
reed warblers are discussed in the fourth supplement
(1811) and the genus Acrocephalus (“pointed head”) erect-
ed.

Systematics. The classification of birds was once again,
as in the Vogelsteller of 1789, basically ecological
{“woodland, field, and water birds' f and therefore “old-
fashioned”, for even Ray (1676, 1678) with Willughby had
broken with this tradition and had used structural charac-
ters to divide birds into groups. The Naumanns differen-
tiated a total of 30 groups (“Classes”) of birds in their ge-
ographic region. However, by splitting the pipits from the
larks for the first time on the basis of the former’s tail-
wagging they were employing a behavioral character (very
“modem” in this publication) in the systematic classifi-
cation of a bird grouping. Along the same lines as the Nau-
manns, Bechstein (1805) later introduced the genus name
Anthus for the pipits. The following list documents the
Naumanns’ divisions of the birds of Central Europe, in
which the species dealt with in the eight supplements are
placed in their appropriate group:

I. Woodland birds that have conical bills, eat seeds, and
dehusk the seeds: House, Tree, Rock, and Snow Sparrows,
Chaffinch, Brambling, Greenfinch, Linnet, Twite.

II. Woodland birds that have conical bills with pointed tips,
though not circular in cross-section but vertically flattened;
forage on trees and tall plants, hanging with sharp claws
on seed-heads: Goldfinch, Siskin, Redpoll.

III. Thick-billed birds that forage in trees but also walk
and hop: Hawfinch, Pine Grosbeak, Bullfinch, Common
Crossbill, Parrot Crossbill, Common Rosefinch.

IV. Buntings, upper mandible with ridges in the hard palate
and smaller than the lower mandible. Seeds are cut in the
bill and dehusked. Usually walk but sometimes hop: Com,

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Reed, Lapland, and Snow Buntings, Yellowhammer, Or-
tolan, Dunnock.

V. Pigeons and doves: Wood Pigeon, Stock Dove, Turtle
Dove.

VI. Woodland game-birds: short, curved bill, feed on buds,
plants, berries, larvae, worms, etc, seeds; walk and nev-
er hop: Capercaillie, Black Grouse, Hazel Grouse, Pheas-
ant.

VII. Woodland birds that feed on insects and larvae etc.
in trees, also seeds. Bill small, hard, wedge-shaped, claws
sharp for climbing and hanging, always hop. Useful birds
that eat many insect pests and their eggs: Great, Blue,
Azure, Marsh, Crested, Long-tailed, and Penduline Tits,
Bearded Reedling.

VIII . Woodpeckers: Black, Green, Grey-headed, Greater
Spotted, Middle Spotted, Lesser Spotted, White-backed,
and Three-toed Woodpeckers, Wryneck, Nuthatch, Wall-
creeper, and treecreeper.

IX. Juniper birds: hop, like to eat juniper berries: Mistle,
Song, and Rock Thrushes, Dusky Thrush (subspecies eu-
nomus). Fieldfare, Redwing, Blackbird, Ring Ouzel,
Waxwing.

X. Woodland birds that feed on insects, larvae, worms, etc.
and berries: Barred and Garden Warblers, Blackcap, Com-
mon Whitethroat, Lesser Whitethroat, Willow and Wood
Warblers, Chiffchaff, Nightingale, Thrush Nightingale,
Robin, Bluethroat, Common and Black Redstarts.

XI. Woodland birds that live on earthworms and insects,
bill elongated and pointed, walk on the ground (do not
hop): Common Starling, Rosy Starling, Hoopoe, Bee-
eater, Grey, Yellow, and PiedAVhite Wagtails.

XII . Woodland birds, flycatchers and berry eaters, bill
pointed and broad at the base. They sit still and wait for
prey: Golden Oriole, Icterine Warbler, Spotted Flycatch-
er, Pied Flycatcher.

XIII. Swallows, bill broad, catch their prey in flight: Com-
mon Swift, House and Sand Martins, Barn Swallow,
Nightjar.

XIV. Woodland birds that eat only worms, larvae, insects,
etc., but not berries; hop only: Cuckoo, Great and Eurasian
Reed Warblers, Marsh, Aquatic, Sedge, River, and
Grasshopper Warblers, Wren, Fire-/Goldcrest, Northern
Wheatear, Whinchat, Stonechat.

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XV. Game-birds, eat seeds, green plants, worms, larvae,
etc., never perch in trees: Great Bustard, Macqueen’s Bus-
tard, Common Crane, Black-bellied Sandgrouse, Grey
Partridge, Ptarmigan, Quail, Com Crake, Collared Prat-
incole.

XVI. Larks, farmland birds that feed on seeds, green
shoots of crops and other plants, with small hard bills.
Walk rather slowly: Sky, Wood, and Crested Larks, Snow
Bunting.

XVII. Larks with thin bills, feed on worms, insects, etc.,
not seeds, move their tails like a wagtail; they form a tran-
sition from the larks to the wagtails: Tawny, Meadow, and
Tree Pipits.

XVIII . Farmland birds that are snipe-like, walk slowly:
Curlew, Golden Plover, Dotterel.

XIX. Plovers, form a transition from curlews to snipes and
sandpipers: Lapwing, Grey Plover, Turnstone, Common
Ringed/Little Ringed Plover.

XX. Snipes, bill long, soft, pliable, bill tip can be opened
separately: Eurasian Woodcock, Great, Common, and Jack
Snipes.

XXI. Water snipes [waders], long-legged, walk slowly:
Curlew, Bar-tailed Godwit, Greenshank, Spotted Red-
shank, Common Redshank, Black-tailed Godwit, Black-
winged Stilt, Ruff, Red Knot, Sanderling,
[Schwarzbrauner Sandlaufer] Wood Sandpiper?, Marsh,
Curlew, Green, Common, and Broad-billed Sandpipers,
Dunlin, Eittle Stint, Red-necked Phalarope.

XXII. Herons and storks: White and Black Storks, Grey,
Purple, Night, and Squacco Herons, Great White Egret,
Eittle Egret, Spoonbill, Eurasian Bittern, Eittle Bittern,
Glossy Ibis.

XXIII. Waterhens, feed on water plants and water insects:
Moorhen, Coot, Water Rail, Spotted Crake, Eittle/Baillon’s
Crake.

XXIV. Gulls and terns: Black-headed, Eittle, Common,
Glaucous, and Eesser Black-backed Gulls, [Kleine bunte
Mevel\ Kittiwake, Arctic Skua, Caspian, Common, Eit-
tle, and Black Terns. In supplements: White Pelican, Great
Cormorant, Northern Gannet.

XXV. Swans, geese, and ducks: Mute Swan, Whooper
Swan, Common and Ruddy Shelducks, Grey, Bean, Brent,
White-fronted, Snow, and Barnacle Geese, Mallard,
Gadwall, Garganey, Eurasian Teal, Shoveler, Wigeon, Pin-

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tail, Long-tailed Duck, Harlequin Duck, [Grofie
Tauchente] Goosander/Red-breasted Merganser ?, Com-
mon Eider, Goldeneye, Tufted Duck, Pochard, Ferrugi-
nous Duck, Greater Scaup, Common and Velvet Scoters,
Red-crested Pochard, Goosander, Red-breasted Mer-
ganser, Smew.

XXVI. '' Divers’’’ \ Common Guillemot, Black Guillemot,
Puffin, Little Auk, [Gesprenkelter Seetaucher] Yellow-
billed Loon ?, Great Northern, Red-throated, and Black-
throated Loons, Great Crested, Red-necked, Black-
necked, and Little Grebes, [Schwarzbrauner Taucher]
Slavonian Grebe ?, Kingfisher, Dipper.

XXVII . "'Semi-raptors'”'. Raven and crows: Common
Raven, Carrion and Hooded Crows, Rook, Western Jack-
daw, Magpie, Eurasian Jay, Nutcracker, European Roller.

XXVIII. Shrikes'. Transition species from semi-raptors to
the raptors proper. Great Grey, Lesser Grey, Woodchat, and
Red-backed Shrikes.

XXIX. Raptors'. Black Vulture, Griffon Vulture, [Gemein-
er Adler] Black Kite ?, White-tailed, Golden, Short-toed,
and Lesser Spotted Eagles, Osprey, Gyrfalcon, Saker,
Peregrine, and Red-footed Falcons, \Blaufalkel], Hobby,
Merlin, Common Kestrel, Goshawk, Sparrowhawk,
[Halbweyhe = Hen, Montagu’s, and Pallid Harriers],
Marsh Harrier, Red Kite, Common, Rough-legged, and
Honey Buzzards, [Eulenfalkel].

XXX. Chvls'. Eagle, Snowy, Long-eared, Scops, Short-
eared, Tawny, Bam, Little, Tengmalm’s, Pygmy, Ural, and
Hawk Owls.

Friedrich Tiedemann (1810, 1814). The first textbook
of general ornithology after J. H. Zorn’s Petino-Theolo-
gie (1742-1743) was written by Friedrich Tiedemann, Pro-
fessor of Anatomy and Physiology at the universities of
Landshut and, from 1816, Heidelberg. He published ma-
jor works on the anatomy of fish, reptiles, amphibians, and
birds, later working on the physiology of the digestive sys-
tem of mammals. He was no field ornithologist, but at the
start of his career he brought out a detailed und

Naturgeschichte der Vogel (1810, 1814) [Anatomy and
natural history of birds] in two volumes as part of an un-
finished zoological textbook. The first volume on the
anatomy of birds was based on his own extensive stud-
ies, the second was a detailed representation of the life and
distribution of birds gleaned from all of the German and
international literature (“Reproduction and growth”,
“Metamorphosis of birds”, “Occurrence and distribution”,
and “Movements of birds”). In this work Tiedemann quot-
ed widely from the contributions to the natural history of
birds, nests, nest building, etc. by Zorn, Derham, and oth-

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er 18th-century physico-theologians, and also included
much from recent ornithological literature on European
and tropical birds (Stresemann 1951: 297-303).

Like Zom 70 years before, Tiedemann described the
purposiveness (Zweckmdfigkeit) of avian body stmctures,
concluding that “the entire anatomy of the bird is organ-
ized for flight. The head is small, light, and ends in a more
or less pointed bill, a form that makes cutting through the
air very much easier” (p. 348). During flight the wings act
like paddles, sails, or parachutes, the tail like the rudder
of a ship. A muscle contracts the toes when the knee joint
is flexed thereby fastening the sleeping bird to the branch.
“When sleeping, birds mostly put the head under one
wing, so that the line of the center of gravity passes be-
tween the feef ’ (p. 365).

Birds breed once or several times in spring, earlier in
southern Europe than farther north, but the crossbill breeds
in winter and tropical birds have completely different
breeding periods. Many males have a breeding plumage,
sing, and perform a courtship display. Nest types, sites,
and construction are described, as are tropical communal
nests. The form of the nests of Dipper, Wren, and Long-
tailed Tit are presented, using the descriptions of Zom,
Derham, and Bechstein. The nest-building instinct is in-
nate: “We must suppose that birds are driven by a blind,
innate compulsion and determination to build their nests,
but to explain this is still outside the compass of our
knowledge” (p. 60; with reference to H.S. Reimams 1760).
Tiedemann knew that smaller birds in South America lay
fewer eggs (2 to a maximum of 4) than those in Europe,
and that young females lay fewer eggs than older ones of
the same species. Birds that breed several times in a year
lay fewer eggs in later clutches than in the first clutch.
“The degree of development of birds at hatching depends
on their way of life and where they live” (p. 75). Ground-
nesting species have relatively large eggs and the young
are relatively advanced in their development at hatching,
while the young that remain long in the nest are born
naked and helpless. The “shape of eggs before incubation”
and “malformed eggs” are discussed, and the section “On
breeding” supplies information on the respective share in
incubation of females and males. The (designed, since pro-
tective) covering of the eggs when temporarily leaving the
nest was well known to the author from Moorhen, Caper-
caillie, ducks, geese, and swans, as was the similarly pur-
posive “distracting” display in species with an open nest
(after Zorn). The varying number of eggs in the nest of
different species and the differing incubation periods are
illustrated in tables (pp. 63-72 and 138-139). The length
of incubation depends on the stage of development the
young are in on hatching, hence that of nidicolous song-
birds is relatively short, that of nidifugous species rela-
tively long.

The section on the “Metamorphosis of birds” from
hatching to death deals with the rearing of young by their

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parents, growth, molt, song development, etc.: “Young
birds learn [....] the actual song successively, mostly
through imitation of the song of their parents. [....] When
young songbirds are removed from their parents they can
easily learn the song of other birds with which they might
come together” (pp. 288-289). Juvenile plumage often re-
sembles that of females; molting waterbirds “often lose
all their flight feathers simultaneously, so have to hide
themselves in the reeds”, and the color of the winter
plumage of many birds is often quite different from the
color in spring; diurnal rhythm (singing and foraging in
the morning, going to roost at twilight).

Morphological adaptations “designed for purpose” in
birds are explained by the influence of climate, especial-
ly temperature (p. 567): in northern realms, for protection
against cold, birds have a thick covering of feathers rich
in down, which also covers the feet. Birds in the tropics
have less feathers and a lighter plumage, but there can be
luxurious feather growth in particular parts of the body
(e.g. the long “tail” feathers of peacocks, pheasants, and
birds of paradise). Feather coloration in the far north is
often white, gray, brown, or black, in lower latitudes how-
ever there is a dominance of lively and frequently irides-
cent colors.

After a long discussion regarding the global distribution
of birds, Tiedemann’s conclusion was that “through its
structure and formation, according to its own particular
climatic and physical conditions, each part of the Earth
has brought forth its own particular forms of plants and
animals” (p. 566). Therefore in any particular area the dis-
tribution of birds depends on its plant and animal produc-
tion (Mauersberger 1980: 11). Tiedemann ends his work
(1814) with a long bibliography of ornithological litera-
ture from each country and a discussion of the movements
and migration of birds in Europe/North Africa, Asia, North
America, and the southern hemisphere.

No other author in the 19th century attempted again to
write such a general study of ornithology, bringing the
great variety of aspects concerned together under overall
headings, as Erwin Stresemann was to do so brilliantly
over a hundred years later in his handbook Aves
(1927-1934). When examining Tiedemann’s explanations
of many facts we must distinguish between the function-
al and historical explanations that underlie every biolog-
ical phenomenon (Mayr 1982). The emergence of the
''zweckmafiige [adaptive/purposive] bark-coloration” in
the plumage of Wryneck and treecreepers in the course
of evolution, as well as the camouflage colors of ground-
breeding species and their “protective effecf ’, is explained
today by natural selection, while according to the 1 8th cen-
tury physico-theologians such phenomena were simply
seen as due to the wisdom and goodness of the Creator.
These are the historical, evolutionary-biological (ultimate)
explanations of the development of such plumage colors.
Their functional, immediate (proximate) explanations are

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to be found in the basic physiological processes control-
ling the differentiation of feather pigmentation and its dep-
osition on the growing feather. Similarly, there are certain
physiological hormonal reasons why a migratory bird
should begin its journey on a particular autumn night
(functional explanation), but the historical one of why it
has to leave its breeding site and migrate to different win-
ter quarters at all lies in its “genetic program”, which has
developed through natural selection in the course of evo-
lution.

In many cases Tiedemann discussed the “fittedness” of
avian structures (historical explanations) without using the
terms adaptedness or purposiveness, such as the stream-
lined form of the body as an adaptation to flight, the au-
tomatic clasping of the toes of arboreal species when the
knee joint is bent, and so on. In other cases he speaks as
a physiologist and tries to find functional reasons for cer-
tain phenomena, without attempting to consider possible
historical explanations for these cases. His functional ex-
planations were bound to fail because knowledge of the
entire physiological-chemical foundation of biological
phenomena was completely inadequate at the time. Stre-
semann (1951: 302-303) quotes several such examples
from Tiedemann’s book (1814, pp. 14, 72-73, 541, 572,
594, 597-599), though without emphasizing the impor-
tant difference between functional and historical explana-
tions. For example, Tiedemann looked for the origin of
the differing sizes of clutches among birds in their diet
(smaller number of eggs with animal food, larger number
with vegetable food; functional explanation), and Strese-
mann accused him of ignoring the biologically determined
greater or lesser reproduction requirement of the various
species (historical explanation). Tiedemann further be-
lieved that plumage color was determined by diet and tem-
perature (carbon in its multifarious degrees of oxidation
and the ambient temperature; functional causes), while
Stresemann remarked, “That plumage coloration in many
cases could be related to the necessity of a bird to protect
itself (Zorn 1742) is not even considered by our physiol-
ogist” (p. 302; historical cause). For the biologists - who
were mostly physiologists - working around the year 1 800
there was no alternative to a physical cause for traits, since
for them the teleological (historical) explanation of adap-
tations by the physico-theologians was no explanation at
all. It was only evolutionary biology grounded on natu-
ral selection that made an understanding of adaptedness
in nature possible. But Tiedemann and his contemporaries
had not yet reached that level of thinking, although he him-
self had very concrete ideas on the evolution of animals.
When he wrote that “With every great upheaval of the
Earth animals have become extinct, as the bones of mam-
moths, the Ohiothiere [mastodons], the Paldotherian
[horse- or tapir-like mammals], the Anoplotherian [pig-
or cow-like mammals], the Megatherian, the Megalonix
[two giant ground sloths], the Ornithocephalus [a genus

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of pterosaur] and many other animals conclusively prove.
But after each such upheaval it seems that new animals
have been created, presumably mainly via gradual meta-
morphosis and alteration of the older animals that had sur-
vived into new animal forms, effected by new climatic and

physical influences” (p. 322); “ species are subject

to metamorphosis over time just as individuals are” (1814:
325).

(6.) The “Golden Age” of field ornithology (1820-1850)

After 1750 and into the 19th century there was in Europe
a fairly large number of breeding birds that are now rare,
e.g. Short-toed Eagle (Circaetus gallicus), Ruff {Philo-
machus pugnax). Great Snipe {Gallinago media). Gull-
billed Tern (Gelochelidon nilotica). Stone-curlew (Burhi-
nus oedicnemus), European Roller {Garrulus glandarius),
Eurasian Hoopoe (Epupa epops). Rock Thrush (Montico-
la saxatilis), and Aquatic Warbler {Acrocephalus paludi-
cola). They inhabited the so-called common or unenclosed
land. This comprised a wide variety of landscapes, rang-
ing from gravel beds, impoverished grassland and heath-
lands, moors and bogs, riverine forest, and open wood-
land, accounting for approximately two-thirds of the area
of Central Europe and characterizing the appearance of
the countryside at that time. What remains of these habi-
tats today are mere remnants representing a small fraction
of the original (Schulze-Hagen 2005).

During the period 1820-1850 several ornithologists
were in contact with each other internationally, working
on a set of productive questions. They, for the first time,
constituted a discipline of ornithology (Farber 1982).

Johann Friedrich Nanmann (1780-1857) (Fig. 26). The
“Golden Age” (1820-1850) of Central European field or-
nithology wintnessed the appearance of the outstanding
works of Johann Friedrich Naumann (1820-1 844, 1 860),
Christian EudwigBrehm (1820-1 822, 1823-1824, 1831),
and Frederik (Friderich) Faber (1822, 1824-1827,
1 825-1 826). These publications laid the ground for future
research in the coming decades. The 1 820s heralded a first
great flowering of field ornithology in Germany.

Naumann’s 12-volume Naturgeschichte der Vogel
Deutschlands (1820-1 844, with addenda 1 844-1 854 and
a supplementary volume in 1 860) [Natural history of the
birds of Germany] was the titanic labor of one man that
served later generations as a handbook on the life and
habits of European birds (Fig. 27). For his names, see on-
line Appendix 3. Praise for the Naturgeschichte by two
great 19th-century British ornithologists was all but
boundless:

“[B]y far the most important work of this or any other pe-
riod was the publication of Naumann ’s Birds of Ger-

Bonn zoological Bulletin - Supplementum 59: 1-116

Fig. 26. Portrait of J.F. Naumann (from Thomsen & Strese-
mann 1957).

many’, which was commenced in 1820 and completed in
1844. Twelve octavo volumes of about 600 pages each tes-
tify to the industry of the author, whilst a careful study of
the content proves him to have possessed a knowledge of
the various plumages of the birds of which he treats, their
habits, songs, call-notes, food, and all the numerous de-
tails of their history, which a lifetime devoted to their ob-
servation was able to teach, not only unrivalled by any
author before or since, but far above and beyond all hope
of rivalry [...] Had this work only been translated into
English, half the nonsense that subsequent ornithologists
have written on birds would never have appeared (Hen-
ry Seebohm 1885: XII). “This Naturgeschichte der Vogel
Deutschlands, being almost wholly re-written by his son
J.F. Naumann, is by far the best thing of the kind as yet
produced in any country. The fulness and accuracy of the
text, combined with the neat beauty of its coloured plates,
have gone far to promote the study of Ornithology in Ger-
many” (Alfred Newton 1896: 17), and “for fulness of
treatment, perspicuity, and general accuracy, the work of
Johann Friedrich Nauman has not been surpassed (Al-
fred Newton 1905: 37; see also Thomsen & Stresemann
1957: 180-181).

In a manner of speaking, Naumann senior had got his son
Johann Friedrich off to a flying start by introducing him
to ornithology at the tender age of 8 or 9 years old, pass-
ing on all his accumulated wisdom and enabling the young
man to build on an already substantial foundation. In great

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Jurgen Haffer (t) et al.

n<t[t[<r acFcllf^D fti n litililittr.

^aturflcfd)i(^>te

0 tr

gjoflcl ©cutfd;lttnl)g,

r a 4

eigcneii ^rfa&rungfn rntmorfen.

S u c 1 ^ im 4

Hm^rarhlht, fltprbTlll, fr^f Wirm.fcrf,

fM, mH wit attf.n no* t>« (istnbwtii ftriri*i«i<ii

uiib (iffledKitin atfciitiw^tn aBir fctnlf*«n HftH itH<H

ftaillHl>trf(i}«b(n^(i[(ll, auf# 3if“t
1;i;fi n

6 4 f f m 0 e M *

Fig. 27. Title page of J.F. Naumann, volume 1 (1820).

detail, the voluminous text of the Naturgeschichte deals
with the name, synonyms, description, occurrence, habits,
diet, breeding, predators, hunting, and beneficial and
harmful effects for every species. This pattern in describ-
ing bird species was probably taken by Naumann from
Blasius Merrem (Thomsen & Stresemann 1957: 93). The
anatomist C.L. Nitzsch, and after his death R. Wagner,
helped with contributions on the anatomy of the various
families dealt with in individual volumes. In the foreword
to the second volume in 1 822, Naumann discusses the, for
him, unbridgeable opposition between the representatives
of the two great branches of ornithology: the field ornithol-
ogists and the systematists, criticizing the continual
changes being made to the “artificial construction” that
is avian systematics:

“Everyone thinks they can have a try, and if someone has
seen a couple of hundred mounted birds then he wants to
be a reformer and improve the system. — Go out into the
field, see the animals in their lives and habits; how dif-
ferent then will you judge when you have found that things

are not what they appeared to be in the cabinet [ ] It

has always seemed to me that the intellectual and schol-
arly philosopher of nature [= systematist] cannot be rec-
onciled with the son of nature [= field ornithologist] who
simply follows the straight path that nature has marked

for him. — Whoever has made it his aim not just to hunt
birds for his collection but also to observe them in their
haunts and in their ways, and to study their habits in na-
ture, will find as much pleasure in this as work and will
have little time left to occupy himself with the scribblings
of the creators of systems ” (1 822: I-II).

The way of life of birds. The focus of Naumann’s work
lay in the portrayal of the way of life of birds, which he
described with affection and in great detail. Habitat pref-
erence, diet, nesting habits, and relationship to their sur-
roundings are minutely discussed, but there is little trace
of the earlier theoretical impulses of J.H. Zorn or F. Faber
in Naumann’s approach, which is more painstaking ob-
servation than causal research (Mauersberger 1980). The
competitive relationships between several species is
again something that Naumann appears uninterested in.
Nevertheless, there are the beginnings of a causal outlook
in areas of ecology in some of Naumann’s findings, e.g.
the relation between diet and foraging method. Habitat
preference can be discerned in Naumann’s texts and for-
aging sites are clearly recognizable (tall herbs, treetops,
ground), as are the composition of the diet, nest site, nest
material, and the many different ways of movement and
forms of behavior in birds.

Illustrations. The birds in the Naturgeschichte are illus-
trated on a total of 391 excellent hand-colored copper-
plates, 379 of which are by J.F. Naumann and which he
engraved anew for this work as well as making many im-
provements in their drawing compared with earlier ver-
sions. The additional 12 plates in the supplementary vol-
ume of 1 860 are by F. Sturm (see Neuer Naumann, Vol. 1 ,
p. XXV, 1897). Naumann achieved a high degree of per-
fection, especially in his treatment of plumages, so that
his plates are counted among “the most lifelike and accu-
rate representations of birds ever made” (Nissen 1953,
1957). However Naumann’s paintings - of raptors in par-
ticular - were the subject of some critical remarks by one
contemporary ornithologist. Hermann Schlegel (1849) in
Leiden, who had himself published large-format illustrat-
ed books, wrote:

“Among the older German artists, Johann Friedrich Nau-
mann and Susemihl deserve special mention. The former
has managed, through long and careful study of the liv-
ing bird, to produce small, often very characteristic,
painstaking illustrations, but they reveal the hand of the
naturalist rather than the artist. They are usually lacking
in spirit, liveliness, and elegance. The engraving is most-
ly hard and unsure, the embellishments meager; all in all
even the drawing, with the exception of the waterbirds,
marshbirds, and songbirds, leaves much to be desired, and
not infrequently is quite disfigured by Naumann ’s habit of
drawing the eyes much larger than they are in nature. This

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is especially the case in his birds of prey, which (as in most
works) leave the most to be desired, and often are even
less than mediocre. ”

Ludwig (2000: 101) saw the reason for the stiff represen-
tation of the raptors in the fact that Naumann had only
“limited artistic ambitions, stuck to traditional patterns of
illustration, and did not develop his own pictorial forms
corresponding to his demands. He did not present birds
as if he were drawing living creatures moving through
their environment but mounted specimens on their
pedestals placed before white walls”. While it is actually
the case that in these instances Naumann had indeed cre-
ated plates for ornithologists, and not as an artist,
Schlegel had specifically excluded Naumann’s plates of
waterbirds, marshbirds, and songbirds from this criticism.
Here the birds appear as living creatures, and their habi-
tat is often suggested in the background. On the whole
Naumann’s bird paintings are evidently illustrations for
his scientific Naturgeschichte and not works of art in their
own right.

New Descriptions. In the course of their efforts to exact-
ly differentiate the species and subspecies of birds, father
and son Naumann (later Johann Friedrich alone) described
a host of new forms, and gave them scientific names.
Some of these names have retained their validity while
others have failed to gain permanent recognition. Later
work showed that many of them referred to already known
species or subspecies'®:

Cygnus xanthorinus J. F. Naumann, 1 842 [= C. c. cygnus
(L.), Whooper Swan]

Cygnus melanorhinus J. F. Naumann, 1 842 [= C. bewickii
Yarrell /columbianus, Bewick’s Swan]

Anser intermedius J. F. Naumann, 1842 \= A. albifrons
flavirostris Dalgety & Scott, 1948", Greenland White-
fronted Goose]

Anser minutus J. F. Naumann, 1842 \=A. erythropus (L.),
Lesser White-fronted Goose]

Anas leucopis J. F. Naumann, 1799 \=Aythya nyroca (L.),
Ferruginous Duck]

Anas merganser J. F. Naumann, 1799 [= Alopochen ae-
gyptiaca (L.), Egyptian Goose]

Buteo leucurus J. F. Naumann, 1 853 [= B. r rufinus (Cret-
zschmar). Long-legged Buzzard]

Limosa baueri J. F. Naumann, 1836 [= L. lapponica ba-
Mm Naumann, 1836, Bar-tailed Godwit]

Tringa macroura J. A. & J. F. Naumann, 1811 [= Bartra-
mia longicauda (Bechstein, 1812), Upland Sandpiper]
Scolopax Glottis minor J. A. Naumann, 1799 [= Tringa
stagnatilis (Bechstein, 1 803), Marsh Sandpiper]
Phalaropus angustirostris J. F. Naumann, 1 836 [= Ph. lo-
batus (L.), Red-necked Phalarope]

Bonn zoological Bulletin - Supplementum 59: 1-116

Mormon [= Fratercula] corniculata J. F. Naumann, 1821,
Horned Puffin]

Lariis glaucescens J. F. Naumann, 1 840, Glaucous-winged
Gull

Larus cachinnans michahellis J. F. Naumann, 1840 [=
Larus michahellis J. F. Naumann, 1840, Yellow-legged
Gull

Sterna macrura J. F. Naumann, 1819 [= S. paradisaea
Pontoppidan, 1763, Arctic Tern]

Sterna fluviatilis J. F. Naumann, 1819 [= 5. /z. hirundo L.,
Common Tern]

Acrocephalus lacustris J. F. Naumann, 1811 [=A. a. arun-
dinaceus (L.), Great Reed Warbler]

Acrocephalus stagnatilis J. F. Naumann, 1811 [= Locustel-
la fluviatilis (Wolf, 1810), River Warbler]

Sylvia (Calamoherpe) horticula J. F. Naumann, 1853 [=
Acrocephalus s. scirpaceus (Hermann, 1804), Eurasian
Reed Warbler]

Sylvia cariceti J. F. Naumann, 1821 \= Acrocephalus palu-
dicola (Vieillot, 1817), Aquatic Warbler]

Sylvia ruficapilla J. F. Naumann, 1 853 [= S. a. atricapil-
la (E.), Blackcap]

Troglodytes musculus J. F. Naumann, 1823 [= T aedon
Naumann, 1823, Southern House Wren]
Troglodytes stellaris J. F. Naumann, 1 823 [= Cistothorus
platensis stellaris 1823, Sedge Wren]

Troglodytes Bechsteinii J. F. Naumann, 1 822 [= T rufi-
collis atrogularis Jarocki, 1819, Rufous-browed Wren]

The Naumanns described the Upland Sandpiper under the
name Tringa macroura from a bird that had been collect-
ed as a “vagranf ’ on the River Werra in Hessen and sent
to Herr von Minckwitz in Silesia (see Nachtrag 5: 274,
1811). J.F. Naumann very likely examined and sketched
the specimen during his visit there in 1805. The mount-
ed Homed Puffin that Naumann used in his initial descrip-
tion of the species was among a collection of seabirds
“from the waters around Kamchatka” that he had received
from Peter von Woldicke (Brunsbiittel) and that most
probably had been obtained during the Russian expedi-
tion of 1 8 1 5-1 818 (Hildebrand! 200 1:39). The Glaucous-
winged Gull was described by Naumann from North
American specimens in the Berlin Zoological Museum in
the 10th volume of his Naturgeschichte in 1840.

Naumann brought three skins of a laughing-dove-type
columbid from Turkey home with him from his trip to
Hungary in 1835. He had noticed them in the National
Museum in Budapest and sent a description and colored
illustration of the bird to his Hungarian acquaintance E.
von Frivaldsky, who used both of them in his publication
(1838) on the Collared Dove under the name Columba
risoriavm. decaocto Friv. (Stresemann 1953; Thomsen &
Stresemann 1957: 126-127).

A series of bird species and subspecies were named by
contemporaries or later ornithologists in J.F. Naumann’s

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honor. Some remain valid, while others were syn-
onymized:

Fratercula arctica naumanni Norton, 1901, Atlantic Puf-
fin

Falco naumanni Fleischer, 1818, Lesser Kestrel
Turdus naumanni Temminck, 1 820, Naumann’s Thrush
Gallinula naumanni Schinz, 1821 [= Porzana pusilla in-
termedia (Hermann), Baillon’s Crake
Acridotheres tristis naumanni Dementiev, 1958, Common
Myna

Sylvia naumanni Muller, 1851 [= S. a. atricapilla (L.),
Blackcap]

Troglodytes naumanni Brehm, 1855 [= T. t. troglodytes
(L.), Winter Wren]

‘‘The Naumann CuW\ The more substantial books deal-
ing with the avifauna of Germany which appeared in the
latter half of the 19th century were all more or less ex-
tracts from “Naumann”. Many ornithologists were con-
vinced that almost everything worth knowing about Ger-
many’s birds was now available: “Because of the smooth
polish of the descriptions, [Naumann’s Naturgeschichte]
had more of an instructing than a stimulating effect” (Stre-
semann 1951: 314, 352). Even Altum’s interesting ideas
in his book Der Vogel und sein Leben (1868) [The bird
and its life] on the impact of instincts in the life of birds
found no resonance among his colleagues. Thus an uncrit-
ical “Naumann cuh” (Heinroth 1917) had arisen among
ornithologists which put a brake on progress in the disci-
pline and culminated in a new edition of Naumann’s
Naturgeschichte (1897-1905) in 12 folio volumes, with
new illustrations by a variety of artists. The copper plates
of Naumann’s bird paintings were no longer extant so new
originals had to be commissioned from a series of artists.
His text was reprinted but many passages, mostly regard-
ing faunistics, were inserted. This new edition was sub-
ject to considerable criticism from various sides, but, as
Hartert wrote (1930: 4): “One can be as critical as one likes
about the so-called NaumanF (and there is indeed
much in it to criticize!) but it must be admitted that it has
made Naumann’s name better known than ever before and
has reached many more hands than the ‘Old Naumann'
would ever have done.”

Also the printing quality of the New Naumann was at-
tacked, as the following example from more recent years
illustrates:

“As a bibliophilic enterprise this edition fails in every cat-
egory. Printing quality, typography, print area, everything
shows a dreadfully low level of taste. Although excellent
work has been done by artists like Otto Kleinschmidt, An-
ton Goring, J. Keulemanns, Stephan von Necsey Bruno
Geisler, Oskar von Riesenthal, and E. van Maes, the col-
ors of the chromolithographic plates are blatantly over-
done and strive only for noisy effect” (Marholz 1965: 384).

Bonn zoological Bulletin - Supplementum 59; 1-116

Schalow (1909) presents an interesting survey of Nau-
mann’s status and the research prompted by his work. In
1930, on the 150th anniversary of Naumann’s birth, L.
Schuster, E. Hartert, O. Heinroth, E. von Boxberger, O.
Kleinschmidt, and O. Reiser honored his achievements
from a number of different perspectives in the journal
Beitrdge zur Fortpflanzungsbiologie der Vogel (Vol. 6:
1-9), but also pointed out various shortcomings in his
work, such as the “simply invented” incubation periods
for many species. “Naumann gives a nicely rounded off
account for almost every bird and admits no gaps in his
observational knowledge, since how else could he man-
age to claim that the completely naked nestling of the
Black Woodpecker was covered in down?”, asked Hein-
roth (1930: 5). It was he who “finally helped [ornitholog-
ical progress] in Germany to breach the wall erected by
the Naumann cuh” (Stresemann 1951: 352), mainly
through the weighty 4-volume work by him and his wife
on Die Vogel Mitteleuropas in alien Lebens- und Entwick-
lungsstufen photographisch aufgenommen und in ihrem
Seelenleben bei der Aufzucht vom Ei ab beobachtet
(1924-1933) [The birds of Central Europe photographed
in every stage of life and development and observed in
their behavior from the egg onward], which contained
much new material on the biology of the European birds
that he raised together with his wife Magdalena. Hein-
roth’s (1917) critique of some statements in Naumann’s
work does little to damage its reputation; the author of a
publication of such magnitude inevitably suffers the fate
of a few errors here and there in its volumes:

“He did not notice the voice differences beween the sex-
es in ducks and did not believe that the young Cuckoo
ejected its fellow nestlings from the nest. As was gener-
ally believed in his day, he ascribed conscious action to
birds. ”

Even the young G. Stein (1928: 129) dared to find fault
with Naumann: his account of the breeding biology of the
Common Sandpiper “must be regarded as erroneous in its
most important points”, he wrote.

In a historical essay dealing with statements on the in-
cubation periods in a series of bird species in the works
of past ornithological authors, Margaret Nice (1954)
showed that certain false assertions have been repeated in
the literature from Aristotle right down to the present day
(see Table 2, Haffer 2006: 36). The duration of incuba-
tion was underestimated by many early authors, especial-
ly in raptors and some marshbirds, the mistaken assump-
tion being that they assumed that the body size of a bird
or volume of its egg were the important determining fac-
tors. While these parameters do determine the length of
the incubation time to some extent, there is considerable
variation in different avian families. Zorn (1742) gave
some correct information concerning incubation based on

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67

his own field observations, but they remained unregard-
ed by later authors. Bechstein and the Naumanns estimat-
ed incubation duration in many species on the basis of
body size, and they were mostly wrong. Only since the
publications of Heinroth and Niethammer have correct in-
cubation times been given for the majority of species. Of
the 11 incubation durations given by Heinroth (1922) all
are correct, while of the 15 set down in Niethammer’s
Handbuch (1937-1942) only that of Black Tern (CMido-
nias niger) is mistaken (14-17 instead of the correct 20-22
days).

Christian Ludwig Brehm (1787-1864)

“Old Brehm”, a Protestant parson in Renthendorf (Thiirin-
gen), was a late physico-theologian (Fig. 28). The study
of nature, and of birds in particular, was for him “a true
divine service”, allowing him to see “the infinite wisdom
of the Almighty even in insignificant and seemingly chaot-
ic things”. “The naturalist must follow the Creator and try
to recognize His footsteps everywhere. The more pious
our heart, the more will the veil be lifted that hides the
Works of God from our feeble eyes” (Brehm 1827).

Like his fellow divine J.H. Zorn one hundred years ear-
lier, Brehm was a teleologist who everywhere saw “the
most perfect expressions of Purpose”. He asserted that
“Every creature is fitted in every way to the place in which
it lives, and for the food that it eats [. . . ] In cold countries
plumages are thicker than in warm climes. The Ptarmi-
gan that live in the north and in the Alps are yellow, brown
and black in the summer, like the rocks among which they
live, and white in the winter”. He compared the details of
the climbing apparatus of the Nuthatch and the woodpeck-
ers (1822, 1827; see Stresemann 1951: 305-306). Hence
Brehm became, like Zorn before him, a pioneer of func-
tional or biological morphology, in that he studied the
adaptations of birds to their environmental conditions. His
3-volume Beitrdge zur Vogelkunde (1820-1 822) [Contri-
butions to ornithology] had a lasting influence on the
course of the young discipline. Thanks to his extensive col-
lection of skins - around 9000 specimens at the end of his
life - he was able to study molt, juvenile and adult
plumages, and individual variations in many species.
Brehm edited the first ornithological journal, Ornis, oder
das Neueste und Wichtigste der Vogelkunde (1824-1827)
[Ornis, or the latest and most important information on
ornithology], but it ceased publication after only three is-
sues.

As an ornithologist Brehm was a meticulous observer.
For example he demonstrated that birds could simultane-
ously be good biological species and morphologically very
similar (sibling species), such as Common and Short-toed
Treecreepers, Marsh and Willow Tits, Firecrest and Gold-
crest, or Crested and Thekla Larks. Despite considerable
opposition from several colleagues he insisted that these
birds belonged to separate biological species and occurred

Bonn zoological Bulletin - Supplementum 59: 1-116

Fig. 28. Portrait of C.F. Brehm (from Haffer 200 1 ; Brehm-Ge-
denkstatte, Renthendorf, Thilringen).

sympatrically without interbreeding. He was eventually
proved right, although it would take more than half a cen-
tury until the last skeptics admitted that the two treecreep-
ers and the two tit species each belonged to different
species. Common Nightingale and Thrush Nightingale
(Sprosser) had been known to naturalists as separate
species since the time of J.L. Frisch (1733), when it was
also realized that the very similar leaf warblers {Phyllo-
scopus) as well as the dark-colored flycatchers (Ficedu-
la) should probably be split into different species.

During the 1 820s and 1 830s Brehm assumed that each
one of his morphospecies - European Robin, Common
Chaffinch, Common Redstart, etc. - represented biolog-
ical entities whose subspecies replaced each other to cre-
ate in each case a geographical-ecological mosaic. Since
he believed (in contrast to most of his contemporaries and
to us today) that along their contact zones these subspecies
did not interbreed, he treated his subspecies as (in today’s
language) parapatric species, and most of his morphos-
pecies as Artenkreise (“species circles”, or here super-
species). In this matter the main thing for us is to com-
prehend his way of thinking and to understand his over-
all concept. With his assertion that morphospecies “split”
into several “genera” (= subspecies) Brehm did not real-

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Jurgen Haffer (t) et al.

ly mean a “dissolution” of the morphospecies concerned
but simply their taxonomic classification or subdivision.
The great majority of his numerous ecological “sub-
species”, which each belong to a particular morphospecies
and, according to Brehm, are morphologically distinguish-
able and replace each other geographically in (e.g.) conif-
erous or deciduous woodland or bushy landscapes, were
“ficticious” and were not confirmed by his successors*^,
but 55 of his geographical subspecies are still currently
recognized (Haffer 1996, 2003, 2006).

Several 20th-century authorities believed that in his
books of 1 83 1 and 1 832 Brehm had actually divided most
Central European bird species into several separate
species, so that, according to his ideas, in a single habi-
tat one could see, alongside each other, a variety of species
of Blackbird, Robin, Chaffinch, etc. This is a false alle-
gation or a misunderstanding of Brehm’s interpretation,
because in the 1 830s he never held such an opinion as far
as the great majority of bird species were concerned. In
Brehm’s view at the time, in any particular habitat there
mostly existed only one representative of a morphospecies,
i.e. only one Blackbird, Robin, or Eurasian Jay. Accord-
ing to his view in 1 83 1 , of 3 1 1 of his subspecies of Ger-
man “songbirds” (in today’s sense), 289 replace each oth-
er geographically or ecologically within their respective
morphospecies, while for only 22 (7%) subspecies does
he claim that they live alongside another subspecies of
their morphospecies in the same habitat (syntopically).
However in quite a few cases his statements on distribu-
tion are very vague, so these assertions, plus a few oth-
ers, should be taken cum grano salts.

In the course of the 1 840s he became increasingly un-
clear in his writings about the geographical-ecological sep-
aration of a number of his subspecies, confidently assert-
ing that in several cases different subspecies of a morphos-
pecies occurred together in the same habitat, breeding but
not hybridizing. However in many instances he himself
was uncertain whether he was dealing with different taxa
or with varieties (individual variations) within a popula-
tion, writing: “they could be called subspecies or vari-
eties”. Then, in the 1850s, he actually interpreted most of
these “subspecies” as individual variations within a sin-
gle taxon, thus taking back his earlier opinion that these
“subspecies” of a morphospecies represented separate
sympatric taxa. Yet Brehm was not always consistent and
his taxonomic views, as we have seen, altered substantial-
ly over time. This must be taken into account when con-
sidering the theoretical ideas regarding his species and
“subspecies”. His postulations from the 1830s on the one
hand, and those mainly from the 1850s on the other, dif-
fer enormously from each other, though in both cases they
are clear and comprehensible. His views in the 1840s,
however, are today difficult to follow. He was dealing at
length with several cases of sympatric subspecies of a mor-
phospecies, and his writings are best regarded as “transi-

Bonn zoological Bulletin - Supplementum 59; 1-116

tional” between what he believed formerly, in the 1830s,
and latterly, in the 1850s. It was in this last decade that
he began to adopt ternary nomenclature, giving subspecies
three names (genus, species, subspecies), hence differen-
tiating them from the binary-named species.

In his review of C.E. Brehm’s ideas on avian species,
Eck (2006), in tune with several authors in the 20th cen-
tury, only took account of the reproductive aspect, giving
attention neither to the change over time in Brehm’s think-
ing nor to the geographical-ecological occurrence of the
individual forms (subspecies) within Brehm’s morphos-
pecies. The reader might gain the false impression that all
of Brehm’s forms were mere fiction, yet the number of
morphospecies of Central European birds distinguished by
Brehm agrees almost exactly with the number of biolog-
ical species.

Friderich Faber (1796-1828). The Dane Friderich
(Frederik) Faber had a theoretical mind, and in his short
life made many important observations on the biology and
ecology of Arctic birds (Helms 1928, 1934; Stresemann
1951 : 308-309). Stresemann (1928: 182) called him “one
of the most innovative ornithologists” of the 19th centu-
ry, who attempted to create a comparative biology of
northern birds “ in order to reveal the underlying causes
of what is visible, and in doing so he raised himself high
above his famous contemporaries like Johann Friedrich
Naumann und Christian Eudwig Brehm”. Faber traveled
in Iceland from May 1819 until September 1821 with the
intention of carrying out exact observations to obtain re-
sults that would be generally applicable. Faber, like the
philosopher Immanuel Kant, was convinced that “all of
Nature is nothing more than a linking together of phenom-
ena according to laws, and there is absolutely no irregu-
larity” (Faber 1825-1826, Foreword, p. IX). He published
in Germany, initially in Oken’s journal Isis, and belonged
to the group around C.E. Brehm and J.F. Naumann, with
whom he was in regular correspondence during the 1 820s.
In 1823 he also visited the Naumanns in Kothen and
Ziebigk.

Faber had clear ideas about biological species and wrote:
“Individual birds freely mating with each other in nature
belong to one species” (1825: 117-118). Species can vary
geographically, i.e. morphologically differentiated region-
al groupings (populations) do not necessarily belong to an-
other species (as some ornithologists of the time believed,
based on a strictly morphological species concept). In a
letter to J.F. Naumann, Faber wrote:

“I call a bird a species when individuals freely and nat-
urally mate and, in addition, produce young capable of re-
producing themselves; it is certainly the case with U[ria]
troile [= Uralia aalge. Common Murre] and U. tr. leucoph-
thalmus [= ‘Bridled’ Common Murre] that they mate with
each other but because the latter is simply a race of the
former they do not lose their diagnostic features, so that

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when [they] are paired with each other the young some-
times lack a white eye ring and sometimes have one. These
2 birds resemble each other almost completely in their his-
tory and form; but this is not the case with U. Brunnichii
[= U. lomvia. Thick-billed Murre] and both of the others.
Uria tr leuc. eggs are just like those of U. troile but in
both of them the color of the eggs varies almost from one
individual to the next” (26 March 1822; see Klein 1910).

Faber (1825-1826) distinguished between sedentary
birds, non-directional dispersers, and migratory birds. The
“migratory instinct” causes the last of these to depart in
autumn while the “home-sickness instinct” brings them
back to their breeding grounds in spring. Avian migration
always takes place from the poles toward the equator, nev-
er the reverse. The further north a bird breeds the earlier
it begins its autumn migration and the later it returns in
spring. Male songbirds arrive in Iceland a few days ear-
lier than females. In the breeding quarters the “home-sick-
ness instinct” is replaced by the “mating instinct”, lead-
ing to mating, nest building, egg laying, incubation, and
raising the young. In monogamous birds, Faber distin-
guished between (a) double monogamy (both sexes raise
the young together; e.g. coastal species like auks and cor-
morants), (b) intermediate monogamy (the young are led
to the sea immediately after hatching, where they forage
for food on their own; e.g. freshwater birds like Colym-
bus [= Gavia], Podiceps, Fulica), and (c) single
monogamy (females raise the young on their own; e.g.
Cygnus, Anser, Anas, Mergus). In addition, Faber talks
about birds’ eggs: the number in the clutch of various
species, absolute and relative size (compared with the size
of the bird in question), shape, surface texture, and color.
Synoptic (dichotomous) tables illustrate (1) mating,
breeding, and feeding situations of Icelandic birds, (2)
walking ability, (3) flying ability, and (4) swimming abil-
ity of boreal waterbirds.

In his Prodromus der islandischen Ornithologie (1822)
(Fig. 29) [Preliminary study of Icelandic ornithology^
Faber gathered together short statements on the ecology
of individual species, and in his Beytrdge zur arctischen
Zoologie (\S>24, 1826, 1827) [Contributions to Arctic zo-
ology] he published extensive accounts - short mono-
graphs in essence - on all the birds he had observed {Podi-
ceps, Fulica, Phalaropus, Puffinus, Procellaria [= Hydro-
bates], Uria, Carbo, Colymbus [= Gavia], Passeres, Fal-
co, Strix, Mormon [= Fratercula], Alca). “All the qualties
required to create the genuine expert were united in
Faber”, wrote Stresemann (1951; 308), “total command
of the literature, a thorough grounding in anatomy, a sharp
eye in the study of bird skins, and an understanding
amounting to genius for what the biologist must look for
in the field if he wishes to understand and not only de-
scribe. More critical than the enthusiastic Brehm, a finer
mind than the quiet Naumann, Faber unquestionably oc-
cupies the first place in this illustrious company.”

ttt

\

©efd)t^te SSogd
^ i fl n t

bet liitfruirtn

Jtop cn^aig en/ 1822* *

3(uf AvUtn ScilafTctA

* IttEudt bti Pa

Fig. 29. Title page of Faber (1822).

Constantin Wilhelm Lambert Gloger (1803-1863).

Gloger studied the avifauna of Silesia, especially in the
mountainous Riesengebirge (1827) on today’s
Polish/Czech Republic border (Karkonoski in Polish,
Krkonose in Czech), and published a handbook of the
landbirds of Europe (1 834). He had a heated dispute with
C.L. Brehm about how to establish species boundaries in
birds, relatively broadly or relatively narrowly (see below).
But both of them had the same theoretical concept of what
is generally meant by “species”, namely reproductive com-
munities (on a typological basis), a point of view wide-
spread among German ornithologists around 1 800, build-
ing on the works of the 17th-century John Ray and the
18th-century G. Buffon (e.g. Blumenbach, Zimmer-
mann, Illiger, and Oken).

Gloger (1833: X) wrote: “Animals which mate with
each other in the wild (though not only exceptionally) be-
long to the same species”. And Brehm: “Those creatures
that regularly mate with each other in a free and natural
situation are of one and the same species [. . . ] No one, least
of all Herr Gloger, can raise any serious objection to this
definition of a species”.

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Jurgen Haffer (t) et al.

So Gloger and Brehm supported the same species con-
cept in theory, but in practice both ornithologists had very
different opinions on the boundaries of the taxonomic cat-
egory “species”.'^ The taxonomic category of species in
Gloger’s system was broad and inclusive. For many
species it encompassed geographical “varieties”, which he
described in words but did not award their own taxonom-
ic names, given the often continuous or clinal gradients
of difference between these subspecies. By contrast, C.L.
Brehm’s taxonomic species category was narrow, gener-
ally encompassing only a single taxonomic form or sub-
species, because he assumed that subspecies of a morphos-
pecies would not hybridize with each other. Hence Brehm
was an early taxonomic “splitter”, while Gloger was an
early “lumper”, who brought together a relatively large
number of geographical forms to create very broad poly-
morphic species taxa. In many cases he interpreted vari-
eties as subspecies that later would be shown to be species,
such as:

Anthus pratensis (inch A. cervinus and rufogularis)
Anthus aquaticus [= A. petrosus ?] (incl. A. littoralis)
Motacilla alba (incl. M. lugens)

Motacilla flava (incl. M. melanocephala \=feldegg ?])
Parus major (incl.g P. monticolus)

Sturnus vulgaris (incl. S. unicolor)

Cinclus aquaticus [= cinclus] (incl. C. pallasii)

Corvus monedula (incl. C. dauuricus)

Emberiza hortulana (including E. caesia)

Passer domesticus (incl. P. hispaniolensis and italiae)

These facts indicate that Brehm and Gloger stood for con-
trary and extreme positions in questions of taxonomy, yet
in the final analysis both of them were mistaken. Brehm
defined species too narrowly, and Gloger often too broad-
ly. The “golden mean” that later ornithologists would ar-
rive at on the basis of additional information lay between
the extreme viewpoints of these two doyens of ornithol-
ogy.

Gloger introduced some genus and subspecies names that
are still valid (in bold)''^-

Taoniscus Gloger, 1842 - Dwarf Tinamou
Buteo buteo vulpinus (Gloger, 1 833) - “Steppe Buzzard”
Necrosyrtes Gloger, 1841 - Hooded Vulture
Tympanuchus Gloger, 1841 - prairie chickens
Bugeranus Gloger, 1 842 - Wattled Crane
Neomorphus Gloger, 1 827 - ground cuckoos
Bubo bubo sibiricus (Gloger, 1833) - “Siberian Eagle-
Owl”

Rhinoplax Gloger, 1841 - Helmeted Hornbill
Pelargopsis Gloger, 1841 - kingfishers
Nystactes Gloger, 1 827 - pufllbirds
Xipholena Gloger, 1841 - cotingas

Bonn zoological Bulletin - Supplementum 59: 1-116

Motacilla alba lugens Gloger, 1829 - ""Trauerbachstel-
re” [= M. (a.) yarrellii - Pied Wagtail; M. lugens - Black-
backed Wagtail]

Terpsiphone Gloger, 1827 - paradise flycatchers

It goes without saying that Gloger - just like Brehm and
the great majority of biologists prior to the appearance of
Charles Darwin’s epochal book in 1859 - was a typolo-
gist or essentialist, believing in the constancy (invariabil-
ity) of species, and certainly not an evolutionist. To illus-
trate his view in this regard he quoted with great approval
the contemporary botanist E. Meyer, who had written:

“The immutable integrity of species is the only fixed point
around which [...] their varieties revolve in continuous os-
cillation and dynamic, so that even an observer feels dizzy
watching it in action [...] Nothing is secure if the ground
is not firm ” (see Gloger 1833: 135).

Gloger devoted a most interesting book of lasting value
to geographical variation in birds, with the title DasAbdn-
dern der Vogel durch Einfiufi des Klima’s (1833) [The
variation of birds under the influence of climate^. Here he
explained in some depth his view that “those few people
who [. . . ] wish to regard climatic varieties as species must
be seen as following the wrong path. What one cannot de-
limit one ought not to separate!” (p. 5). More than one-
third of the species known to Gloger vary geographical-
ly (p. 137); he listed them in his handbook of 1 834, where
they are described in detail. Regarding the actual appear-
ance of geographical variation, he noted that darker col-
ors (black, black-brown, gray, brown, rust red) are more
intensely developed in those forms of a species living in
warmer climates. Conversely, in cold northern climes
plumage becomes paler and whiter (pp. 12, 15). The pelt
of mammals too varies with climate in the same fashion
as plumage color in birds (p. 38). This phenomenon was
formulated as Gloger ’s Rule by Rensch( 1929: 152, 1934:
25), under which name it has been discussed in many ma-
jor works of zoogeography and evolution (e.g. Huxley
1942: 213; Mayr 1942: 90, 1967: 256, 1984: 200, 449;
Rensch 1954: 43, 47). Gloger himself (1833: 73-78) stat-
ed that “voice, song, color, and other life qualities, resi-
dence, etc., all can change”, both individually in the same
place or geographically. However, he did not believe that
the geographical differences in the various subspecies of
a particular species of bird were genetically fixed, so that
if members of a form were introduced into an area occu-
pied by another form of the same species “then after a few
years they would either resemble the form native to that
place, or their offspring conceived there would resemble
them in the second or third generation [. . . ] climatic species
cannot exist, only simple varieties” (1833: 106, 107).

Eike Brehm (1827), Gloger (1829) pointed out that “the
females of many open-nesting species of ducks and fowl

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are inconspicuously colored; hence Nature gives her spe-
cial protection to the parent bird that is both most endan-
gered and most important for the continuation of the
species [. . . ] The color of the eggs is in many cases more
or less clearly adapted to the surroundings, especially in
ground-nesters like Nightingale and Sky Lark” (Strese-
mann 1951; 325),

Gloger popularized practical bird conservation in the
1840s (Barthelmess 1981), and when Jean Cabanis
founded the Journal fur Ornithologie in 1 853, Gloger was
invited to be a collaborator (see below). In the following
years he published numerous commentaries and reviews
on avian biology.

Professor Lichtenstein and Gloger ’s Rule. Gloger had
grown up in the country in Upper Silesia, had attended
grammar or high school in NeiBe, and from autumn 1821
studied zoology in Breslau (Wroclaw). Of particular im-
portance to him was the winter semester of 1 824/25, when
he matriculated at the University of Berlin and for 8
months was a student of H. Lichtenstein, the professor of
zoology (Moller 1972). Since the founding there of the Zo-
ological Museum in 1810, the professor had a consider-
able collection of Eurasian birds and mammals at his dis-
posal, and in his lectures and courses often drew attention
to the striking geographical variations in the coloration of
many species due to the influence of different climate
types. This phenomenon had already been pointed out by
Peter Simon Pallas (1741-1811), the great scientific ex-
plorer of Siberia and the Far East, in his important
Zoographia Rosso-Asiatica of 1 8 1 1 . Eichtenstein himself
published hardly anything on these important conformi-
ties, but was in full agreement with his student Gloger us-
ing the Berlin material to study geographical variation in
the coloration of birds and mammals and later publishing
his findings in a long paper. Eichtenstein wrote a foreword
to the work, in which he stated his agreement to those el-
ements in Gloger’s thinking which he (Eichtenstein) had
formulated in lectures, conversations, and letters. Follow-
ing his return to Breslau in the summer of 1825, Gloger
wrote a long series of letters to Eichtenstein, which are
held in Berlin, although the professor’s replies are miss-
ing.

A high percentage of bird species follow Gloger’s Rule,
whose adaptive basis is a protective or camouflage col-
oration to confuse predators, prey animals, or competitors.
More heavily pigmented feathers and hair are also advan-
tageous in combating damage from bacteria, which are
commoner in warm humid climates than in dry ones (Burtt
& Ichida 2004).

Ornithological book projects left unfinished by Gloger.

In the early 1 830s Gloger started several major book proj-
ects, but in each case, following publication of their first
parts, abruptly abandoned and never finished them:

Bonn zoological Bulletin - Supplementum 59: 1-116

(1) Vollstandiges Handbuch der Naturgeschichte der
Vogel Europa ’s, mit besonderer Riicksicht auf Deutsch-
land [Complete handbook of the natural history of the
birds of Europe, with special emphasis on Germany].
Only Volume 1 (1 834a) on the landbirds was published
(Fig. 30). In it Gloger followed the research findings of
C.E. Nitzsch, in that he distinguished between “Singing
passerines. Aves passerinae melodusae’' and “Passerines
without a larynx. Aves passerinae anomalae’\ Therefore
for the first time in a generalized handbook the
hirundines and the swifts appeared well separated from
each other. Several anonymous contemporary reviews
of this excellent publication were very positive, prais-
ing the richness of the contents and looking forward to
the appearance of the second volume on the waterbirds
(see the journals Reportorium der gesammten deutschen
Literatur (Qd. E.G. Gersdorf), Vol. 3, 1834, p. 318; Kri-
tische Blatter fur Forst- und Jagdwissenschaft (ed. W.
Pfeil), Vol. 9, 1835, pp. 51-54; Isis (ed. E. Oken), 1835,
columns 413^16). Gloger summarized general aspects
concerning the distribution of European landbirds in an
early lecture (1832a). Volume 2, however, was never
published.

(2) Andeutungen zur zoologischen Geographie, mit beson-
derer Anwendung auf die Verbreitung der Vogel [Sug-
gestions towards a zoological geography, with special
reference to the distribution of birds]. An announcement
of this book appeared on page 2 of the prospectus and
invitation to a subscription to the Handbuch mentioned
above, issued by the book dealer August Schulz & Co.
(Breslau) and dated 1st November 1833. Gloger unfor-
tunately never finished the manuscript of this work and
it was never submitted to the printers. It would have
been the first global overview of the zoogeography of
birds. Only the brief summary of a lecture by Gloger
on the topic ever appeared (1834b). In it he states that
the number of bird species and genera rapidly declines
moving from the tropics to the higher latitudes, but that
the area of distribution of individual species is greater
in these higher latitudes than in the tropics, also that wa-
terbirds have greater ranges than landbirds in the main,
and the number of nocturnal birds increases towards the
equator. Woodpeckers are absent only in the forests of
New Guinea (because of “the smoothness of the bark
and hardness of the wood of the native trees”, accord-
ing to Gloger’s historically interesting ecological expla-
nation!).

(3) Gemeinniitziges Hand- und Hilfsbuch der
Naturgeschichte [Practical handbook and manual of
natural history], which was intended to be a represen-
tation of the entire animal kingdom (though mostly of
all mammals and birds). Once again only Volume 1 of
this handbook appeared after a considerable delay: 1841
(pp. 1-400) and 1842 (pp. 401-496 and pp.
I-XXXXIV). On p. Ill of the Foreword to this handbook

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Jurgen Haffer (t) et al.

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i>er ^

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ffiilplKbt tfi JtlifirliAnl tr{1i5lHp«if*i^Jire!iHL[4m' llMtvmi* tn SiluTlItlrl^ Unb ISrr filrffirfB
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©teS[au 1834,

Bti €Sctiu(j utib ^antp,

AvtEuilt hi nrof:. bh^

Fig. 30. Title page of Gloger (1834).

and manual Gloger explains that he had been unable to
make headway with several ongoing projects because
of an eye ailment, but has been able to dictate this par-
ticular manuscript without carrying out further research.
Even if various health problems in the 1 830s had inter-
fered with his work, it remains difficult to understand
why he didn’t turn to the completion of these three un-
finished books later in life. What was the reason for this
strange behavior?

Our study of unpublished material in the Geheimes Staat-
sarchiv [State Archives] in Berlin has shown that after the
beginning of 1 834 Gloger ’s scientific endeavors were sud-
denly diverted towards a completely different subject,
which from then on would command his full attention
(Haffer & Hudde 2007). It was for this reason that he aban-
doned the projects outlined above. From then on his en-
ergies were dedicated to his work on what he called
Gloger ’s Naturliches System der Thierwelt [Gloger ’s nat-
ural system of the animal kingdom^ a theoretical construc-
tion built on natural philosophy which he imagined as the

Bonn zoological Bulletin - Supplementum 59; 1-116

high point of his intellectual life and career. This was now
his life’s work, for which, at the age of only 30, he was
to sacrifice his extremely promising ornithological future,
as we must confirm in retrospect.

Grammar school teacher in Breslau. In July 1830, the
same year in which he gained his PhD, Gloger was ap-
pointed to the position of natural history teacher at the
Matthias Gymnasium in Breslau. However he regarded the
job only as a transitional occupation until he would be able
to take up a post as university lecturer or professor of zo-
ology. This apparently became so obvious to his superi-
ors that a few years later they recorded that Gloger “is not
suited for this school and can in no way be commended
in the conscientious undertaking of his teaching duties or
in his official diligence. He gives nothing in return for his
remuneration” and shows a “complacent evaluation and
self-conceited notion of his own worth” (Provincial
School Board of Breslau, 9 August and 14 October 1 836;
personal file on Gloger, Staatsarchiv). Again and again
during the 1 830s Gloger wrote petitions for the financial
support of his scientific research and applied unsuccess-
fully for the position of associate university professor.
These documents can still be studied in the Berlin State
Archives, in the file on Gloger from the “Royal Ministry
of Spiritual, Educational, and Medical Affairs”. In Sep-
tember 1 842, when the minister concerned. Dr. Eichhorn,
pressured by Gloger’s insistence, finally approved a 3-year
stipend of 600 Reichsthaler per year “towards the further-
ing of his projecf ’, Gloger happily left his permanent, yet
detested post at the Breslau grammar school on 3 1 De-
cember 1842. He immediately moved to Berlin as a pri-
vate scholar “in order to be able to devote all my time and
energy to the service of science in a place with access to
the required literature” (Gloger on 25 March 1861; Rep.
87 B, No. 19998, Sheet 287; Staatsarchiv). This “service”
referred to the long-promised elaboration of “Gloger’s
Natural System of the Animal Kingdom”.

Natural philosophy as the supposed highpoint of
Gloger’s life and work; the role of Professors Steffens
and Nees von Esenbeck as Gloger’s teachers. Between
ca. 1 800 and 1 840 many German biologists were influ-
enced in their thinking by the Romantic Zeitgeist and by
the idealistic natural philosophy of Friedrich Wilhelm
Schelling and Eorenz Oken. They taught the unity of na-
ture and the human mind, believing that what was required
was “to show the genesis of the world out of the human
mind [. . . ] An entirely new perception of Nature must arise,
based on Idealism” (Kiihn 1948: 216). They believed that
deductive knowledge of nature was possible and that the
genesis of the animal world may be understood through
theoretical thinking without laboriously assembling an in-
ductive basis of observed facts. When the scientific un-
tenability of such notions was later recognized there was

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much complaining about “the pointless waste of great en-
ergies and valuable time” and “Schelling’s corrupting in-
fluence” on the biology of the day in such works as Ideen
zur Philosophie der Natur (1797) [Ideas towards a phi-
losophy of nature^ or System des transzendentalen Ideal-
ismus (1800) [System of transcendental idealism].

As pointed out above, since the early 1830s Gloger too
had fallen under the influence of this biological Zeitgeist,
having clearly been “seduced” by the teachings of his pro-
fessors at the University of Breslau, above all Henrich
Steffens (1773-1845) and C.G.D. Nees von Esenbeck
(1776-1858), both prominent representatives of the nat-
ural philosophy school in Germany. The former taught in
Breslau between 1811 and 1832, when he was called to
Berlin. He saw Man as a living unity of mind and nature,
and one of his published works dealt with the main prin-
ciples of “philosophical science”, a glorification of
Schelling’s worldview (Liebe, in Hennicke 1893). “Be-
cause of Steffens’ almost paternal affection for me from
the very beginning, I have been completely indebted to
him,” Gloger wrote on 3 April 1 830 to Profesor Lichten-
stein in Berlin, and again on 26 September of the same
year: “Prof. Steffens, who shows the interest of a father
in me and who does everything in his power for me”. Al-
so in 1 830, Steffens contacted the Minister of State with
a recommendation for financial support for Gloger, say-
ing that he saw in him “a most excellent future teacher of
natural history” (Staatsarchiv).

The well-known natural philosopher Nees von Esenbeck
also taught at Breslau University from 1 830 to 1 85 1 . “He
has always been meticulous, more meticulous than any-
one else, has known about my project from the very start
[= 1 834] and has most attentively followed its progress”,
wrote Gloger (Letter 1 to the Crown Prince in 1838; Staat-
sarchiv). The natural philosophy of his professors’ lectures
and writings had heavily influenced Gloger’s thinking
since the early 1830s.

Work on Gloger’s Natural System of the Animal King-
dom. On 7 January 1 834, the 30-year-old Gloger believed
that he had been granted “an enlightened inspiration by
Providence'^ concerning the discovery of the system un-
derlying all of nature”, which he regarded as the “hoped-
for turning point for the better” in his life. He immediate-
ly started on a new undertaking, “which consists of a large-
scale zoological systematics and will contain the unexpect-
edly rich results of a happy idea in a moment of illumi-
nation” (letters 1 and 3 to Crown Prince Friedrich Wil-
helm in 1839"^).

Gloger felt that he had been found worthy of “a discov-
ery far more influential than any granted to previous nat-
uralists”. Providence had chosen him “to achieve for the
first time not only the most difficult task but one that had
been thought to be almost impossible!”. He felt himself

Bonn zoological Bulletin - Supplementum 59: 1-116

to be “a weak tool for such an important task, though one
chosen by a higher power and equipped at least with the
best of intentions”. Ultimately he saw in his system a
“proof of the necessity of the existence of God” (letter 3
to the Crown Prince).

In his applications for support to the Prussian Acade-
my of Sciences in Berlin in October/November 1835 and
January 1838, Gloger explained the principles of his sys-
tematics, which he said would have to be in agreement
with “simple empiricism as well as a refined natural phi-
losophy” {Staatsarchiv).

The position of individual animals or forms in the sys-
tem, either “lower” or “higher”, is always determined by
“one and the same numerical principle”. All forms occu-
py a definite and unalterable position, and in this the use
of particularly problematical taxa is essential, since their
position makes that of neighboring taxa evident. The sys-
tematic arrangement never follows a simple line but in-
stead is constructed of ever smaller parts of largely par-
allel and analogous lines of forms of equal number, and
this is true for genera as well as species and varieties, and
hence an arrangement in the form of tables is best suited
for depicting the natural system.

In addition, from a step-wise arranged natural system,
Gloger was hoping to find the underlying formative rules
{ursprungliche Bildungsgesetze). The natural system and
the history of nature would then be perfectly congruent
and be as one.

According to the explanations in his letter to the Crown
Prince of 11 December 1838, this natural classification
“will have to develop into a calculation with forms (here
organic structures) similar to the role played by algebra
in connection with mathematics, a calculating and con-
structing in quantities and forms. It has in common with
those two that a majority of forms and factors needs to be
known in order to allow the calculation and deduction of
those entities that are still unknown”.

Following his “enlightened inspiration” of January
1 834, whose exact nature he never revealed, as early as
29 March of that year Gloger presented the Ministry of
Spiritual, Educational, and Medical Affairs with a tabu-
lar overview of the vertebrates and of the entire animal
kingdom based on his “new method”. Lichtenstein re-
ceived a letter on 1 8 June 1 836 accompanied by special
tables on the system of mammals, listing essential gen-
era, while on 6 July the Prussian minister von Altenstein
was sent five tables of the mammals glued together plus
a universal table of the entire “world system”, which can
be inspected today in the Department of Ornithology of
the Berlin Museum of Natural History. For a detailed ac-
count see Glaubrecht & Haffer (2010), which forms the
basis of much of the remainder of this chapter.

Gloger’s Universal Table of the World-System divides
Seyn [Being or Existence] into Schopfer (Ursache) [Cre-
ator (Cause)] and Welt (Wirkung) [Cosmos (Effect)]', the

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latter consists of (a) systems of fixed stars and
galaxies/nebulae and (b) planets with no life and planets
with life, i.e. the Earth. The zoological tables carry the gen-
eral title Gloger ’s Natural System of the Animal Kingdom.
The mammals for example are divided into non-walking
(flying and swimming) and walking (terrestrial) mammals.
The first group includes those with wing membranes (i.e.
bats) and flipper feet (seals, whales, etc.). The terrestrial
mammals are classed as higher terrestrial mammals and
lower terrestrial mammals, distinguished from each oth-
er by (e.g) complete or incomplete rows of teeth, presence
or absence of hooves, and similar morphological criteria.
A detailed main table contains “mammals down to all gen-
era”. These tables are similar to dichotomous identifica-
tion keys, but fail to make the huge significance given to
them by Gloger clear to a modern observer. But above all
it is impossible to discern how he actually utilized his “nat-
ural philosophical numerical principle” to arrive at this
classification.

Other natural philosophers at this time also predicted the
existence of undiscovered genera or species on the basis
of certain “numerical principles”, such as 4 or 5 genera
per family or species per genus (Stresemann 1951:
1 84-1 85). But Gloger’s “calculation with forms”, his pre-
dictions, and systematic categorization of unknown taxa
are not closely explained and his method cannot be
gleaned from the study of his tables. The only concrete
example in the substantial bundle of letters and manu-
scripts surviving is the family of “toothed” pangolins,
whose existence Gloger said he had “worked out” using
his system before their discovery (letter to Lichtenstein
of 14 February 1838).

These unpublished explanations of his system by Gloger
himself, only briefly summarized here, are marked by
much pathos, immense long-windedness, and few com-
prehensible facts. Hence his requests for financial support
were often met with skepticism and reservations. On 18
August 1 842 the Prussian Academy of Sciences wrote to
the minister responsible about Gloger’s plan to work on
the systematics of vertebrates, pointing out that his idea
was based on the personal error that the apparent success
of his treatment of a small part of the mammals permit-
ted him to be optimistic and to immediately expand his
research to the entire animal kingdom. Herr Gloger ought
to rather employ his excellent observational talent,
“which has enabled him to gain outstanding knowledge
in the area of the higher animals, in specialized studies,
namely the completion of his natural history of European
birds”. To the Academy it appeared “dubious to vouch in
advance for the alleged success of his enterprise, which
after all only aims at a formal framework” (Staatsarchiv).
The zoologists of the Academy, among them C.G. Ehren-
berg in particular, had seen through the emptiness of
Gloger’s project on natural philosophy.

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Gloger apparently continued to work on his ambitious
plans for a system encompassing the whole of the animal
world until at least 1 850, since in that year he wrote to
J.F. Naumann, informing him that he was still occupied
with his “natural system” and so would be doing no fur-
ther work on the second volume of his European hand-
book (Moller 1972: 57-58). The increasing criticism of
natural philosophy throughout Germany in the 1 850s prob-
ably made his situation ever more difficult, so that Gloger
finally abandoned his dream of a “Natural System of the
Animal Kingdom”.

Work on pest control and animal conservation. Gloger’s
theoretical studies based on natural philosophy produced
no concrete results, and at the end of 1 845 his stipend was
exhausted. When the Ministry refused any further pay-
ments he was forced to think of some way of providing
an income for himself, and started to write papers on pest
control and the protection of economically useful animals
to sell to the Ministry of Agriculture, which did indeed pro-
vide him with small and irregular payments for a certain
length of time. He then turned to writing on bee-keeping
and the cultivation of hemp, on game protection and agri-
culture, and to papers on damage caused by vermin, es-
pecially voles and mice, the conservation of birds, in par-
ticular hole-nesters, and other useful animals (Haffer &
Hudde 2007).

In doing this Gloger acted in the interest of the balance
of nature and for the conservation and protection of all an-
imals useful for this balance, so he was one of the
founders of an advanced ecological form of animal con-
servation (Barthelmess 1981). His suggestions of howto
combat pests through artificially increasing the numbers
of their natural enemies were based on ideas of Professor
Lichtenstein, who, in his unpublished Bemerkiingen zu den
Berichten und Gutachten uber die Feldmause am Rhein
im Jahre 1822 [Remarks on the articles and reports about
the voles on the Rhine in the year 1822] (1823) had writ-
ten:

“Without doubt the natural methods, i.e. those most im-
mediately supplied by nature, are best [...] They consist
primarily in the deliberate encouragement of an increase
in numbers of the natural enemies of the voles. ” It ought
therefore to be “deprecated in the extreme to shoot a Spar-
rowhawk. Goshawk, Buzzard, Kestrel, or owl. [...] An in-
crease in polecats, martens, and weasels would do a great
deal towards the eradication of the voles. They should be
caught in an organized fashion and released in the fields
where the voles are doing damage ” (Rep 87 B, No. 19998,
pp. 27-30; Staatsarchiv)

The response in Germany to Gloger’s popularly written
articles in daily newspapers and in agricultural and forestry
publications was divided, though abroad it was more

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Fig. 31. Portrait of Constantin W.L. Gloger in 1862 with the
Order of St. Stanislaus (from Glaubrecht & Haffer 2010; Mu-
seum fur Naturkunde Berlin, Historische Bild- und Schriftgut-
sammlungen).

friendly. In 1 862 the Russian Czar even awarded Gloger
the St. Stanislaus Order 3rd Class for his work on pest con-
trol (Fig. 31).

Return to ornithology: biology of birds. When Professor
Jean Cabanis, ornithologist at the Museum of Natural His-
tory in Berlin, founded the Journal fur Ornithologie in
1852 (the first issue appeared in 1853) he invited Gloger
to be a collaborator, doubtless paying him a small fee for
his help and for his numerous contributions (a total of 128
after all!) to this new journal during the last ten years of
his life. He wrote on some of his observations in Silesia
decades earlier as well as on various aspects of avian bi-
ology in general, stimulated by his study of the ornitho-
logical literature. He apparently lacked the money even
to undertake excursions or make field observations in the
surroundings of Berlin.

Among the subjects he dealt with in the first year of the
Journal were the acrobatic climbing ability of Little Bit-
tern in its reedbed habitat. Northern Pintails swimming just
below the water surface, or hybridization between duck
species. In the following year (1854) there were articles
on various aspects of the reproduction of the Common
Cuckoo, the hybridization of Western Capercaillie and
Black Grouse, the attraction of shiny objects for some
birds (which Gloger thought could be connected to their
diet, e.g. metallically iridescent beetles), plus remarks on

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the food of Great Spotted Woodpecker and gulls dropping
hard-shelled mollusks on rocky ground (1855). In an in-
teresting note he drew attention to the fact that (accord-
ing to Nilsson) young curlews can already hear, recognize,
and respond to the calls of their parents while they are still
in the egg (1856), and later in the same year he wrote about
family bonds in birds and the supplementary molt in the
Red Grouse Lagopus 1. scotica in Scotland. In later years
he discussed bird migration across the sea (1857) and
“cock nests” in birds (1859). Gloger explained the greater
length of wing and tail feathers in immature Golden and
White-tailed Eagles by their need for more effective flight
feathers than the adults because they move around so
much, while older birds become increasingly sedentary
(1860); for further details see Haffer & Hudde (2007).

But the small incomes he received for his contributions
to the Journal fur Ornithologie and his papers for the Min-
istry were eventually insufficient to cover even the most
basic necessities. Ratzeburg (1868: 201, footnote; 1874:
81, footnote) rather unfairly described Gloger during that
decade as “having gone astray”, and of living “idly” or
“indolently”. Gloger attended at least two Deutsche Or-
nithologen-Gesellschaft conferences, 1851 in Berlin
(where he was even reconciled with his old adversary C.L.
Brehm, thanks to J.F. Naumann acting as go-between) and
1856 in Kothen (where he delivered a verbose lecture on
the definition of species, though the conference was un-
able to agree on the taxonomic limits of the species tax-
on), but otherwise lived a very secluded life as a lonely
private scholar and bachelor, having failed in his three-
decade long attempt to classify nature. (Moller 1972: 82)
related his sad end: “He lodges with Widow Schulze in
Berlin [-Charlottenburg], Mauerstr. 80, and she, along with
her son, sometimes has to support him.” Finally his pover-
ty was compounded by sickness and Gloger died on 30
December 1863 of “abdominal dropsy”.

C.W.L. Gloger ’s approach to species - even if he regard-
ed them as reproductive communities - was essentially ty-
pological and he delimited many bird species in polytyp-
ic fashion, which was quite in contrast to other ornithol-
ogists of his time. In what we would call today “lumping”,
he subsumed many of the geographically variable climat-
ic varieties (subspecies) of birds within one species, hence
he argued against taxonomic names for subspecies. This
caused later authorities, for example Stresemann (1951:
72), to judge Gloger ’s view as having led systematic or-
nithology into a dead end, whereas we propose that his
approach stimulated fruitful and ultimately helpful discus-
sions on species taxa and concepts and on the practical
question of how to delimit species. Both the synthesis of
similarities as well as the analysis of geographical differ-
ences between populations are necessary and lead to fresh
knowledge in ornithology. The historical significance of
Gloger’s early ornithological work is now being general-
ly acknowledged (Haffer 1992, 1997b, 2001, 2006; Haf-

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fer & Hudde 2007). In his later life his tragic struggle with
classification holds both a message and a warning. High-
lighting the danger of fatal error and complete failure, his
Natural System of the Animal Kingdom stands as one of
the many alternative classification schemes and attempts
of the 19th century, representing systematists’ continuous
endeavor to bring order into the seemingly chaotic assem-
blage of animal taxa. For a more detailed account, includ-
ing illustrations of Gloger’s system and tables, see
Glaubrecht & Haffer (2010).

(7.) Ornithology of the late 19th and early 20th cen-
tnries

(7a.) Fannistics and life histories of birds

Bird collections and natural history museums. Bird col-
lections in museums form the basis for systematic and zoo-
geographical studies, for research on individual and geo-
graphic variation, studies of plumage color patterns, bio-
diversity and other topics. Public and private bird collec-
tions were very important during the period when ornithol-
ogy originated as a separate branch of science. Ornithol-
ogists of the early 1 9th century who owned bird collec-
tions were, among others. Count Hoffmannsegg near Dres-
den, Bernhard Meyer in Offenbach, and Christian Ludwig
Brehm in Renthendorf (Thiiringen).

When the University of Berlin was founded in 1810,
Count Hoffmannsegg presented his collections of birds
and insects to the new Museum of Zoology (under Carl
Illiger and later M.H.C. Lichtenstein) as a foundation on
which to build in future years. Similary, Bernhard Mey-
er’s collection formed the basis of the Senckenberg Mu-
seum in Frankfurt a.M. under P.J. Cretzschmar. In 1825,
four other public centers existed besides those mentioned
above: Darmstadt (under J.J. Kaup), Munich (under J.B.
Spix and J. Wagler), Dresden (under L. Reichenbach), and
Halle (under C.L. Nitzsch). These and other museum col-
lections in Germany grew steadily during the 19th and
20th centuries, especially after Germany had established
colonies in Africa, New Guinea, and the Pacific Ocean
during the early 1880s. These colonies were lost with the
outcome of the First World War. In 1916, the Senckenberg
Museum was able to purchase Count von Berlepsch’s col-
lection (built up from 1860 to 1910) comprising 55 000
specimens, the largest private bird collection ever assem-
bled in Germany. Other private collections were those of
O. Kleinschmidt (now in the museums of Bonn and Dres-
den), V. von Tschusi zu Schmidthoffen (now in the Vien-
na museum), and the old collections of Prince Maximil-
ian zu Wied-Neuwied (purchased by the American Mu-
seum of Natural History, New York; a total of 4000 spec-
imens), and of C.L. Brehm (now in the museums of New
York and Bonn; the original total was 9000 specimens).

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Scientists and commercial collectors from central Europe
traveled widely overseas, contributing to the growth of the
regional knowledge of the avifaunas of the world (Stre-
semann 1975; Mearns & Mearns 1998; Hinkelmann
2000). See Fig. 32.

As listed by Mearns & Mearns (1998), the German col-
lections among the important museum collections in Cen-
tral Europe today are of the order of 60 000 bird speci-
mens (Munich, Stuttgart), 75 000 (Bonn, Dresden), or 90
000 (Frankfurt a. M.). Only the collections in the Muse-
ums of Natural History in Vienna and Berlin reached to-
tals of 100 000 and 140 000 bird specimens respectively.
Additional museum collections are those in Braun-
schweig (30 000), Hamburg (30 000), Basle (26 000), Bre-
men (20 000), Halberstadt (Museum Heineanum, 1 8 000),
Kothen (Naumann Museum), Halle, and Bern. Besides the
bird collection of the Zoological Museum of the Univer-
sity of Hamburg, an important private “South Sea Muse-
um” of the wealthy merchant J.C. Godeffroy existed in the
city from 1861 until 1 885. It was founded to augment the
knowledge of ethnology and zoology of Oceania. G. Hart-
laub and O. Finsch described (often in the Journal des Mu-
seums Godeffroy 1871-1879) the many ornithological
novelties which professional collectors and the captains
of Godeffroy’s merchantmen sent home. After the com-
pany experienced economic difficulties this museum was
closed and the valuable collections were auctioned off in
1885 and scattered all over the world (Stresemann 1975:
229). The following publications deal with the history of
the main bird collections in Germany mentioned above:
Munich (Hellmayr 1928; Reichholf 1992), Bonn (Rhein-
wald & van den Elzen 1984), Dresden (A.B. Meyer 1 897),
Frankfurt a.M. (Naumberg 1931; Steinbacher 1967),
Berlin (Stresemann 1922; Ahrens 1925; Mauersberger
1994a), Braunschweig (Blasius 1897; Boettger 1954; Ha-
jmassy 1983), Bremen (Duncker 1953), Halberstadt
(Quaisser & Nicolai 2006; Nicolai et al. 2009), and Halle
(Taschenberg 1894; Piechocki 1971). See also (e.g.)
Rheinwald (2003), Roselaar (2003), and Steinheimer
(2006). The sizes of these museum collections are mod-
est compared to the collections of the British Museum in
Tring or of the American Museum of Natural History in
New York, each of which contains about 1 million spec-
imens. However the old European collections are very im-
portant scientifically because they contain the type spec-
imens on which many species and subspecies of birds are
based. Besides the “skin” collections which preserve the
skin and plumage, bill and feet of a bird, museums have
also built up collections of skeletons and of birds preserved
in alcohol serving as research material for anatomical and
functional-morphological studies.

Oology. The collection and study of avian eggs was quite
fashionable among private collectors in Europe during the
19th century, and a large series of well-illustrated books

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Total bird collGCtton Total bird collection

Fig. 32. Growth of the bird collections of some museums in Central Europe. Numbers indicate major collections incorporated
into the respective museums: 1 -Natterer; 2 - “Novara”; 3 - Reischek; 4 - Tschusi and Grauer; 5 - Schiebel; 6 - Seilern; 7 - Rup-
pell; 8 - Berlepsch; 9 - Erlanger; 10 - Kleinschmidt; 11 - C.E. Brehm; 12 - indicates losses during and immediately after the end
of the Second World War (from Haffer 2001, which see for sources).

on the subject has been published (Zorn 1742, 1743; Klein
1766; Gunther 1772; Naumann & Buhle 1818-1828;
Schinz 1819-1830; Thienemann etal. 1825-1838; Thiene-
mann 1845-1856; Morris 1853-1856; Baedecker et al.
1855-1867; des Murs 1860; Graessner 1860, 1880; Wol-
ley & NewTon 1864-1907; Reichenau 1880; Oates &
Hume 1889; Poynting 1895-1896; Seebohm 1896;
Nehrkom 1899; Rey 1899-1905; Oates & Ogilvie-Grant
1901-1912; Dresser 1905-1910; Krause 1905-1913; Jour-
dain 1906-1909; Pelt Lechner 1910-1914; Koenig
1931-1932; Kobayashi & Ishizawa 1932-1940; Helle-
brekers 1949; Hoogerwerf 1949; Matousek 1956; Schon-
wetter 1960-1992; Verheyen 1967; Makatsch 1974,
1976; Cramp etal. 1977-1994; Mikhailov 1997). Schon-
wetter’s 4-volume Handbiich der Oologie (1960-1992) is
more satisfactory than previously published ones. This,
however, does not make him the “founder of scientific ool-
ogy”, as Piechocki (1999) stated in his title without any
Justification to substantiate his claim. The founders are
rather among the oologists of the mid- 18th century listed
above (Nitze 2000).

The Journal fur Ornithologie regularly carried scientif-
ic notes and articles on (e.g.) the individual and geograph-
ical variations in egg color in particular species, the char-
acteristics of the egg of the brood-parasitic Common
Cuckoo (relatively small size, strong shell, similarity in
color to the host’s eggs), differences in the structure of the

eggshell in closely related species, the eggs of bird species
from overseas and other topics. Blasius (1861) discussed
at a DO-G meeting the question whether or not birds’ eggs
have species-specific characteristics. The first discussions
of the significance of measurements and weights of eggs
were those of Reichenow (1870) and Nathusius (1882).
The latter’s studies on the structure of eggshells in differ-
ent groups of birds were combined and republished in an
English translation almost a century later (Tyler 1964). An
oological Journal, Zeitschrift fur Oologie und Ornitholo-
gie, appeared from 1891 to 1924 (volumes 1-29). Some
oologists published heated discussions in the Journal fur
Ornithologie (1877-1879) about the applicability of Dar-
win’s theories, especially his proposed mechanism of nat-
ural selection, to the evolution of the color and shape of
birds’ eggs (F. Kutter for and W. von Nathusius against;
see also Kutter 1889 and Hartert 1890). Some of the cur-
rent uses of extant egg collections are display, research,
and identification as discussed by Walters (1994). Rahn
et al. (1985a, b; 1988a, b; 1989a, b) analyzed and evalu-
ated Schonwetter’s data from various oological and phys-
iological viewpoints. The gradual thinning of eggshells of
several common European species during the last few
decades due to environmental influences (DDT) was doc-
umented on the basis of historical egg collections (Green
1998).

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Behavioral observations. Another controversy among
Central European ornithologists from the late 1 860s on-
ward was sparked by the evolutionary and anthropomor-
phic views of animal behavior as expressed by Alfred Ed-
mund Brehm (1829-1884, the son of Christian Eudwig
Brehm, in his popular books on Das Leben der Vogel
(1861, 1867) [The life of birds], Thiere des Waldes,
Wirbelthiere (1864) [Forest animals, vertebrates], Illus-
triertes Thierleben, Vogel (1866-1867) [Illustrated ani-
mal life, birds], and Brehms Thierleben, Vogel
(1876-1879) [Brehm ’s animal life, birds]. His adversary
was the zoology professor Bernard Ahum (1824-1900),
who countered Brehm ’s interpretation in his book on Der
Vogel und sein Leben (1 868) [The bird and its life]. Both
ornithologists made excellent observations on bird behav-
ior but differed profoundly in their interpretations. Natu-
ral-theological (Ahum) and anthropomorphic views
(Brehm) were both held by many scientists in Europe from
the 18th into the 19th centuries without major disputes.
It was the great popularity of Brehm’ s Animal Life which
provoked Ahum into writing his teleological and anti-Dar-
winian book, leading to a vigorous public debate. Brehm
himself (1868, 1876: 20-24), A. & K. Muller (1868,
1890), and K. RuB (1868) attacked Ahum, the latter an-
swering these criticisms in later editions of his book. The
discussions on Instinct and deliberate actions of higher
animals by A. & K. Muller (1869: 8-16) indicates that
their views were less extreme than appears from their high-
ly emotional book review (1868). They, in fact, did ac-
cept the existence of instinctive behavior in animals that
has remained constant for “an unimaginable time” and
stated that “instinct as a cogent law dominates the entire
animal world”. They went one step beyond Ahum when
they claimed that “instincts are peculiar not only to the
animal’s soul but even man acts at times instinctively”.
However, they continued, at least higher animals also act
to a higher or lesser degree deliberately. The animal’s free
self-determination proves the indubitable relationship of
its soul with that of man (see also A. & K. Muller 1890:
69-81).

A.E. Brehm, who adhered to a Darwinian view of evo-
lution, was a great popularizer of the study of animal life.
The two main aspects of his work were (1) his emphasis
on the habits of animals and (2) his anthropomorphic in-
terpretation of animal behavior. In his writings Brehm fre-
quently cited his own observations of birds, also those of
his father C.E. Brehm and of J.F. Naumann among Ger-
man ornithologists, as well as the writings of naturalist
travelers overseas like Heuglin, Audubon, Radde, Gould,
Burmeister, Wallace, Azara, Darwin and many others. He
combined his personal expedition experience in northeast-
ern Africa, Spain, Norway and later in Siberia with that
of many other adventurers, writing brilliantly and enthu-
siastically about animals as living beings in their environ-
ments in a style accessible to the general public. His pub-

Bonn zoological Bulletin - Supplementum 59: 1-116

Usher distributed Brehm’s book in several languages and
in large editions worldwide.

Brehm’s anthropomorphic interpretation of animal be-
havior was derived from the romantic natural philosoph-
ical views prevailing among many European zoologists
of the first half of the 19th century (e.g. C.E. Brehm, E.
Oken, J.F. Naumann, R Scheitlin; see Stresemann 1975:
319, Jahn & Wolf 1979, and especially Schulze 2009). Ac-
cording to Brehm, in their actions animals reveal “sym-
pathy”, “compassion”, “love”, “hatred”, “gratitude”,
“vanity”, “sense of honor”, “pride”, etc. He disliked the
predominantly analytical academic zoology. The nature of
the ornithological accomplishments of father and son
Brehm during the first and second halves of the 1 9th cen-
tury were totally different from each other, and a direct
comparison of their writings, their significance and influ-
ence, as attempted by some authors (e.g. Dathe 1989),
would seem impossible. Brehm senior was a researcher
and systematist who addressed a relatively small group of
colleagues. As mentioned above, he also published many
functional observations on birds (Stresemann 1975: 302),
discussions of their plumages and color changes, and he
discovered the specific distinctness of the members of sev-
eral pairs of sibling species among the birds of Europe:
(Galerida theklae/G. cristata, Parus salicarius/P. palus-
tris [= Poecile montana/Poecile palustris], Regulus ign-
icapilla/R. regulus, Certhia brachydactyla/C. familiaris).
Although the names of G. theklae and R. ignicapillus were
introduced into the literature by his son A.E. Brehm (1857)
and his colleague J.C. Temminck (1820) respectively, the
specific distinctness of these two species had been estab-
lished by the elder Brehm. This is clear in the case of the
Regulus species (where Temminck referred to the manu-
script of C.E. Brehm), but less so in the crested larks. In-
direct evidence is provided by the labels of the type spec-
imens of G theklae which J.H. examined at the Ameri-
can Museum of Natural History (New York). These birds
had been identified by A.E. Brehm in his pencil handwrit-
ing as “G undata'' (= G cristata)', C.E. Brehm crossed out
his name on the labels and wrote in his characteristic hand-
writing with black ink '"Theklae". With respect to ornithol-
ogy as a science, Christian Eudwig Brehm’s fame far ex-
ceeds that of his son, even though the latter’s books have
been read by many more people than those of his father.

In his book Der Vogel und sein Leben (11th edition
1937) Bernard Altrum opposed Darwinism and A.E.
Brehm’s - and many other authors’- anthropomorphic
views of animal behavior (Kraus 1914). As a natural the-
ologian he defended a teleological interpretation of a bal-
anced harmony of creation but, with respect to the behav-
ior of animals, he had the merit of reintroducing into or-
nithology the concept of instinct and innate behavior pat-
terns. Ahum insisted that when a bird is singing it does
not express its feelings and has no conscious intention.
“The animal does not think, does not reflect, does not es-

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tablish aims for itself and if it nevertheless behaves pur-
posively, then someone else must have thought for it” (An-
imal non agit, sed agitur, “an animal does not act, but is
acted upon”, or more freely “An animal does not act by
its own volition, but reacts to stimuli [drives]”; Mayr
1935). Altum thought that birds are no machines (as his
opponents understood him as saying) but living organisms
capable of adapting to local conditions to a certain degree.
We emphasize that Altum’s rejections of Darwinism did
not affect his praiseworthy battle against Brehm’s anthro-
pomorphism. Ahum discovered territory in the life of birds
and its adaptive meaning. His ecological studies on the
food of raptors and owls as well as his discussions of the
“economic value” of birds were other important contri-
butions at a time when the bird protection movement was
gathering momentum.

We know today: “Alfred Edmund Brehm rightly stated
that, in their emotions, higher animals may be very sim-
ilar to humans. However, it was not consistent to base on
this similarity his view that the behavior of animals can
be guided by reason and morals, and even better so than
in humans. By contrast, Bernard Ahum perceived correct-
ly that the species-specific instinctive behavior patterns
of birds occur blindly and without any understanding of
their functions. But he did not realize that in humans this
is also partly true and followed Descartes’s opinion of an-
imals being soulless machines. ... Many an exasperated
conflict of opinion has been unmasked, through Charles
Darwin’s insights, as a spurious dispute. Today, total una-
nimity prevails among scientists about his theories”
(Lorenz 1974: XIV; see also 1973: 10). Lorenz also stat-
ed that “Birds are much more stupid than people believe;
however, in their feelings and passions birds differ much
less from humans than many people presume”. Lucanus
(1911, 1925, 1926) and Bolsche (1924) commented sim-
ilarly on the debate between Brehm and Ahum stating that
both were right to some degree, but wrong in many re-
spects; as so often in life the “golden mean” appeared to
Lucanus the correct approach.

The work and publications of Brehm and Ahum repre-
sent the first highlight of behavioral observations of birds
in Europe (besides the pioneering work of Pemau and
Zom that we have already covered). However, neither a
science of animal psychology nor the idea of experimen-
tal field studies existed during their lifetimes. Moreover,
most scientific ornithologists of the late 19th century were
of the opinion that studies of birds in the field were only
“second class” and do not contribute important biologi-
cal data, thereby indirectly discouraging such work, as ex-
pressed for instance in the following well-known quote:

“Popular ornithology is the more entertaining, with its
savor of the wildwood, green fields, the riverside and se-
ashore, bird songs and the many fascinating things con-
nected with out-of-door-Nature. But systematic ornitho-
logy, being a component part of biology - the science of

Bonn zoological Bulletin - Supplementum 59: 1-116

life - is more instructive and therefore more importanf ’
(Ridgway 1901:1-2).

Even though Ahum’s book of 1 868 was read very wide-
ly during those years, and six editions had appeared be-
fore 1900 (five additional ones were issued up to 1937),
none of the Central European ornithologists of the late
19th century picked up his suggestions regarding the in-
stinctive behavior of birds and began to study, describe,
and analyze birds’ activities in detail. German ornitholo-
gy concentrated instead on the study of local fauna, avian
phenology, and old-fashioned biology. Publications like
J. Rennies ’s book on nest building. Bird-architecture
(1844; a German translation entitled Die Baukunst der
Vogel appeared in 1 847), had no influence. In this book-
let the author deviated from the usual systematic treatment
and classified birds on the basis of their differing man-
ners of nest construction (digging, platform-building,
braiding, weaving, stitching, cementing, etc.). Conclud-
ing, the author stated that the instinct which guides the
birds is not blind but is closely related to reason, if not
really the same talent. However the abilities of animals
always remain the same, whereas those of mankind
progress during the course of time.

Bird migration continued to be explained by “inherit-
ed habits” until Gatke (1891) eventually concluded that
migrants act on the basis of instincts, as the teleologists,
including Ahum (1868), had maintained long before (Stre-
semann 1975: 325, 333, 345). The detailed study of in-
stinctive behavior in birds commenced in Germany, when
Valentin Haecker discussed bird song in 1 900 and Oskar
Heinroth spoke about his researches on the behavior of
the Anatidae at the Fifth International Ornithological Con-
gress in Berlin in 1910. In Britain, a new generation of
field ornithologists (e.g. E. Selous, F. Kirkman, and E.
Howard) emphasized the need for detailed observations
of birds’ habits and behavior from around 1901 (Strese-
mann 1975: 342-344; see also the section on the “David
Lack and the New Avian Biology” below).

Increasing emphasis on ecology, behavior, quantifica-
tion of occurrence, and species richness: 1900-1999.
Many ornithologists at the turn of the 20th century be-
lieved that most or all that can be known about European
birds had already been discovered and compiled, especial-
ly in Naumann’s classic handbook (1820-1 860). This mis-
conception led to the publication of a slightly revised edi-
tion of this work in 12 folio volumes between 1897 and
1905. Actually the available ornithological knowledge at
that time represented no more than the first foundations
of study, and served only to outline in a general way the
life histories of the more common bird species in the re-
gion. The literature of the late 19th century usually fails
with regard to the biological details of nest life, ecology,
and social behavior patterns. One man opened a breach
in the walls erected by the Naumann cult, as Stresemann

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Jurgen Haffer (t) et al.

(1975: 345) put it, and he was Oskar Heinroth
(1871-1945), who had pointed out certain shortcomings
inNaumann’s work (Heinroth 1917, 1930). He was inter-
ested in the details of life history and habits of birds, i.e.
things not found inNaumann’s and Brehm’s publications.

The large handbooks on or including the birds of Cen-
tral Europe published during the 20th century are those
of Hartert (1903-1922, three volumes and one supplemen-
tary volume with F. Steinbacher, 1932-1938) with an em-
phasis on systematics, Niethammer (1937-1942, three vol-
umes) and Glutz von Blotzheim & Bauer (1966-1997, 14
volumes). They are characterized by an increasing empha-
sis on quantifying the occurrence of bird species in dif-
ferent areas and documenting ecological and behavioral
patterns of each species based on the rich literature that
had become available. The 14 volumes of the Handbuch
der Vogel Mitteleuropas summarize the entire knowledge
of Central European birds in a highly organized manner
and truly represent a “New Naumann” for the 20th cen-
tury, influencing the preparation of the nine-volume British
handbook edited by Cramp et al. (1977-1994) as well as
that of other handbooks published in neighboring coun-
tries (e.g. Romania and Czechoslovakia); see Schulze-Ha-
gen (2013). Chapters on the grouse, partridges, and allies
have been translated into French and widely used in field-
work on these birds in the French Alps.

The handbooks have been supplemented by numerous
other works on the avifaunas of Germany, Austria, and
Switzerland as well as of many individual German states
(e.g. Krohn 1925; Kuhk 1939; Tischler 1941; Heyder
1952; Klafs & Stiibs (1979, 1987); Rutschke (1983, 1987);
Knorre et al. (1986); Holzinger et al. (1987-). We list here
a selection of important titles in chronological order: Fa-
tio et al. (1889-1956); Kleinschmidt (1905-1937, 1913,
1934); Goeldi (1914); Schnurre (1921); Friderich (1902-
1904, 1923); Heinroth & Heinroth (1924-1933); Groeb-
bels (1938); Noll (1941-1942); Bauer & Rokitansky
(1951); Corti(1952); Berndt& Meise (1959-1966); Stre-
semann, Portenko et al. (1960-2000ff); Glutz von Blotz-
heim (1962); Rokitansl<y (1964); Wiist (1970); Wolters
(1975-1982); Schifferli et al. (1980); Bezzel (1982); Dvo-
rak et al. (1993); Winkler (1999); Bauer & Berthold
(1996); Eck (1996); Schmid et al. (1998); Bauer et al.
(2005). Summaries of quantitative mapping efforts in most
European countries have been edited by Hagemeijer &
Blair (1997). Bezzel (1982) discussed the composition of
the Central European avifauna, its occurrence in the cul-
tural landscape, its long-term dynamics, species richness,
and population densities and their variation on test plots
of different size, while Bairlein (1996) summarized the re-
search results into the physiological ecology and synecol-
ogy of European birds. Other disciplines that developed
during the 20th century are ecomorphology (e.g. Bock
1977, 1994; Eeisler 1977), ethology, the study of orien-
tation behavior in migratory birds, and population biolo-

Bonn zoological Bulletin - Supplementum 59; 1-116

gy (discussed in some detail below). Brief reviews of or-
nithological work after the Second World War with an em-
phasis on the political separation of East and West Ger-
many are those of Neumann (1995) and Rutschke (1998).

General prehistorical and historical aspects of birdlife
in Central Europe are an increase in diversity due to the
settlement of large areas by humans, and the ensuing al-
teration of closed woodland into highly structured mosa-
ic landscapes several centuries ago (Bezzel 1982; Berthold
1990a; Gatter 2000). Declines in the populations of birds
and a reduction in diversity began during the first half of
the 19th century caused by direct persecution and habitat
destruction, leading to a period of accelerated decline due
to almost complete ecosystem destruction by humans in-
to the 20th and 21st centuries. Global warming in the fu-
ture will certainly have a severe effect on the composi-
tion of the Central European avifauna (see e.g. Bairlein
2011; Sybertz & Reich 2011; Worm worth & §ekercioglu
2011). Some predictions may possibly be formulated on
the basis of the results that will be obtained through a study
of the effects of the “Eittle Ice Age” of the 16th century
and other climatic changes on the European bird fauna
(Kinzelbach 1995c).

(7b.) Avian systematics

Microsystematics and speciation. During the 19th centu-
ry, the specialists in the museums of Eondon, Eeiden, Paris
and Berlin by necessity worked with relatively few spec-
imens from far-distant collecting stations. Most of them
described any morphologically different bird as a differ-
ent species, sometimes even male and female of one
species. The widespread occurrence of geographical vari-
ation within species was unknown or little known to these
workers, most of whom applied a narrow morphospecies
concept, (e.g.) G.R. Gray, R.B. Sharpe, RE. Sclater in
Britain, C.J. Temminck in the Netherlands, E.R Vieillot
and R.P. Eesson in France, J. Cabanis and A. Reichenow
in Germany. Intermediate specimens between such
“species” were regarded as hybrids possessing no more
significance than any other abnormal animal. These mu-
seum ornithologists greatly increased our knowledge of
the regional diversity of the avifaunas of the world but
none of them pondered seriously the problem of distin-
guishing real species from local varieties. This was done
clearly and in great detail on the basis of long personal
experience and extensive bird collecting by several Eu-
ropean explorer-naturalists who, during the late 18th and
the first half of the 19th centuries, traveled in Russia,
Siberia, and the Far East, men such as Pallas, von Mid-
dendorff, Eversmann, von Schrenck, Radde, or Seebohm.

Hardly any of these explorer-naturalists had a museum
affiliation in Europe. They published the results of their
studies privately in costly expedition reports with limit-

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81

ed distributions. Therefore their consistent emphasis on
broadly defined polytypic species entities of Eurasian bird
and mammal species, together with their impressive data-
base on geographical variation, did not have the impact
among fellow workers that would have been desirable.
Even though the explorer-naturalists found some support
from Gloger (1833) and Schlegel (1844), the museum spe-
cialists’ view on narrowly defined morphospecies in their
influential publications continued to dominate systemat-
ic ornithology, and the work of the explorer-naturalists fell
into oblivion. Eventually, however, their taxonomic
views prevailed decades later, mainly through the influ-
ence of the explorer-naturalists of North America (e.g.
Audubon, Nuttall), via museum men like Baird, Coues,
Allen, and Ridgway.

Essential istic (typological) views dominated systemat-
ic ornithology in Europe during the 19th century, irrespec-
tive of whether taxonomists applied narrow (Einnaeus
“school”) or wide species limits (Pallas-Schlegel “school”;
see Fig. 33).

It was assumed that a Platonic “type” (essence) under-
lay each species; these were conceived as immutable nat-
ural entities which had independent origins (Mayr 1982;
Haffer 1997a,b). The replacement of these “schools” by
modern Darwinian views through the work of the See-
bohm-Hartert school beginning in the 1 890s caused con-
siderable friction. This is obvious from the contrast be-
tween Hartert and most of his ornithological colleagues
in Britain, as well as between Stresemann (1889-1972)
and Kleinschmidt (1870-1954) in Germany. No such dif-
ficulties developed in North America, where the evolution-
ary Baird-Coues school replaced, but did not overlap with,
the earlier essentialistic Wilson school.

Johann Heinrich Blasius (1809-1 870), Professor of Zo-
ology in Braunschweig, was one of the spokesmen for
German ornithology during the 1860s. Eike most of his
colleagues he was a creationist and taught that an unshake-
able order rules organic nature, as it also rules the worlds
of crystals and stars. Animal species were assumed to be
rigidly delimited and to represent proof of an eternal or-
der from the beginning of the world; no species ever gave
rise to a new species. Hermann Schlegel in Eeiden was
convinced, like C.E. Brehm and the entomologist Schaum
in Germany, as well as Agassiz in North America, that all
geographical varieties had also existed since the beginning
of creation and were immutable (Stresemann 1975). Bla-
sius and Bernard Altum (1824-1900) emphatically op-
posed Darwin’s theories when the young Gustav Jaeger
(1832-1917) dared to discuss these new views at the an-
nual meeting of the Deutsche Ornithologen-Gesellschaft
in Stuttgart (September 1860). Eike Blasius, Altum and
Otto Kleinschmidt (1926) also propagated essentialistic
(typological) views. However they had no followers and
the Pallas-Schlegel school ceased to exist when Klein-
schmidt died in 1954 (Haffer 1997b).

Bonn zoological Bulletin - Supplementum 59: 1-116

Because Kleinschmidt delimited species taxa widely, he
joined forces with the early members of the Darwinian
Seebohm-Hartert school in their struggle against the Ein-
naeus school around 1900. Its early members were Har-
tert himself, Schalow, A.B Meyer, Hellmayr and, a few
years later, Stresemann (Seebohm had died in 1 895). They
applied trinomial nomenclature, designating subspecies of
polytypic species with a third name, as proposed by
Schlegel (1844) and the ornithologists of the North Amer-
ican Baird-Coues school. Others declined the use of sub-
species names, despite the fact that they applied similar-
ly wide taxonomic species limits. They recommended (as
F. Faber and C. Gloger had done earlier in the 19th cen-
tury) describing in words the geographical variation of
widespread species without the application of formal sub-
species names, because they believed that “subspecies” are
merely “climatic varieties” with no genetic basis. On the
other hand, some workers of the Einnaeus school did not
deny the existence of geographical varieties of species in
nature, yet they assigned binominal names to these sub-
species as C.E. Brehm had done decades earlier (Haffer
1992: 120, 1996); for example J. Cabanis of the Zoolog-
ical Museum in Berlin who published on ""Merops men-
talis nov. subsp.” {J. Ornithol. 37 [1889]: 76). It is clear
that the meaning of binomial names as applied during the
19th century varied appreciably between different system-
atists. Zoologists accepted trinominal nomenclature for
subspecies only from the early 20th century onward.

History of the biological species concept. Many ornithol-
ogists and other naturalists of the 19th century applied a
biological species concept based on the interbreeding of
conspecific individuals and reproductive isolation between
the representatives of different species; Gloger, for in-
stance, stated in 1 833 that “What under natural conditions
regularly pairs, always belongs to one species”. By stress-
ing “regularly” he wanted to eliminate complications due
to occasional hybridization. Henry Seebohm in Britain was
the first ornithologist to emphasize geographical isolation
as the conditio sine qua non for speciation to occur, and
he came close to a biological species concept when he
wrote that during geographical isolation two forms may
“become so far separated that, should their areas of dis-
tribution again overlap they will nevertheless not inter-
breed, and the two species may be considered to be com-
pletely separated” (1881: X). See also Wagner (1 889).

The entomologist Eimer (1889:16) said “Species are
groups of individuals which are so modified that success-
ful interbreeding with individuals of other such groups is
no longer possible” (similarly Standfuss 1896: 336, 353).
Two British entomologists, Karl Jordan (bom and educat-
ed in Germany) and Edward Poulton, discussed the na-
ture of biological species in great detail in several articles
published in 1896, 1903, and 1905 (see Mayr 1982). Their
work was in the tradition of Darwin’s concepts of grad-

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03 haffer BZB-emI 02.12.13 13:42 Seite 82

82

Jurgen Haffer (t) et af

North America

Europe

Year

Year

— 2000

B

Evolutionary microtaxonomy

2000 —

iftilill

— 1950

— 1900

— 1850

— 1800

VMmWv

MtHerf

1 1^'

£

r •

Ia*

► 1*1.’

L

fV

f

Jr ■

• ;

Tschusi

-

*■

1950 —

- Klemschmidt

\

Sharpe \

\

Sclalcr '

1900

-Schleget \

Cabants '

Midd

I

Gray I

Bonaparte ( 1850

Gloger

I

Lesson I
I

Temmlnck j
I

Vieillot I
r

Pallas

1800

A1

I

Linnaeus |

Ballon

^Essentialistic microtaxonomy

Fig. 33. Research traditions (“schools”) of systematic ornithology during the 19th and 20th centuries. A. Typological (essentia-
listic) microtaxonomy; A1 . Linnaeus school, A2. Pallas-Schlegel school, A3. Wilson school; B. Evolutionary microtaxonomy; B1 .
Baird-Coues school, B2. Seebohm-Hartert school. The main publishing periods of major ornithologists are indicated symbolical-
ly. Most presently active representatives of evolutionary microtaxonomy are indicated anonymously. Bech. - Bechstein, Bias. -
J.H. Blasius, Midd. - Middeldorff, Nau. -Naumann, Stres. - Stresemann. (From Haffer 1997a).

Bonn zoological Bulletin - Supplementum 59; 1-116

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The Development of Ornithology and Species Knowledge in Central Europe

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ual evolution and speciation through the differentiation of
populations in geographical isolation (allopatric specia-
tion). Ludwig Doederlein (1902) emphasized that - as long
as we remain at a particular time level (in the case of fos-
sils or extant animals) - species are sharply delimited from
one another and produce no fertile hybrids. He also dis-
cussed in detail geographical variations in relation to the
mobility of the species concerned.

Since pre-Darwinian times ornithologists had recog-
nized the need to collect “series” of specimens of a species
from individual localities, i.e. more or less extensive pop-
ulation samples, to understand individual and geograph-
ical variation. Thus by 1 900 the populational analysis of
species was routine for students of mammals, birds, fish-
es, snails and some other groups (Mayr 1980: 127). Ernst
Hartert who - under the influence of Seebohm, the North
American ornithologists, and later K. Jordan - delimited
species taxa broadly, published a list of the biological
species of birds inhabiting the Palearctic Region
(1903-1922). L. Plate (1914) conceived the species “phys-
iologically”, stating that a species comprises all individ-
uals which reproduce together sexually; a common bond
between them facilitates mutual recognition and sexual re-
production. As early as 1919, Stresemann also adopted a
biological species concept: “Forms which have reached
the species level have diverged physiologically to the ex-
tent that, as proven in nature, they can come together again
without interbreeding. Morphological divergence is inde-
pendent of physiological divergence” (1919: 64, 66). In
the case of allopatric taxa, their taxonomic rank as sub-
species or species is to be determined by inference based
upon several auxiliary criteria. In several articles he in-
terpreted the increased variability of certain populations
of Central European birds as due to secondary contact and
hybridization of the parent populations involved. Strese-
mann criticized the recognition of formally named sub-
species in continuously (clinahy) varying populations of
birds. During the 1920s his investigations of polymorphic
species (where he labeled the morphs as “mutations”)
linked ornithology with genetics. He treated species as ag-
gregates of populations which often vary clinahy and dis-
cussed population phenomena, such as the population con-
tinuum zones of secondary intergradation and geograph-
ical isolates. Through this work, Stresemann (followed by
B. Rensch and E. Mayr) introduced the methodology of
the “New Systematics” into ornithology during the 1920s
and 1930s, and it was these ornithologists who made the
decisive contributions to the development of the biolog-
ical species concept and the problems of speciation (Mayr
1982; Haffer 1992). From 1913 onward, Stresemann rou-
tinely applied the theory of allopatric speciation, i.e. the
origin of species from small and geographically isolated
populations. He was the first to develop a model of spe-
ciation for European subspecies and species of birds from
populations isolated in Mediterranean refugia during gla-

Bonn zoological Bulletin - Supplementum 59: 1-116

cial periods in the Pleistocene (Haffer et al. 2000). Kre-
mentsov (1994: 36^1) completely neglected the contri-
bution of Central and western European zoologists to the
development of the New Systematics and the biological
species concept when he compared the work of Russian
and “western” (i.e. North American) biologists of the ear-
ly 20th century. His discussions are totally inadequate, par-
ticularly where he claims that many of these achievements
originated with the publications of Russian entomologists
at that time.

The discovery of geographically representative forms
which do not or only rarely hybridize along their zones
of contact, and for this reason represent biological
species (Rensch 1928, 1929), eventually led to a reversal
of the excessive taxonomic “lumping” tendency among
ornithologists, which had reached a peak in Europe un-
der the influence of Kleinschmidt’s (1926) views. Klein-
schmidt delimited very broad Formenkreise (“species cir-
cles” = polytypic species), which often included as sub-
species geographically representative species. The publi-
cations of Rensch (1929, 1934) and later Mayr (1942)
were the first comprehensive statements of population sys-
tematics in Europe.

Comparable to the events in North America, an “Evo-
lutionary Synthesis” between the experimental-reduction-
ist tradition of the geneticists and the observational-ho-
listic naturalists (biological systematists and paleontolo-
gists) also occurred in Germany between 1937 and 1950
(Junker & Engels 1999; Junker 2000; Reif et al. 2000). A
crucial contribution toward the “synthetic” programs of
those years was the close cooperation between Stresemann
and the population geneticist N.W. Timofeeff-Ressovsky
(1900-1981) in Berlin during the period 1940-1945 (Haf-
fer 1994a, 1999). They discussed the genetic basis of ge-
ographical variation and genetic effects during range ex-
pansion in various bird species. Following a lecture by
Stresemann (1943) on the species concept and on the im-
plications for the speciation process of ecological differ-
ences in subspecies and species (see Stresemann 1943),
Timofeeff-Ressovsky (1943) emphasized how fruitful the
cooperation had been between the more theoretically ori-
ented genetic-evolutionary teams and the systematists and
zoogeographers during recent years, leading to full
agreement regarding the nature of the evolutionary
mechanisms. A joint study by Stresemann & Timofeeff-
Ressovsky (1947) on speciation in birds was completed
in 1944 but not published until after the end of the Sec-
ond World War. Timofeeflf-Ressovsky’s arrest by the So-
viet authorities in 1945 prevented further studies in this
series that were planned.

Examples of ornithologists in Central Europe who con-
ducted studies of birds on the basis of the New System-
atics and within the framework of the Seebohm-Hartert
school during the second half of the 20th century are W.
Meise, G. Niethammer, and G. Diesselhorst (birds of

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varoius regions of the world), J. Martens, S. Eck, A. Hel-
big, M. Kaiser, and A. Gebauer (Palearctic birds), H.
Schifter and R. van den Elzen (African birds), J. Haffer,
C. Konig, and K.-E. Schuchmann (Neotropical birds).
Many of these studies included the application of mod-
ern methods of morphological and vocal analyses as well
as molecular DNA analyses.

Several alternative notions of the general nature of
species (theoretical species concepts) conceive these units
as created entities (typological concept), as groups of non-
intergrading populations at any particular time level (bi-
ological concept), or as phyletic lineages through geolog-
ical time (phyletic and evolutionary concepts). These the-
oretical concepts need to be distinguished from the dif-
ferent (wide to narrow) taxonomic delimitation of indi-
vidual species taxa (Table 1). Under each of these theo-
retical species concepts biologists have delimited and are
delimiting “narrow” and “wide” species taxa (splitters and
lumpers respectively) depending on whether these system-
atists place the taxonomic species limit at relatively “low”
or “high” levels of microtaxonomic differentiation respec-
tively. A species limit at a fairly high level of differenti-
ation results in relatively few species taxa, each species
comprising wide arrays of variously differentiated geo-
graphical representatives. On the other hand, a taxonom-
ic species limit at a low level of differentiation results in
more numerous, internally rather uniform and narrowly
defined species taxa. Zoologists advocating different the-
oretical species concepts might in practice delimit species
taxa in a similar manner. Those, on the other hand, adher-
ing to the same theoretical species concept might delim-
it species taxa quite differently.

Problems regarding the nature of species in animals gen-
erally, and the delimitation of species taxa in particular,
have again been the subject of extensive discussion in the
recent literature (Haffer 1992, 1997c; Helbig 2000). In a
few cases, detailed work has indicated that the members
of closely related European birds can no longer be con-
sidered as representing subspecies of one biological
species (as has been the case for a long time) but are
species Just beyond the level of species differentiation (e.g.
Oenanthe hispanica/O. pleschanka and Phylloscopus
collybita/Ph. brehmii). However, it is important to real-
ize that, in these cases, both the earlier and modern tax-
onomic interpretations refer to the same species limit un-
der the biological species concept. On the other hand, the
limit of “phylogenetic species” is placed at a much low-
er level of differentiation. Even taxa intergrading along
broad zones of hybridization (subspecies under the
biospecies concept) are considered as binomially named
“species” under the phylogenetic species concept.

Zoogeography. The methods of a static 19th century zoo-
geography led to the subdivision of the world into numer-
ous faunal regions and subregions of animal distribution
based on faunal lists and their greater or lesser resem-
blance (e.g. Reichenow 1888; Schalow 1897). Beginning
during the 1910s, Erwin Stresemann viewed zoogeograph-
ical problems within ornithology from a dynamic-ecolog-
ical point of view. His method of zoogeographical analy-
sis stressed the need to examine the dispersal abilities and
distributional ranges of individual species, as well as the
ecological and geological history of particular regions to
understand the zoogeographical history and differentiation

Species concepts

Specieii limit*)

Typological

Evolutionary

Historical

morphological

biological

cladisiic paleontological

\Mde

Gloger 1833, 1834
Middendorff 185(Js

Kleinschmidt 1920s

Geyr 1924
Meinertzhagcn 1954
Eck 1985

Hcllinayr 1920s
Siresemaini 1919-1927

LnlenTiediate

Brehm 1831

Schlegel 1844
Blasius 1862

Allen 1870.S

Baird I870.s

Cooes 1870s

Sccbohm 1880s

Hartcrt 1903-1922

Sircsemann 1928 IT.

Iknnig 1966 Simpson 1961

and to 1 lowers and other

Willmann 1985 palcoiviologists

Dubois 1871

Ridgway 1870s
Dwighi 1918, 1925

Rensch 1929, 1934

Mayr 1942, 1963

Lack 1944. 1947

Bock 1979, 1986

Narrow

Temminck 1815

Vieillo! 1816

Hartlaub IH77

Sdaier 1880s

Sharpe 1899-1909

Berlepsch 1911
Reichenow 1913

Siepanyan 1974, 1978

Cracrati 1983

Zink 1996. 1997

Table 1. Theoretical species concepts (horizontal) and species limits under different taxonomic species categories (vertical) as ap-
plied by some ornithologists during the 19th and 20th centuries. Years refer to major publications; slightly altered after Haffer (1992).

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The Development of Ornithology and Species Knowledge in Central Europe

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of a genus or family of animals as a dynamic and contin-
uing process. Based on this method he analyzed the zoo-
geography of selected groups of European and Afrotrop-
ical birds in a series of papers published between 1919 and
1929 (e.g. Stresemann & Grote 1929), while F. Steinbach-
er (1927, 1929) studied how the distributional ranges of
selected European bird species resulted from historical
processes. Stresemann returned to this subject in a more
comprehensive manner in his analysis of the avifauna of
the Malay Archipelago that forms the introduction to his
monograph Die Vogel von Celebes (1939) \The birds of
Sulawesi]. In this magnificent work he emphasized the
need to take into consideration the possibility of active
range expansion of island birds across ocean barriers, and
of montane birds across lowland gaps, without the need
to postulate the former existence of land bridges or moun-
tain bridges, as had been done by many previous zoogeo-
graphers. He concluded that - apart from sea-level changes
- several very humid and dry climatic periods had alter-
nated during the Pleistocene, leading to drastic changes
in the distribution of forest and non-forest vegetation and
corresponding faunal movements and differentiation.

Stresemann’ s dynamic-zoogeographical interpretations
of faunal movements and speciation in tropical Africa
(1929, with Grote) and the Malay Archipelago (1939)
were pioneering contributions during those years. The ba-
sic premises of his theories of speciation based on climat-
ic-vegetational fluctuations have been amply confirmed
during later decades for the Palearctic Region (de Eattin
1967), Australia (Keast 1961), tropical Africa (Moreau
1966), and South America (Muller 1973; Hafifer 1974).

Comparative anatomy. The comparative study of the
anatomy of animals is one of the oldest zoological research
traditions, pursued with the aim of establishing a natural
system and of understanding the functioning and biolog-
ical roles of particular structures and organs. Blumenbach
(1779) had emphasized that the animal’s entire habitus
needed to be taken into consideration when judging its sys-
tematic relations, and Blasius Merrem (1788: 4) also as-
serted that “The similarities of all parts of the body tak-
en together, not of individual parts, must determine gen-
era, tribes, orders, and classes” (cited from Stresemann
1975: 107). These positions were again defended by Carl
Illiger (1811) and B. Merrem (1813). In this latter work
the large ratites were placed in a group separate from all
other birds because they lacked a keel on the sternum.
Friedrich Tiedemann (1810, 1814) summarized the then
available anatomical knowledge in a thoroughly re-
searched handbook published in two volumes. Christian
Eudwig Nitzsch (1782-1837), “outstanding both for his
minutely detailed research and for his acute reasoning”
was “one of the most accurate, cautious, and imaginative
morphologists who ever concerned themselves with
avian anatomy” (Stresemann 1975: 236, 308). He pub-

Bonn zoological Bulletin - Supplementum 59: 1-116

lished Osteografische Beitrdge zur Naturgeschichte der
Vogel (1811) [Osteological contributions to the natural
history of birds], a monograph on the nasal gland (1820;
see also Giebel 1 858), and observations on the carotid ar-
tery in birds (1829). In this latter study, he aligned the
swifts with the hummingbirds, moving them a long way
from the swallows. Nitzsch studied each bird that he re-
ceived from a pterylographic (study of feather tracts) and
anatomical point of view, and searched its plumage and
intestines for parasites which he also investigated from
various viewpoints. He contributed detailed chapters on
the anatomy of the genera of European birds treated in J.F.
Naumann’s great Naturgeschichte. After Nitzsch died in
1 837, Rudolph Wagner ( 1 805-1 864) in Munich continued
these contributions for the last volumes. Wagner had pub-
lished a major treatise on the anatomy of birds in 1837.
Several of Nitzsch’s manuscripts were edited and pub-
lished by his colleagues after his death (e.g. Nitzsch 1 840;
Giebel 1858, 1862, 1866). Keyserling & Blasius (1839)
announced that they had discovered a useful character of
the songbirds (except the larks Alaudidae): their “booted”
tarsometatarsus.

Another outstanding zoologist who devoted his atten-
tion, at least temporarily (during the 1 840s), to the anatom-
ical study of birds was Johannes Muller (1801-1 858), who
established the major divisions of the Passeriformes (1847)
based upon characters of the syrinx, separating the New
World (Tyrannidae) and the Old World flycatchers (Mus-
cicapidae). Eater descriptive studies of the syrinx were
those of E. Wunderlich (1884), V. Haecker (1900), and his
student W. Koditz (1925).

Max Fiirbringer (1888) and Hans Gadow (1891, 1893)
had the great merit of augmenting the knowledge of the
anatomy of birds in an unsurpassed manner when they
published in large volumes the results of their painstak-
ing studies over many years. Their classifications are the
basis of most or all later natural systems of the birds pro-
posed during the 20th century. Gadow was interested in
the comparative anatomy and taxonomic relevance of the
avian digestive system, an interest which much later led
Ziswiler (1967; Ziswiler & Famer 1972) to study the
anatomy of the digestive system and the systematic posi-
tion of granivorous songbirds. Eeiber’s (1907) detailed
work on the tongue of the woodpeckers may also be men-
tioned here.

Following the publication of Fiirbringer ’s and Gadow’s
volumes, interest in comparative anatomy with a system-
atic and phylogenetic perspective faded away and only a
few studies appeared in Gennany (e.g. Krause 1901, Mar-
tin 1904) until modern evolutionary morphology flour-
ished again through the work of D. Starck (born in 1908)
and his students (Barnikol, Fiedler, Hofer, Fang) in Frank-
furt a. M. from the mid-20th century (Duncker & Fisch-
er 1999). They studied the morphology of the head (Hofer
1955) and the jaw muscles supplied by the trigeminus

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nerve, concluding that the falcons are more closely relat-
ed to the owls than to the other groups of diurnal raptors
(Starck 1959). The Amazonian Hoatzin (Opisthocomus
hoazin) occupies a very isolated taxonomic position be-
cause of the occurrence of many primitive characters
which, during the course of evolution, have been trans-
formed in other birds through various specializations
(Barnikol 1953). The above authors also assembled evi-
dence to suggest that the Apteryges (Apterygidae and Di-
nornithidae) are widely separated from the other ratites.
Glutz von Blotzheim (1958) believed that the carinate
birds and the ratites evolved separately from different
groups of coerulosaurs.

Macrosystematics and evolution. In his Prodromus
(1811), Carl Illiger (1775-1813) rejected the old-fashioned
beak-foot classification of Linnaeus and followed the po-
sition of his intellectual precursors J.F. Blumenbach and
B. Merrem. He recognized for birds 7 orders, 41 families,
and 147 genera. During the early and mid- 19th century,
many German scientists were influenced by the natural
philosophy of F.W. J. Schelling and Lorenz Oken. In his
Natiirliches System der Vogel [Natural system of birds],
L. Reichenbach (1852) divided each category level into
four subdivisions, whereas J. J. Kaup based his Classifika-
tion der Sdugethiere und Vogel (1844) [Classification of
mammals and birds] on the “sacred number five”. For a
British example of this “quinarian system” see MacLeay
(1819-21, 1825), which was carried into ornithology by
W. Swainson (1836). After 1859, even evolutionists like
A.E. Brehm and Reichenow defended practical systems
that facilitated quick identification. Brehm (1866-1867)
used the categories of crackers (parrots, finches, and
corvids), catchers, searchers, runners, and swimmers, sim-
ilar to the classifications of 1 6th-century authors. The an-
ti-evolutionist E. von Homey er also grouped birds follow-
ing an outdated system (Stresemann 1975: 239). Other
prominent anti-Darwinians were (e.g) Schlegel, Ahum,
and Blasius.

Morphological systematics: In his classic volumes Un-
tersuchungen zur Morphologic und Systematik der Vogel
(1888) [Studies on the morphology and systematics of
birds] Max Filrbringer ( 1 846-1920) kept in mind the con-
nections between form and function as well as the allo-
metric growth of parts. Hans Gadow’s system of birds in
Bronn’s Classen und Ordnungen [Classes and orders] is
similar and the Filrbringer-Gadow system became the ba-
sis of the current classification of birds, the Wetmore-Pe-
ters system (Bock 1990a). Because of the uncertainties of
classification fixation at high taxonomic levels, Strese-
mann concluded: “In view of the continuing absence of
trustworthy information on the relationship of the high-
est categories of birds to each other, it becomes strictly a
matter of convention how to group them into orders. Sci-
ence ends where comparative morphology, comparative

Bonn zoological Bulletin - Supplementum 59; 1-116

physiology, comparative ethology have failed us after
nearly 200 years. The rest is silence” (1959: 227-228).
However the subsequent developments in molecular sci-
ence outlined below show that he was being too pes-
simistic!

His intention was to emphasize that the classic meth-
ods of comparative studies will not fill the considerable
gaps in our understanding of the relationships of many
higher-level taxonomic groups (orders) of birds. In the
meantime biochemical methods of analysis have been ap-
plied to this problem with varied success (egg-white pro-
teins, blood proteins, lipids in the uropygial gland, se-
quencing of mtDNA and nuclear DNA; Sibley & Ahlquist
1990). Bock (1992) is optimistic that progress in
macrosystematics will be achieved on the basis of func-
tional-adaptational investigations of taxonomic characters.
He illustrated this approach with the results of several
studies, (e.g.) the monophylogeny of all paleognathous
birds including the tinamous, the isolated taxonomic po-
sition of the passerine finches, the non-homology of the
foot structure in galbulids and picids, and the isolated po-
sition of the Hoatzin {Opisthocomus hoazin), often
wrongly held to be a cuckoo.

The results of several other attempts to solve macrosys-
tematic problems are here briefly summarized:

Oological systematics: Few avian orders as currently
recognized are corroborated by the characters of the egg
shell, although detailed comparisons are still lacking. On
the other hand, oological characters support the delimita-
tion of many suborders, families, subfamilies, and genera.
In many cases, oology supports the separation of certain
taxa; for instance the eggs of Opisthocomus differ conspic-
uously from those of cuckoos, and the eggs of sandgrouse
(Pteroclidae) differ from those of the pigeons and doves
(Columbidae). In many other cases, oological characters
argue against the lumping of genera or recommend their
splitting. Many other “contradictions” may be understood
as adaptations to breeding in cavities, in open sites, in
colonies, or as a consequence of a change to brood para-
sitism (Meise 1976; see also Kutter 1889, Hartert 1890,
and the long series of oological textbooks published dur-
ing the 19th and 20th centuries outlined above).

Plumage and feather change (molt) : Nitzsch (1 840) an-
alyzed the pterylography of most major groups of birds
and found similarities between (e.g.) hornbills and
hoopoes, gulls and shorebirds, pigeons and sandgrouse,
nightjars and owls, as well as swifts and hummingbirds.
He was one of the first to note pterylographic differences
between the four groups of raptors (diurnal raptors. Old
World vultures. New World vultures, owls), he separated
the bam owls (Tytonidae) from the typical owls, and found
that passerines usually have ten primaries but that the out-
ermost is always shorter than the others and is absent in
some species. Many groups of non-passeriform birds lack
the fifth secondary feather in the wing (diastataxis or

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aquintocubitalism) (Gadow 1893). Steiner (1918, 1956)
and Stephan (1970) studied this phenomenon, which the
latter interpreted functionally; the diastataxic gap devel-
oped several times independently in different groups when
the wing had to be folded more tightly as a result of evo-
lutionary changes in proportions (e.g. a relative lengthen-
ing of the humerus, of the lower arm bones, a relative
shortening of the femur, and other reasons).

From a macro systematic viewpoint, the study of molt
patterns in birds turned out to be rather disappointing. No
single mode of replacement of flight feathers is common
to two orders of birds (Stresemann & Stresemann 1966).
However certain peculiarities of the molt of primaries (P)
characterize a number of families: the falcons (Falconidae)
begin with P4 from where the molt proceeds in both di-
rections, outwardly and inwardly, whereas the hawks (Ac-
cipitridae) change their primaries in a normal descending
order. Cuckoos (Cuculidae) have a transilient mode, while
many parrots (Psittacidae) follow Hampe’s Rule, begin-
ning with P6 from where molting continues outwardly and
inwardly. Each of the three subfamilies of the kingfish-
ers (Alcedinidae) has its own particular molt sequence.
Two more recent publications treating the plumages and
molt of European birds are those of Bub (1981-1988) and
Jenni & Winkler (1994).

Modern ornithology during the 20th century

During the 19th century and until about 1920 ornitholo-
gy and general zoology were widely separated. Few or-
nithologists in Central Europe and other areas of the world
were interested in the research of anatomists, physiolo-
gists, geneticists, and psychologists. Conversely, general
zoologists worldwide viewed ornithology as the scientia
ambilis of amateurs, the results of whose work could not
mean much to researchers in the field of wider zoology.
During the 1920s, however, ornithology rapidly changed
and general biological studies were emphasized over the
earlier systematic-faunistic work. It was soon recognized
that the bird is particularly well suited for studies into the
basic problems of speciation, functional morphology,
physiology, behavior, and orientation of animals. Ornithol-
ogy became an indispensable part of modern biology. This
transformation of ornithology as an occupation for taxo-
nomic and faunistic specialists into a branch of modern
biological science during the 1920s and 1930s was large-
ly led by one man: Erwin Stresemann (1 889-1972). Dur-
ing those years he was Secretary of the German Ornithol-
ogists’ [formerly Ornithological] Society {Deutsche Or-
nithologen-Gesellschaft, DO-G formerly Deutsche Or-
nithologische Gesellschaft, DOG) (Haffer 1994a, 1997a,
2007a,b; Haffer et al. 2000). This accomplishment was of
greater general significance and had broader consequences
than his important ideas on species, speciation and zoo-

Bonn zoological Bulletin - Supplementum 59: 1-116

geography of birds. We emphasize that the “Stresemann
revolution” in ornithology was not restricted to Central Eu-
rope but had a worldwide impact. It was at that time that
ornithology became part of biology, indeed assumed a
leading role in the discipline. For this reason Ernst Mayr
(1997b: 855) stated: “I think one can say truthfully that
no one in the last 100 years has had as profound an im-
pact on world ornithology as Erwin Stresemann”. When
he had just “thrown the switch”, so to speak, and was be-
ginning to work on the new program for ornithology, Stre-
semann said: “Our task will be to work on ornithology giv-
ing up its previous isolation and achieving a respected po-
sition within the system of scientific zoology. ... A wide
field of activity lies ahead of us and thereby the prospect
of a bright future for our science” (Stresemann 1926a:
231). This revolutionary vision led to Stresemann’s far-
sighted support of exciting new research areas like func-
tional morphology, behavior, and orientation of birds, as
well as of promising young scientists like B. Rensch, E.
Mayr, K. Eorenz, G. Kramer and others (see Fig. 33).

(8.) The Stresemann Era

Erwin Stresemann as an innovator

Career. Stresemann accomplished the transformation of
ornithology into a New Biological Ornithology (or New
Avian Biology) by adding avian physiology, functional
morphology, ecology and behavior to the narrower older
ornithology when he (a) published his masterpiece, the
outstanding volume Aves (1927-1934) in the Handbuch
der Zoologie [Handbook of Zoology], (b) skillfully edit-
ed two ornithological Journals from 1922, (c) supervised
a large series of PhD dissertations, and (d) encouraged a
number of other major ornithological projects. The title
of the Festschrift for Stresemann’s 60th birthday, Or-
nithologie als Biologische Wissenschaft (Mayr & Schiiz
1949) [Ornithology as a biological science], acknowl-
edged this major achievement. It is true that the techni-
cal progress during the 20th century (e.g. the availability
of advanced photography, slow motion and color film
cameras, voice recording equipment, color bands, etc.) fa-
cilitated a broadening of research programs in many ar-
eas of ornithology, and that numerous additional oppor-
tunities for research and positions developed after about
1920, and especially after the Second World War. How-
ever, we are here mainly referring to conceptual rather than
technical advances. On the occasion of Stresemann’s 80th
birthday Ernst Mayr (1969) said:

“Despite all emphasis on the freedom of scientific re-
search, leaders are required in science as elsewhere. Or-
nithology of the 1920s badly needed such a leader. . . .
You then opened windows and doors permitting fresh air
to blow through the halls of ornithology. . . . That a new

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epoch was opened for ornithology during the 1920s, is
owed to your creativity and your efficacy. Not three oth-
ers would have been able to achieve what you did all by
yourself’.

In view of the general significance of these develop-
ments, Stresemann’s influence is here documented in some
detail. Although he himself was primarily a systematist
and zoogeographer, his interests also included functional
morphology, physiology, ecology, and ethology. Living
birds and the adaptations to their respective modes of life
always interested him deeply. Through his personal inter-
vention and with his characteristic far-sightedness he pro-
moted several new research traditions and the careers of
promising young scientists during the 1920s and 1930s as
well as the following decades. Although Stresemann was
aware of the new developments initiated through his own
work and that of his students and colleagues during that
time (see Stresemann 1938: 21-22, 1951 : 358), he did not
discuss this paradigm shift in any detail, perhaps becau-
se of personal modesty or because he was still too close
to these events when he wrote the manuscript of his book
on the history of ornithology (1951, 1975) during the la-
te 1940s.

(1) As the new editor of the Ornithologische Monats-
berichte [Monthly ornithological reports] he published
the foWov^'mg Announcement in December 1921: “On-
ly by connecting our discipline with all branches of sci-
ence research will we be able to comprehend the avian
organism and many of its biological characteristics. For
this reason we will publish in future issues of the
Monatsberichte detailed reviews of the more important
publications in the entire field of ornithology including
anatomy and physiology”.

(2) In 1922-1923 he invited B. Rensch from Halle and E.
Mayr in Berlin (then students aged 22 and 1 8 respec-
tively, though the former had already completed his
PhD) to work in his bird department at the Zoological
Museum between semesters. In 1925 he persuaded the
medical student Mayr to switch to zoology and to write
his thesis under Stresemann’s supervision.

(3) Beginning in 1924 he intensified, with O. Heinroth,
the exchange of thoughts and opinions among ornithol-
ogists in the Berlin region by organizing, besides the
public monthly meetings of the DO-G, a second more
technical monthly meeting of DO-G members only,
where many new ideas and publications were discussed
(see J. Ornithol 72 [1924]: 271, 567).

(4) During the 1920s and 1930s he supervised studies and
dissertations dealing with the physiology and function-
al morphology (“biological anatomy”) of birds, most of
them published in the Journal fur Ornithologie.

(5) A letter to K. Lorenz in 1934 (who was then 30 years
old) induced him to quit his position at the Anatomical
Insitute in Vienna in order to study animal behavior.

Bonn zoological Bulletin - Supplementum 59; 1-116

(6) In November 1934 Stresemann invited G. Nietham-
mer (then 26 years old) to come to Berlin and work on
the project of the Handbuch der deutschen Vogelkunde
[Handbook of German ornithology], published
1937-1942. In 1962 he persuaded U. Glutz von
Blotzheim to take over the editorship of the new Hand-
buch der Vogel Mitteleuropas (1966-1997) [Handbook
of the birds of Central Europe].

(7) Stresemann’s verbal criticism of E. von Holst’s ideas
on the flight of birds led to the latter’s construction of
his “artificial birds” (1943; see also J. Ornithol. 120
[1979]: 455-456).

(8) In 1946 he invited G. Kramer (then an assistant pro-
fessor with E. von Holst at the University of Heidelberg)
to coedit with him the Ornithologische Monatsberichte
(the first postwar ornithological journal) and to review
certain foreign publications that had appeared during the
Second World War. Kramer’s critical review (1948 ) of
attempts at understanding the orientation of migratory
birds led him to study this problem himself, beginning
in 1947.

Through his influence and work, Stresemann instigated a
full paradigm shift in world ornithology, the “Stresemann
revolution”. As mentioned above, ornithologists until the
early 20th century were concerned predominantly with
problems of the systematics and distribution of birds in
different regions of the world, i.e. with topics of system-
atic ornithology. Problems of general ornithology were
hardly ever addressed. The questions posed under the new
paradigm, and gradually accepted by ornithologists
worldwide, were totally different from the earlier ones and
concerned (e.g.) functional morphology, adaptations of
birds to their particular lifestyles, physiology, breeding bi-
ology, courtship, orientation, and other behaviors. In his
IOC presidential address, Ernst Mayr (1963: 28 ) stressed
“the shift of the position of ornithology from being mere-
ly a hobby to being a legitimate branch of zoology. Er-
win Stresemann has perhaps done more than anyone else
to bring about this recognition. The numerous young or-
nithologists whom he trained, as zoologists, at the Uni-
versity of Berlin, formed the first school of genuine sci-
entific ornithologists”.

We may ask in retrospect: what circumstances led to this
global paradigm change and how can we explain its oc-
currence? The coincidence of the following historical fac-
tors appears to have been decisive (Haffer 2007a,b; Haf-
fer et al. 2000):

(1) The planning of a Handbuch der Zoologie in 1913 in
which each group of animals was to be treated from a
general biological point of view. Thus the table of con-
tents of the volume on birds, as laid out schematically
by the editor W. Kiikenthal (1861-1922), was that of a

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The Development of Ornithology and Species Knowledge in Central Europe

89

handbook of general ornithology with an emphasis on
anatomy, ontogeny, physiology, and ecology, including
the relations between the bird and its environment (see
Fig. 34).

(2) The author was a young zoologist of the next gener-
ation not entrenched in the research traditions of the pre-
ceding century.

(3) Stresemann perceived his task of writing the manu-
script of the Aves volume (starting after the First World
War in 1919) and his other ornithological activities as
a challenge to develop a “New Avian Biology”, of which
\\\s Aves (1927-1934) was to become the “founding doc-
ument”.

(4) As Secretary (1922-1945) and President (1949-1967)
of the German Ornithologists’ Society, editor of the
Journal fur Ornithologie and of the Ornithologische
Monatsberichte, and supervisor of many PhD students,
as well as in his capacity of senior ornithologist at the
Berlin Zoological Museum, Stresemann occupied a cen-
tral position in German ornithology and, given his per-
sonality, was able to effectively influence the type of re-
search conducted, topics discussed at annual meetings,
and manuscripts published.

(5) Congenial colleagues like Heinroth, Koehler and
Lorenz supported Stresemann’s plans and vision. As the
most prominent professional ornithologist among these
key scientists of the 1920s and 1930s, Stresemann was
the prime mover for these new programs, encouraging
his colleagues and publishing their work and that of their
students in the Journal fur Ornithologie. Heinroth was
director of the Aquarium of the Berlin Zoological Gar-
den and did most of his ornithological work during his
spare time. Lorenz conducted behavioral studies in Al-
tenberg, near Vienna, on a private basis until he became
Professor of Comparative Psychology in Konigsberg in
1940. Kramer (1930) had made behavioral observations
in the Zoological Garden of Berlin and later worked at
the Biological Station in Naples, Italy, from where he
visited Lorenz in Altenberg during the 1930s. During
the mid- 1930s, Otto Koehler (1889-1974), Professor of
Zoology at the University of Konigsberg and a friend
of Stresemann since their student days in Munich, turned
his attention to behavioral studies of birds. With C. Kro-
nacher and K. Lorenz he founded the Zeitschrift fur
Tierpsychologie {toddiy Ethology) in 1936. The first vol-
ume appeared in 1937. These ornithologists and ethol-
ogists remained in close communication with one an-
other over the following decades, when comparative
ethology was being developed as a scientific discipline
(Festetics 1988; Koehler 1988; Wuketits 1990; Thiel-
cke 1991).

(6) Stresemann’s student Ernst Mayr (1904-2005), who
had emigrated to the United States in 1931, transmit-
ted ideas and suggestions stemming from the Strese-
mann circle to several colleagues like Margaret M. Nice,

Bonn zoological Bulletin - Supplementum 59: 1-116

HANDBUCH DER ZOOLOGIE

EINE NATURGESCHICHTE DER
STAMME DES TlERREtCHES

CECDUMDET VOW

Dit WILLY KUKENTHAL

OtKCtfiCU tCQIElUNOSUT
TEiUKD D.awor, AH Dtr UNivtAtrTAT laiiLin

UWTEn MITARSEIT
ZAHLREICHEJ4 FACHCELBWMtBW

hebausoeoebbw von

Dr,THILOKRUMBACH

Sir&ENTEB BAND < ZWEITE HALFTE

Sauropsida: Aves

Ekar^ilcT vcfl

Profr Dr. Ec Tin In Berlin

StsC Fifirfefl I* Text

BERLIN UND LEIPZIG \9Z7 BIS 1934
WALTER DE GRUyTEK ^ CO.

V4JI.UXUfH# V5JI LA-G 3 -HiI - J. V(p.LAftS-

II UC W P tu NC - RftEMtK - J. TlltiMKA - vllT i Cpwr.

Fig. 34. Title page of Stresemann (1927-1934).

J.J. Hickey and others. He organized in 1933 an ornitho-
logical seminar series where current literature was re-
viewed and especially work on bird behavior and life
history studies discussed. Hickey’s influential Guide to
Birdwatching (1943) used many of the suggestions re-
ceived from Mayr. Through his setting up of the San-
ford Hall on the biology of birds at the American Mu-
seum of Natural History (New York) in 1948, Mayr al-
so helped to further the establishment of a New Avian
Biology in North America (Figs. 35, 36).

A modernization of world ornithology in the sense of a
paradigm shift probably would have occurred even with-
out the coincidence of the above historical factors. How-
ever, such a change would presumably not have started
in Germany and not during the 1920s (but one or two
decades later), and probably would have taken place less
rapidly than it actually happened.

In 1930 Stresemann was elected President of the Eighth
International Ornithological Congress (Oxford 1934),
when he was only 40 years old. Other Central European
ornithologists before and after him who also served as
presidents of the IOC were A. Reichenow (1910), K. Im-
melmann (1986), and P. Berthold (1998), while the fol-
lowing presidents had been born and raised in Germany

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Fig. 35. Erwin Stresemann (right) and Ernst Mayr at the XIVth International Ornithological Congress in Oxford, 1966 (from Haf-
fer et al. 2000).

but lived abroad when they were elected: G. Radde (First
IOC in 1 884), E. Hartert (Sixth IOC in 1926), and E. Mayr
(Thirteenth IOC in 1962). In his presidential address in
1934, Stresemann was able to state: “Ornithology is no
longer the scientia amabilis somewhat disdainfully smiled
at by so-called competent general zoologists. We witness
nowadays that geneticists, evolutionists, and animal psy-
chologists learn from ornithologists or at any rate are treat-
ing us as being of equal rank. Eet us rejoice then over this
development, which will assure us a steadily increasing
influx of university students and thereby help us to ob-
tain a very honorable place in the system of zoology”
(1938: 21-22), and later regarding the more recent devel-
opments: “The barriers that protected our special field
were demolished on all sides. Ornithology has progressed
with such breathtaking speed that nothing important can
be achieved in it nowadays except by keeping up with the
pace, without losing sight of the whole” (1951 : 358, 1975:
351).

After Stresemann had taken over the editorship in 1921,
the Journal fur Ornithologie published more and more pa-
pers on biological topics like breeding biology, ecology,
behavior and feeding habits, thereby becoming the
world’s leading ornithological Journal during the 1930s.

Bonn zoological Bulletin - Supplementum 59; 1-116

The Journal set an entirely new style which has been
adopted to a lesser or greater degree by most major or-
nithological journals (Mayr 1963: 28). As Moreau (1959:
33) asserted after spot-checking and comparing the con-
tents of several major journals: the volumes of the Jour-
nal for “1930 and 1940 were far ahead of the Ibis in the
range and the maturity of their contributions. Apart from
many life-history papers, they dealt with breeding seasons,
migration, food, ecology and bio-chemistry”. Other top-
ics that were now frequently discussed included function-
al morphology, orientation, population studies, annual cy-
cles, and - after the sonograph had been developed - bioa-
coustical work, as well as occasional papers on zoogeog-
raphy, fossils and evolution, parasites and, during recent
years, the effect of environmental contamination with
heavy metals.

Comparing the contents of the Journal volumes prior
to and after 1920 reveals the greater length of many of the
early articles, particularly those listing the species repre-
sented in bird collections from overseas or observed dur-
ing an expedition. Also the writing style of the earlier au-
thors was generally more verbose and descriptive com-
pared with that of later contributors. Voluminous descrip-
tive reports from the bird observatories were included in

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New Avian Biology

Years

2000

1950 -

1900 -

Years

2000

- 1950

— 1900

Fig. 36. The “New Avian Biology” 1850-2000.

the Journal during the 1870s and 1880s, and again dur-
ing the first three decades of the 20th century. The num-
ber of articles in the three main research topics of the ear-
lier ornithology (systematics, faunistics in overseas coun-
tries and in Central Europe) diminished greatly during the
1930s. Those that did appear in these fields during that time
differed in approach and were based on extended field ex-
perience of the authors themselves (e.g. on the avifaunas
of Tibet- Schafer 1938, southwest Africa- Hoesch & Ni-
ethammer 1940, and Japan - Jahn 1942). They discussed
many ecological and other biological aspects of the bird
faunas of these regions. The last work of this type pub-
lished in the Journal was that on the avifauna of western
Siberia (Johansen 1943-1961).

Among the topics that occupied early, and a few mod-
ern researchers are the study of egg shells (oology) and
of plumages and molt. During the 1 850s the question was
discussed whether or not feathers may change their col-
ors or whether plumage color change is accomplished ex-

Bonn zoological Bulletin - Supplementum 59: 1-116

clusively through molting (and abrasion). In recent
decades Erwin and Vesta Stresemann studied the molt se-
quence of wing and tail feathers in many groups of birds.
Their monograph on Die Mauser der Vogel [The molt of
birds] appeared as a special issue of the Journal in 1966.

Functional morphology. Early naturalists like von Per-
nau and Zorn during the 18th century, C.E. Brehm and
many others during the first half of the 19th century, based
their observations on the teleological principle (the
search for a purpose) which, given the rudimentary state
of the sciences, led to fairly accurate conclusions for their
times (Stresemann 1975: 293, 302). Through his thought-
ful comparative observations and explanations of (e.g.) the
climbing structures of the Eurasian Nuthatch and the
woodpeckers, Brehm became a precursor of functional
morphology. Other pre-Darwinian naturalists also made
excellent observations on the structure and habits of birds
until, after 1859, Darwin’s theories diverted the attention

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of ornithologists (and zoologists in general) to historical
studies of homologies and the evolutionary relationships
of different groups of birds. However, Fiirbringer’s
(1888) and Gadow’s (1891, 1893) large volumes are of
equal importance to comparative and functional morphol-
ogy. A revival of functional studies took place in Germany
during the early 20th century through the work of Hesse
& Doflein (1910, 1914) on the structure and life of ani-
mals, Virchow (1918) on the spinal column of the Com-
mon Crane, and especially that of the anatomist and or-
nithologist Hans Boker (1886-1939), who called attention
to the direct relationships between environment, behav-
ior, and the morphology of organisms (1935, 1937). Prob-
ably at Stresemann’s suggestion, Boker held a lecture at
the annual ornithological meeting in 1924 on the evolu-
tion of bird flight that was published under the title The
biological anatomy of types of bird flight and their evo-
lution (1927).

Stimulated by the ideas of these workers and based on
his work on the manuscript of the Aves volume, Strese-
mann instigated and supervised a large series of disserta-
tions on avian morphology, embryology, histology, phys-
iology, bird flight, feather structure, and coloration with
a functional perspective, most of them published in the
Journal fur Ornithologie during the 1930s (the full titles
of these dissertations and references are listed in J. Or-
nithol. 114 [1973]: 498^99). However, no Stresemann
“school” of functional morphology developed because as
a museum scientist he was unable to place his students in
university positions, essential for the development of such
an entity. Only Hans Scharnke (1907-1941), the most ac-
tive and promising student in this field, became an assis-
tant professor at the University of Munich, but died soon
after in the Second World War. During later decades fur-
ther topics studied by the following generations of or-
nithologists concerned the functional morphology of the
avian head (Hofer, Barnikol), the oscine jaw muscles
(Fiedler), the avian jaw apparatus (Biihler), respiration (H.-
R. Duncker), ontogeny of birds (M. Starck), functional
morphology of feathers (E. Rutschke), the feathering and
bones of the wing (B. Stephan), and the flight of birds (H.
Oehme, W. Nachtigall, G. Riippell). Two workers in Cen-
tral Europe, contemporaries of Stresemann, who published
excellent functional studies during the course of the 20th
century, were Hans Steiner (1889-1969) and his students
in Zurich (e.g. V. Ziswiler, and his student in turn D.
Homberger), and Adolf Portmann (1897-1982) and his
students (especially E. Sutter) in Basle.

When H. Ruska had published the first electron-micro-
scopical images of biological objects during the 1930s,
Stresemann suggested to his doctoral student F. Frank to
apply this new technique to his study of the coloration of
bird feathers. This led to Frank’s discovery of the struc-
tures causing the blue color of feathers (Frank & Ruska
1939), although its detailed physical basis was only clar-

Bonn zoological Bulletin - Supplementum 59; 1-116

ified later (Schmidt & Ruska 1962). This first application
of the electron microscope initiated a new research tradi-
tion for the study of the iridescent feather coloration of
hummingbirds, peacocks, sunbirds, trogons and starlings
(Durrer 1962; Durrer & Villiger 1962, 1966, 1970), of
birds of paradise and ducks (Rutschke 1966, 1972), and
other birds (Dyck 1976). Otto Volker (1907-1986) ana-
lyzed the biochemical composition and physical proper-
ties of feather pigments in a series of papers, many of
which appeared in the Journal fur Ornithologie between
the 1930s and 1950s (summarized by Volker 1963, 1965).
The latter research tradition had been initiated by Ahum
(1854) and Bogdanow (1858) and was continued by the
work of Valentin Haecker and several of his students be-
tween 1890 and 1924, reviewed by Immelmann (1965).

Since the mid-1970s, a number of ornithologists have
pursued ecological aspects, investigating the adaptiveness
of morphological features, the partitioning of resources by
species, aspects of evolution, and the diversity of taxonom-
ic groups. Ecomorphological research requires full knowl-
edge of the functional and adaptive significances of the
features studied, such as bill, leg, and wing (Winkler 1989;
Bock 1990b). H. Winkler and B. Eeisler combined behav-
ioral observations with experimental and morphological
approaches, studying woodpeckers, migratory songbirds,
reed warblers, weaver finches and kingfishers (Winkler &
Bock 1976; Bock 1977; Eeisler 1977; Eeisler & Winkler
1985, 1991; Winkler & Eeisler 1985, 1992; Eeisler et al.
1987, 1989, 1997; Eey 1988; Winkler 1990).

Physiology. In 1924, comparative studies by V. Haecker,
Professor of Zoology at the University of Halle, indicat-
ed that the readiness of a bird to sing is probably related
to the development of its gonads, i.e. it depends on the pro-
duction of sexual hormones. He also discovered (1926,
1 927) differences in the histology and size of the thyroid
gland of a crow during the course of the year (small dur-
ing spring and early summer, large during fall and win-
ter, i.e. in reverse order to the annual size changes of the
gonads). An increased assimilation through an increase in
thyroid activity in birds under low ambient temperatures
during migration and molt was later confirmed (see Im-
melmann 1965). Haecker (1916, 1924) also studied the
light-dependence of the beginning and end of song in dif-
ferent bird species in the morning and evening. He already
noted that songbirds begin singing in the morning in weak-
er light conditions than when they stop singing in the
evening (when the variation is also more pronounced).

Physiological topics investigated by Stresemann’s stu-
dents included the “Physiology and acoustics of avian vo-
calizations” (Riippell, J. Ornithol., 1933), “The effects of
bacteria in the intestines of birds” (Putzig 1934), and “An-
nual changes in the histology of the thyroid gland in birds”
(Kiichler, J. Ornithol., 1935). In his major textbook,
Groebbels (1932, 1937) emphasized ornithological stud-

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ies from a physiological-biological viewpoint and attempt-
ed to uncover the functional relationships between the bird
and its environment. His work therefore effectively sup-
plemented Stresemann’s Aves. After the Second World
War, Berger studied the physiology and energetics of bird
flight (Berger & Hart 1974), Jacob (1982) the occurrence
and function of stomach oils, Jacob & Ziswiler (1982) the
structure and function of the uropygial gland,
Schwartzkopff (1973) the physiology of hearing in birds,
and Berthold (1975) the physiology of bird migration.
Prinzinger (1990, 1996) and his school analyzed energy
metabolism in birds and other animals. He established the
surprising fact that physiological time is nearly identical
for all birds regardless of mass, when expressed as “en-
ergy metabolism per gram”. Measured in (mass-specific)
physiological time units, all birds, mammals and other an-
imals have about the same duration of their different life
stages and, in this sense, have identical life spans.

Ethology and breeding biology of European birds. When
the German Ornithologists’ [Ornithological] Society DO-
G celebrated its 50th anniversary in 1900, Herman
Schalow (1852-1925) reviewed the progress made, espe-
cially with respect to the knowledge of taxonomy and dis-
tribution of European and non-European birds. He ended
his presentation with the following vista into the future:
In the years to come “the habits and life histories (of birds)
should be studied. . . . Important tasks will then be (1) to
watch these highly organized birds in their instinctive dai-
ly activities, (2) to recognize the relations between an in-
dividual bird and its social environment, and (3) to make
an attempt at perceiving and understanding the independ-
ent, peculiar actions of individual animals in particular cir-
cumstances. We will then have to examine in which way
the behavior of birds, usually considered as the result of
an inexplicable instinct, may be explained. . . as a sensi-
ble utilization of sensory perceptions” (1901: 24-25).

With this remark Schalow probably referred to the then
still unresolved controversy between the views of the Al-
tum group versus those around A.E. Brehm. An author
who resisted the usual anthropomorphic interpretation of
bird behavior during those years was Valentin Haecker
(1864-1927). In discussing the song of birds (1900) he at-
tempted a comparative treatment of song instincts with-
in a Darwinian framework, concluding that sound quali-
ty and rhythm are innate, but that the details of the song
(of Chaffinch and Nightingale) are learned from other
males, usually the father. According to Haecker, the bio-
logical functions of bird song are recognition of conspe-
cific birds as well as attraction and stimulation of females.
However, he did not mention the third important function,
that of territorial advertisement directed at other males,
which had been discovered by Altum (1868), though
Haecker probably did not know this book. In comparative
discussions, he suggested the possible evolutionary devel-

Bonn zoological Bulletin - Supplementum 59: 1-116

opment and differentiation of various song types from sim-
ple calls, of instrumental vocalizations (e.g. drumming of
woodpeckers and the drumming-flight of snipes), of song-
flights, courtship behavior, and of some general behavior
patterns of birds like ruffling of feathers, drooping of
wings, and spreading of tail feathers. He separated sim-
ple reflexes from more complex instincts; see Immelmann
(1965) for a review. The first “psychological” article pub-
lished in the Journal fur Ornithologie was a long paper
by Groebbels (1910), who also treated the development
of bird song from a Darwinian point of view. He empha-
sized the previous neglect of “omitho-psychological” top-
ics despite their general biological importance, and dis-
tinguished between (1) the anthropomorphic interpretation
(e.g. A.E. Brehm), (2) the metaphysical interpretation (e.g.
Altum), and (3) the evolutionary interpretation of behav-
ior patterns in animals (e.g. Darwin, Haecker).

The ornithologist in Germany who initiated the compar-
ative study of behavior (ethology) as a branch of zoolo-
gy was Oskar Heinroth (1871-1945). As a boy he had been
stimulated by Brehms Tierleben to observe motor patterns,
especially in the courtship behavior of ducks and geese
(like Haecker, he apparently did not know Altum’s book
of 1 868). Heinroth continued to pursue ornithological ac-
tivities while he was a student of medicine and later zo-
ology, when he learned about Darwin’s theories. After his
return from an expedition to the Bismarck Archipelago
(1900-1901) he became an assistant to the director of the
Berlin Zoological Garden in 1904 and then Director of the
Aquarium there in 1913, a post which he held until his
death in 1945 (K. Heinroth 1971; Mauersberger 1994b).

When the Fifth International Ornithological Congress
convened in Berlin in 1910, Heinroth summarized his
long-term observations on the behavior of ducks and geese
in a major presentation, a milestone in the comparative
analysis of instinctive behavior in animals within a Dar-
winian framework. He described constant innate behav-
ior patterns which, like morphological structures, charac-
terize certain taxa (species, species groups within genera,
genera, families) and indicate closer or more distant evo-
lutionary relationships.

This paper, published in a congress proceedings (1911)
with rather restricted distribution, was virtually without
any effect on other ornithologists until Oskar and Mag-
dalena Heinroth began to publish another major work: Die
Vogel Mitteleuropas in alien Lebens- und Entwick-
lungsstufen photographisch aufgenommen und in ihrem
Seelenleben bei der Aufzucht vom Ei ab beobachtet (4
vols. 1924-1933) [The birds of Central Europe pho-
tographed in every stage of life and development and ob-
served in their behavior from the egg onward]. Begin-
ning in 1 904, the Heinroths (that is, mainly Magdalena)
had hand-reared individuals of almost ah Central Euro-
pean species from the egg and observed their development
in order to determine to what extent the behavioral char-

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acters (including vocalizations) of a given species are in-
nate or have to be learned by the young bird after hatch-
ing. In these volumes they emphasized “details of habits,
growth and development, feather change, instinctive be-
havior patterns {Triebhandlungen) and psychological fac-
ulties, i.e. aspects which so far have been seldom taken
into consideration” (Heinroth & Heinroth 1924: IX). A de-
tailed presentation of Heinroth’s theoretical views and his
contributions to the development of ethology is included
in the “Russian manuscripf ’ of Lorenz (1992; see also Po-
dos 1994 and Burckhardt 1990, 1999). Stresemann
(1975: 347) concluded that Heinroth’s work was the first
step toward bridging the gap between avian systematics
and avian biology, the two major branches of ornitholo-
gy-

During the 1910s both Heinroth and Stresemann had re-
alized that the data summarized by the authors of the 19th
century represented only a beginning of an understand-
ing of the life histories of even the common birds of Eu-
rope. Much additional work was needed to complete this
task. Stimulated by the work of the Heinroths and encour-
aged by Stresemann as Secretary of the DO-G and editor
of the Journal fur Ornithologie, many ornithologists in Eu-
rope began to observe in detail the nest-life, habits and
growth of birds. The bi-weekly DO-G meetings organized
by Heinroth and Stresemann in Berlin during the 1 920s
and 1930s also helped to spread the new ideas. Numer-
ous excellent and well-illustrated contributions on the
breeding biology of various species appeared in the Jour-
nal, such as those of G. Schiermann on Savi’s Warbler (Lo-
custella luscinioides, 1928^ and Eittle Crake (Porzana par-
va, 1929), H. Siewert on raptors (1928, 1930, 1932, 1933,
1941), R. Dircksen on terns and oystercatchers (1932), F.
Goethe on Herring Gull (Larus argentatus, 1937), or O.
Steinfatt on the Spotted Flycatcher (Muscicapa striata,
1937), Woodcock (Scolopax rusticola, 1938), Fieldfare
(Turdus pilaris, 1941), and Tree Pipit (Anthus trivialis,
1941). The new periodical Beitrdge zur Fortpflanzungs-
biologie der Vogel (1924-1944) [Contributions to the re-
productive biology of birds], ably edited from 1926 on-
ward by Eudwig Schuster (1883-1954), also encouraged
this line of research, at the same time transforming sev-
eral previous egg-collectors into ornithologists (the ear-
lier Zeitschrift fur Oologie had ceased to appear in 1 924).
Early disks with the songs of about 50 bird species record-
ed by Eudwig Koch (1881-1974) were published by Hein-
roth and Koch in 1935-1936 entitled Gefiederte Meister-
sdnger Das erste tonende Lehr- iind Hilfsbuch zur
Beobachtung der heimischen Vogelwelt [Feathered mas-
tersingers; the first self-teaching audio book for the ob-
servation of our birds]. Earlier, A. Voigt (1852-1922), C.
Schmitt (1874-1958), and H. Stadler (1875-1962) had
dealt with graphical methods of the representation of bird
voices (Schmitt & Stadler 1919; Wallschlager 1982).

Jacob von Uexkiill (1921, 1934), studying the interre-

Bonn zoological Bulletin - Supplementum 59; 1-116

lationships between organisms and their environment,
demonstrated that the sense organs of animals fit their sur-
roundings and that only such objects and signals which
possess some significance for the life of the animal are
perceived. Konrad Eorenz (1903-1989) investigated the
social behavior of Jackdaws at his family estate near Vi-
enna (Austria) when, in 1924, he learned about the work
of the Heinroths. He wrote to them immediately, and their
close correspondence from 1930 to 1940 was published
by O. Koenig in 1988. In his early papers on bird behav-
ior published in Journal fur Ornithologie - Observations
on Jackdaws 1927, Contributions to the ethology of so-
cial corvids 1931, Species-specific instinctive behavior
patterns 1932, Bird flight 1933, The companion in the
bird’s world 1935 - Eorenz tried to combine the findings
of Heinroth and von Uexlcull with his own results. Eorenz
had also been corresponding with Stresemann since the
late 1920s and even asked for his advice regarding his fu-
ture career. When in 1934 the director of the Anatomical
Insitute in Vienna, where Eorenz held an assistantship, re-
tired and his successor did not permit him to continue his
previous part-time ethological studies, Eorenz (then 30
years old) turned to Stresemann “as a real friend”. His re-
ply reads as follows:

“My dear Dr. Lorenz, Berlin, 7 March 1934

In view of the new situation there can be no doubt con-
cerning your decision. You must give up anatomy. Your
talents in the fields of animal psychology are so promi-
nent that it would mean an autotomy (and in addition a
biologically detrimental one!) if you were now to be in-
timidated and act ‘rationally ’instead of instinctively. When
the instinctive side is so conspicuously developed in a per-
son as it is in your case, he should be happy about this
gift of God. Don ’t worry and plunge into the water like a
young guillemot, you will surely be able to swim. If it
somehow means an alleviation to you, you are welcome
to pass much of the responsibility on to me, I bear it with
pleasure, ... In full confidence for you. Yours (E. Str.) ”.

Thus Konrad Eorenz continued to analyze species-specif-
ic motor patterns and the social behavior of birds and oth-
er vertebrates. Many examples came from the ducks and
geese, Heinroth’s favorite groups. He summarized the re-
sults of his studies into the evolutionary relationships of
ducks based mainly on their courtship movements in the
Festschrift published on the occasion of Heinroth’s 70th
birthday (1941).

Another founder of comparative ethology was Otto
Koehler (1889-1974); see Hassenstein (1974) and Thiel-
cke (1991). After he had studied the behavior of Parame-
cium during the 1920s, he and his students observed the
breeding behavior of individually marked Ringed Plovers
{Charadrius hiaticula) and Eapwings (Vanellus vanellus)
on the Baltic coast, also conducting experiments on egg-

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rolling and orientation to the nest site (Koehler 1941;
Laven 1941). In the laboratory, the former analyzed the
ability of birds to count (1955) and the development of
song, mainly in Blackbirds (Turdus merula), with conclu-
sions on bird song as a first step towards music and lan-
guage (1951) and pre-verbal thinking in animals (1956).
Besides ethograms of numerous bird species and other ver-
tebrates, major topics of ethology in the following
decades were structure and meaning of vocal performanc-
es, song learning, imitation and dialects, courtship and ter-
ritory, enemy recognition, development and neural mech-
anism of behavior (e.g. Lohrl 1960-61, 1964; Thielcke
1968, 1970; Nicolai 1970; Curio 1976; Bergmann & Helb
1982). The phenomenon of imprinting first studied by
Lorenz was further analyzed by Immelmann (1935-1987)
and his school (Immelmann et al. 1996). He also studied
ecological aspects of the periodic reproduction of birds in
non-uniform environments, in particular the proximate and
ultimate control of reproduction, such as food supply, pre-
dation, endogenous periodicities, or environmental and so-
cial stimuli (Immelmann 1971).

Migration and orientation of birds. The phenology of
long distance migration has interested a host of ornithol-
ogists for over two hundred years (R. Wiltschko 1989).
During the late 1 9th century, efforts were made to record
and map the movements of migratory birds in Central Eu-
rope (twelve annual reports were published in the Jour-
nal fur Ornithologie 25 [1 877] to 40 [1892]). The first In-
ternational Ornithological Congress in Vienna (1884) was
to some extent established in order to set up a network of
European sites to monitor migration, but it came to noth-
ing due to the difficulties of analyzing the collected data
(Haffer 2007a).

As early as 1 859, Middendorff suggested that birds use
the magnetic field of the earth as a means of orientation.
General summaries of the phenomena involved, like mi-
gration “routes”, speed and height of flight during migra-
tion, possible means of orientation by diurnal and noctur-
nal migrants were published by E. von Homey er (1881)
and Gatke (1891), the latter after having observed bird mi-
gration on the island of Helgoland since 1 837. After bird-
banding schemes had been established at the newly found-
ed observation stations in Rossitten (today Rybachiy in
Russia) in 1901 and on Helgoland in 1910, annual reports
were published in the Journal fur Ornithologie until 1924,
during which period the study of bird migration was boost-
ed considerably (Eucanus 1922; Thienemann 1927,
1930). After the Second World War, the work of the ob-
servatory of Rossitten was transferred to Radolfzell (Bo-
densee/Eake Constance). Atlases based on the banding and
recovery reports appeared in 1931 (Schiiz & Weigold), in
1975-1985, and 1995 (Zink; Zink & Bairlein). More re-
cent summaries of bird migration studies are those of
Schiiz et al. (1971) and Berthold (1990b), which include

Bonn zoological Bulletin - Supplementum 59: 1-116

the results of radar observations introduced by Sutter
(1957).

The well-planned and successful displacement studies
of W. Riippell (1935, 1937) initiated the experimental
analysis of orientation behavior of migratory birds. Dur-
ing recent decades, laboratory experiments have estab-
lished that the direction and duration of movements in
many birds are based on endogenous programs. Markers
for orientation probably are the sun (Kramer, Schmidt-
Koenig), the stars (Sauer), and/or the magnetic field of the
earth (Merkel & Wiltschko 1965); see also Schmidt-
Koenig (1979), Berthold (1991), Wiltschko (1995),
Wiltschko & Wiltschko (1995, 1999). To compensate for
the apparent movement of the sun and the stars across the
sky, migratory birds use an endogenous “clock” and in this
way are able to maintain certain compass directions
(Kramer, Hoffmann). Internal and external Zeitgeber
(Aschoff) for circadian and circannual rhythms constitute
the mechanism of the “clock” and also regulate breeding
and molting cycles (Gwinner 1975, 1986, 1990; Berthold
1996).

The “knowledge” of the distance to be traveled in the
genetically determined direction probably results from an
endogenously determined amount of energy. When these
reserves have been consumed the bird has reached its win-
ter quarters (“vector navigation”), as already suggested on
the basis of theoretical considerations without experimen-
tal proof by several earlier authors:

“The migrating bird does not need guidance, as it mere-
ly follows an instinct which automatically prescribes the
direction. . . . The bird does not endeavor to reach a par-
ticular goal; instead, the goal of the journey results by it-
self when the migration instinct ceases” (Eucanus 1922:
132, 134; see also Stresemann 1927-1934: 692, 704).

Meanwhile these suggestions have been confirmed
through quantitative measurements of the energy used by
caged birds - of various species wintering in different ar-
eas - during the period of their “migratory restlessness”
(Zugunruhe) of differing durations, and the flight direc-
tions that they had maintained (Gwinner 1968, 1990).
Two-way selective breeding experiments with warblers in-
dicated a high influence of genetic factors for the control
of migration habits and yielded large selection responses
(Berthold et al. 1990, 1992 ). These results suggest the po-
tential for a rapid microevolution of migratory behavior
in the event of environmental changes.

Population biology of European birds. When Margaret
Morse Nice (1883-1974) visited Berlin and the Zoolog-
ical Museum in early 1932, Stresemann was fascinated by
the description of her population studies of the Song Spar-
row (Melospiza melodia) using individually marked birds
near her home in North America. He encouraged her to
write up the results obtained so far and published them in
German in the Journal fur Ornithologie (1933-1934), sev-

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eral years prior to the publication of the two volumes of
her famous monograph in the United States (1937-1942).
As IOC President, he invited her to give a paper at the
forthcoming Eighth International Ornithological Congress
in Oxford (1934) and encouraged similar studies in Ger-
many.

Gottfried Schiermann (1881-1946) had an unusual abil-
ity to locate nests: “One day, just going through the woods
with him, he discovered 54 occupied bird nests with me”
(Mayr 1997a). He determined the population density of
all species inhabiting various forest types around Berlin
by counting the nests of each species in his study sites
(Schiermann 1930, 1934, 1942^3). Ecological and pop-
ulation studies were later conducted on raptors by (e.g.)
Uttendorfer and co-workers (1939, 1952), Schnurre
(1935-38), and Wendland (1952-53, 1984).

Observations of color-banded birds were made on the
Ringed Plover (Charadrius hiaticula) on the shore of the
Baltic Sea (Koehler 1941), the Yellow Wagtail {Motacil-
la flavd) on Helgoland (Drost 1948), the Yellowhammer
(Emberiza citronella) near Munich (Diesselhortst 1949,
1950), the Whincat (Saxicola rubetrd) near Heidelberg
(Schmidt-Koenig & Hantge 1954), and the Kentish Plover
{Charadrius alexandrinus) on the island of Oldeoog (Rit-
tinghaus 1956). A long series of population studies of hole-
nesting birds was initiated by Berndt & Frieling (1939)
and continued after the Second World War by many work-
ers (e.g. Creutz 1955 and Winkel 1989). Bezzefs book on
Vogel in der Kulturlandschaft (1982) [Birds in the man-
made landscape^ includes a comprehensive discussion of
the results of modem population studies of central Euro-
pean birds, such as data on density and abundance, age
composition, mortality and recruitment, distribution pat-
terns and dynamics. In recent years, PH. Becker marked
Common Terns {Sterna hirundo) with passive transpon-
ders to obtain data on population dynamics (e.g. Wendeln
& Becker 1998).

The modem bird census programs and atlas projects per-
mit quantitative estimates of distribution and population
trends for many species of European birds.

Relations between avicnltnre and ornithological re-
search

The keeping and breeding of non-domesticated birds in
cages (aviaries) and at ponds in parks has a long tradition
in Central Europe, starting in the Middle Ages (Eauffer
1939), and was widespread when von Pemau (1660-1731)
studied local birds at his home, even experimenting with
some of them. He published his influential book in 1702
(Stresemann 1925, 1951; Thielcke 1988). Almost a cen-
tury later, J.M. Bechstein issued a Naturgeschichte der
Stubenvdgel (1794) [Natural history of cage birds] which
appeared in several German and English editions during

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the following decades. The keeping of cage birds was very
popular in many parts of Central Europe during the first
half of the 19th century, when C.E. Brehm published the
Handbuch fur den Liebhaber der Stuben-, Haus- und alter
der Zdhmung werthen Vogel (1832) [Handbook for lovers
of cage birds and all birds that can be usefully tamed].
Its attraction increased during the second half of the 19th
century because of the growing importation of cage birds
from overseas. The comprehensive publications by A.E.
Brehm on Gefangene Vogel (1870-1876) [Captive birds,
2 volumes] and K. RuB on Fremdlandische Stubenvdgel
{ 1 870-1 876) [Exotic cage birds, 4 volumes] furthered the
wide distribution of bird-keeping in Europe. These activ-
ities of bird fanciers increased scienctific knowledge on
the plumages and molt of young and adult birds of many
foreign species, as well as details of their general behav-
iors and breeding biology. The experiences gathered by
bird fanciers when keeping “simple” and “difficuh”
species in captivity were the foundation of several impor-
tant research traditions during the 20th century:

(1) In the 1920s, Hans Duncker (1881-1961) actively
sought to engage bird-keepers in scientific research to
investigate the genetics of plumage color (Duncker
1923, 1924; Birkhead et al. 2003);

(2) The ethological studies of O. & M. Heinroth, K.
Eorenz, H. Eohrl, J. Nicolai, E. Thaler, K. Immelmann
and others who themselves gathered additional avicul-
tural experiences;

(3) The experimental work on song development in song-
birds (Thielcke, Messmer, and others);

(4) The experimental studies of G. Kramer, F. Sauer, F.
Merkel, W. & R. Wiltschko, E. Gwinner, and R Berthold
on the orientation of migratory birds, on circadian and
circannual rhythms, as well as on the physiological and
genetic basis of migratory behavior in birds.

The successes of these recent research traditions were un-
thinkable without the long avicultural tradition in Central
Europe (Eohrl 1989). The DO-G therefore supported in
official statements the lawful keeping and breeding of cage
birds {J. Ornithol. 119 [1978]: 131 and 128 [1987]:
401^02). However, there exists no historical treatment
of European aviculture documenting the beginning, expan-
sion, and decline of the keeping of cage birds as a popu-
lar pastime together with remarks on the increase in ex-
perience of handling, feeding and breeding of such birds.
Regarding the history of the domesticated Canary and its
races see Stresemann (1923, 1952), also Birkhead (2003)
and Birkhead et al. (2003); on other domesticated birds
see Bezzel & Prinzinger (1990: 514-516) and Birkhead
& van Balen (2008). Eauffer (1939) mentioned some ear-
ly records of cage birds in medieval reports and described
aspects of aviculture in Thiiringen during the 20th centu-
ry. The paintings of 1 7th and 1 8th-century artists docu-

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ment various species of European birds in cages. In 1794,
72 species of cage birds from overseas were kept in Cen-
tral Europe; this number had increased to 230 in 1 870 and
to 1450 species in 1920 (Bezzel & Prinzinger 1990: 515).
Additional publications refer to the relationships between
man and birds in the Harz uplands (Knolle 1980), the
Erzgebirge (Granitz 1981), and in the Oberbergisches
Eand (Fastenrath-Brunohl 1936), but an overall treatment
is lacking. Wille & Spormann (2008) give a very compre-
hensive recent account of the Harz Chaffinch-singing com-
petitions.

Applied ornithology

Here, applied ornithology comprises both bird protection
and the prevention of damage by birds. Barthelmess
(1981) published a thorough and enjoyable historical
monograph on these topics. We will concentrate in this
section on the first of these.

In the millinery trade, the fashion of wearing feathers,
wings, or entire bird skins on ladies’ hats disappeared dur-
ing the early 20th century as a result of worldwide protests
against the destruction of whole populations of birds of
paradise, herons, egrets, grebes, etc. The organized bird
protection movement in Central Europe was initiated af-
ter 1850 by the publications of Gloger (1858a,b; 1865),
Altum (1872, 1877, 1878, 1879, 1889), and K.T. Eiebe
(see Hennicke 1 893), who furthered the general idea of
bird protection and mobilized favorable public opinion.
The IOC also played an important role in this area. The
question of bird species being “useful” or “harmful” was
at the center of the discussion. Altum based his opposi-
tion against the negative attitude toward raptors and owls
on his entomological experience and the results of his eco-
logical studies of the food of many of these birds. Eiebe
also demanded a reasonable protection of raptors based
on his view of nature as a harmonic whole where these
birds have their role too. No animal is “useful” or “harm-
ful” he said. Bau (1905) presented a clear and sensible
summary of these discussions during the late 19th centu-
ry, which in many cases had more of an emotional than a
technical basis. Eocal bird protection associations devel-
oped from 1870 onward; the German Bund fur Vogelschutz
was founded in 1899 and the Schweizerische Bund fur
Naturschutz in 1909.

Eike most of the above scholars, Hans Freiherr von
Berlepsch (1899), a cousin of Count von Berlepsch, em-
phasized the decrease of many bird species because of the
decreasing availability of suitable habitats in Central Eu-
rope (rather than because of active persecution by man)
and recommended many corresponding measures. The
best way toward a general bird protection appeared to be
via landscape protection. Important ornithological base-
line data for such an ecological approach were presented

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in the books of Jager (1874), Schnurre (1921), Groebbels
(1938), Noll (1941-1942), and Bezzel (1982), as well as
by the studies of the food of raptors by Uttendorfer and
colleagues (1939, 1952); see also Niethammer (1951,
1963).

Economic bird protection as an aid against insect
plagues originally envisioned a direct influence of insect-
eating birds and therefore recommended an artificial in-
crease in their population densities. However birds
proved ineffective once an infestation had broken out,
though they may be prophylactically effective (especial-
ly species of Paridae) through their destruction of eggs and
larvae during the winter months (Bruns 1963). The ini-
tiatives of influential personalities and private organiza-
tions led to the realization that long-term studies based on
scientific methods were necessary to resolve the multiple
connections between individual ecological factors and
functions. Quantitative data on the population dynamics
of European songbirds over recent decades indicate neg-
ative trends in population size for 20 species; only 4
species showed positive trends and 11 had quasi-stable
populations (Berthold 1990a; Berthold et al. 1999). See
especially Bauer et al. (2005).

Bird artists

Although bird artists are not researchers, their work has
always been intimately connected with that of ornitholo-
gists (Murr 1938; Mengel 1980; Nottmeyer-Einden 1994;
Schulze-Hagen 2000). For this reason the names of the
more important bird artists are mentioned here (for details
see Anker 1938, Nissen 1953, and Jackson 1999). The
Frisch brothers (and one of the sons) prepared many pleas-
ing illustrations for their father Johann Eeonard Frisch’s
Vorstellung der Vogel in Teutschland (1733-1764) [Birds
in Germany], most of which were based on observations
of living birds. This was also the case in several histori-
cally interesting collections of bird paintings of the 1 8th
century which never found a publisher: {\)Aviarium prus-
sicum by J.T. Klein (1685-1759), comprising the paint-
ings of S. Niedenthal, is preserved in Erlangen; (2) J.C.
von Mannlich’s (1741-1822) Sammlung europdischer
Vogel [Collection of European birds] originally contained
more than 300 excellent plates, only a few which could
be traced (Schlenker & Baumeister 1995); (3) the numer-
ous excellent paintings of Georg(e) Forster (1754-1794),
writer, naturalist and geographer, which he prepared as as-
sistant to his father J.R. Forster (1729-1798) during Cap-
tain James Cook’s second Pacific expedition (1772-1775)
and which are now at the British Museum and in several
archives (Steiner & Baege 1971; Whitehead 1978; Forster
2007).

A. Gabler and J.M. Hergenroder provided excellent il-
lustrations for J. Wolf & B. Meyer’s Naturgeschichte der

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Vogel Deutschlands (1805-1 821) [Natural history of the
birds of Germany]; see Schlenker (2000). Their work
reached the quality of the realistic and magnificent plates
of J.F. Naumann, who was simultaneously author and artist
of the most celebrated German ornithological handbook
published during the 19th century. J.C. Susemihl began
to paint the plates for Bor[c]khausen’s Teutsche Ornitholo-
gie in 1795, almost at the same time as Naumann started
the illustrations for his father’s Naturgeschichte. The Ger-
many of the first half of the 19th century did not have the
connections with tropical countries which Britian and
France had, which is why very few illustrations of the col-
orful birds inhabiting equatorial regions appeared there
during the first half of the 19th century (e.g. in the works
of Spix 1824-1825 andRiippell 1845).

Joseph Wolf (1820-1899) was equally able to portray
a bird in a scientifically accurate manner and in a pleas-
ing way from an artistic point of view. He was born in the
Eifel region and even as a young man contributed excel-
lent illustrations to the books of E. Riippell (Frankfurt a.
M.), J.J. Kaup (Darmstadt), and H. Schlegel (Eeiden) be-
tween 1 839 and 1 847. He continued his career in Eondon,
becoming the greatest animal painter of the 19th century
(Schulze-Hagen & Geus 2000). F.W.J. Baedecker
(1788-1865) published beautiful plates illustrating the
eggs of European birds (1855-1867), while his equally
outstanding bird paintings remained unpublished (Baege
1969). Other bird artists of the 19th century include R.
Kretschmer, F. and A. Specht, W. Heubach, W. Kuhnert,
and G. Miitzel, who also painted many representatives of
other animal groups. The poorly reproduced chromolith-
ographic plates accompanying the second edition of Nau-
mann’s work (1897-1905) were based on the high-qual-
ity paintings of B. Geisler, A. Goring, O. Kleinschmidt,
O. von Riesenthal and others. E. Aichele and W. Heubach
illustrated O. Fehringer’s Die Vogel Mitteleuropas
(1922-1931) [The birds of Central Europe]. O. Klein-
schmidt’ s bird paintings with which he illustrated his
books on the songbirds (1913) and raptors (1934) of Ger-
many, as well as those of his monograph series Berajah
(1905-1937) are well executed and excellently repro-
duced. Other Central European artists of the 20th centu-
ry include O. Natorp, H. Kirchner, F. Murr, and F. Weick
and W.D. Daunicht, the last two distinguished by their
painstaking work for the immense Handbuch of Glutz von
Blotzheim and Bauer (1966-1997).

Ornithological research centers in Central Enrope

Public and private natural history museums and collections
were the centers of ornithological research during the 19th
century, when very few salaried ornithological positions
existed. Besides professional ornithologists at the public
museums, private scholars advanced systematic ornithol-

Bonn zoological Bulletin - Supplementum 59; 1-116

ogy and influenced young scientists and their careers in
Europe. As an example we can mention Hans Count von
Berlepsch (1850-1915), who actively encouraged and sup-
ported E. Hartert, K. Jordan, and O. Kleinschmidt during
the 1890s and the young C. E. Hellmayr from 1900 on-
ward. Walter Rothschild employed Hartert at his private
museum in Tring, England in 1 892. During the early 20th
century, the museums of Vienna, Bonn, Dresden, and
Frankfurt a. M. took over the private collections of V. von
Tschusi zu Schmidhoffen, A. Koenig, O. Kleinschmidt,
and Count Berlepsch respectively, where (together with
other materials) they continue to be available for modern
research on systematics, geographical variation, and zoo-
geography.

During the decades following the Second World War the
emphasis of ornithological research gradually shifted away
from the museums to large laboratories at the universities
and other institutions, with studies of the ontogeny of be-
havior patterns, the phenomenon of imprinting, the genet-
ics of migration, or the mechanisms of orientation and nav-
igation in migratory birds. Research at the universities is
carried out at zoological institutes and depends on the sci-
entific interests of the respective professor. It is therefore
less formalized and less continuous than at museums and
similar institutions. Some universities nowadays offer in-
troductory and advanced courses in ornithology.

Most state institutions dedicated to ornithological re-
search were founded to study bird migration or applied
problems of bird protection. While these objectives con-
tinue to be pursued, most of these institutions now include
in their programs other aspects of the biology of birds.

Natural history museums and their bird collections have
important roles to play in current programs. They docu-
ment the occurrence of common, rare, threatened, and ex-
tinct species and continue to provide the basis for much
modem research, especially using molecular genetic tech-
niques (e.g. Berlioz 1960; Miller 1985; Remsen 1995;
Meams & Meams 1998: 393 ff ; Collar et al. 2003; Rhein-
wald 2003; Roselaar 2003; van den Elzen et al. 2005;
Joseph 2011; Frahnert et al. 2013). In addition, many mu-
seum scientists study live representatives of their special-
ty groups in the field and/or in aviaries. Systematic sur-
veys of particular groups of birds worldwide, or hand-
books of the avifauna of a particular region, are prepared
with constant reference to museum collections. The illus-
trations in modern field guides are painted on the basis of
museum specimens, and we all carry with us into the field
“miniature bird collections” transformed by artists and sci-
entists into identification books. The current research on
biodiversity is a Joint effort by zoologists of different
spezializations (including museum workers), demonstrat-
ing again the continued importance of museum collections.

Traditionally ornithologists, like other scientists, have
been men. This imbalance has been remedied to some ex-
tent only recently. Even in earlier times a few outstand-

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ing Central European female travelers and scientists ad-
vanced the cause of ornithology, like Ida Pfeiffer
(1797-1 855), who collected on Madagascar and many oth-
er parts of the world, Amalie Dietrich (1821-1891), who
sent birds and other animals (as well as plants) from Aus-
tralia to Germany, Dr. Emilie Snethlage (1869-1929), who
studied the bird fauna of Amazonia, and Dr. Maria
Koepcke (1924-1971), who worked on the birds of the
lowland deserts and forests as well as the Andes moun-
tains of Peru.

Bird-banding programs during spring and autumn mi-
gration periods continue with the help of numerous vol-
unteers among non-professional ornithologists. Many of
these also dedicate themselves to other avifaunistic work
at more sophisticated levels than their predecessors in for-
mer times. The results of their work represent the quan-
titative database for long-term studies and for detailed dis-
tribution atlases at local, regional, or international levels.
Many ornithologists without an institutional affiliation
conduct studies on the breeding biology, ecology, food
habits, or molt of selected species. Most members of lo-
cal and national ornithological societies and formal work-
ing groups are non-professional ornithologists. On a per-
centage basis, the number of articles published by non-
salaried ornithologists (amateurs) in the national journals
has decreased since the Second World War while, at the
same time, those of university-trained ornithologists have
increased. This is a result of specialization, improved ed-
ucation levels, augmented funding, and increased numbers
of professional ornithologists at research institutes and
conservation agencies compared with the situation prior
to the Second World War. In general, ornithology contin-
ues to combine professional scientists and spare time re-
searchers in one optimal, synergistic group working at high
scientific levels (Bairlein & Prinzinger 1999).

David Lack and the New Avian Biology in the United
Kingdom (1942-1947)

From the 1920s to the early 1940s, the journal of the
British Ornithologists’ Union, Ibis, continued to publish
mainly conventional papers on geographical ornithology
consisting of expedition reports and detailed specimen
lists, but hardly any biological articles. The New Avian
Biology seemingly went unnoticed in Britain. Several
British ornithologists, generally field ornithologists work-
ing outside the official ornithological establishment in the
United Kingdom, had published important “biological”
books and articles (mainly in the journal British Birds
founded in 1907) on the lives of British birds since about
1900. This literature, however, remained on the fringes of
British ornithology and had no influence on the editorial
policy of the Ibis. Examples are several works by Edmund
Selous (1901, 1905a,b) on the behavior of birds and on

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sexual selection; Eliot Howard (1907-14) on the behav-
ior of warblers and on territory theory; Frederick Kirkman
and his co-workers (Kirkman et al. 1910-13) on British
birds; Eagle Clarke (1912) on bird migration; Julian Hux-
ley (1914, 1922, 1925) on the courtship behavior of the
Great Crested Grebe, Red-throated Diver/Eoon, Pied Av-
ocet, and other birds; Edgar Chance (1922, 1940) on the
breeding habits of the Common Cuckoo; James Burckitt
(1924-26) on an individually marked population of the
European Robin, or Max Nicholson on How birds live
(1927), The study of birds (1929), and The art of bird-
watching (1931), and James Fisher’s bestseller of 1941
Watching Birds. The biological significance of these stud-
ies was generally recognized in Britain only many years
later (Eack 1959).

It is interesting to note that although most of these works
were reviewed favorably and in detail in Ibis, the reviews
did not lead to a change in the editorial policy of that jour-
nal or in the conservative attitude of the older generation
of British ornithologists. William E. Sclater edited /Z)A un-
der the traditional policy from 1913 until 1930, followed
by Claud B. Ticehurst until 1941. The latter also resisted
suggestions to modernize the journal, but his successor
Claude H.B. Grant nominated David Eack (1910-1973)
as “Biological Assistant to the Editor” in 1942 in order to
attract biological articles to Ibis.

As an undergraduate (in natural sciences) at Cambridge
(1929-1933), Eack had watched birds in the surroundings
of the city, discovering for instance double-brooding in the
European Nightjar. He later pursued ecological and life-
history studies of British birds during his spare time as a
schoolmaster at Dartington Hall in Devon. His publica-
tions dealt with habitat selection (his term: Eack 1933) and
with the territory theory (Eack & Eack 1933, with his fa-
ther Eambert Eack). In 1937 he drew up a “Review of bird
census work and bird population problems” in Ibis, his fa-
mous book The life of the Robin appeared in 1943. His
papers of the 1930s, which included theoretical matter and
interpretations, appeared in non-ornithological journals
presumably because they would have been rejected by the
editor of Ibis. In these papers Eack quoted from the Ger-
man-language ornithological literature and was certainly
aware of the modern biological contents of Stresemann’s
Journal fur Ornithologie. He spent four months at the
Senckenberg Museum in Frankfurt a. M. in 1929, having
learned to speak German fluently (Eack 1973; Thorpe
1974).

In 1937, Huxley recommended to Eack an ecological
study of the species differences and other aspects of the
Galapagos finches in the field, and obtained funds for
hack’s expedition to the islands from December 1938 un-
til April 1939. Eack had met Ernst Mayr during a trip to
the USA in 1935 and on his return journey from the Gala-
pagos Islands stayed with him in New York for several
weeks in 1939. Mayr tried to spread Stresemann’s

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“gospel” in the United States, where he influenced some
of the New York birders and encouraged Margaret Nice
to write up her population study of the Song Sparrow,
which he published as editor in two volumes of the Trans-
actions of the Linnaean Society of New York (Nice 1937,
1943).

Discussions of Lack’s expedition results {Darwin’s
Finches, Lack 1947) influenced Mayr’s interpretations of
bill-size differences in these finches. As he explained in
his appreciation of Lack’s work (1973: 433): “The empha-
sis in species studies during the 1930s and 1940s was
strongly on the nature of isolating mechanisms and their
origins. It was David Lack more than anyone else who re-
stored balance by emphasizing the importance of ecolog-
ical compatibility between species”. It was Lack who
brought ecology into the evolutionary synthesis, as Mayr
(1947) remarked in his article on “Ecological factors in
speciation”.

Lack suggested publishing his Galapagos findings in the
Ibis, “but received so discouraging a reply from the late
editor [Ticehurst] (before he saw the paper), and at the
same time so pressing an invitation from the CAS [Cali-
fornia Academy of Sciences], that [he] accepted the lat-
ter” (Lack 1941: 637). Ticehurst had objected to expla-
nations and interpretations if they ventured beyond the
realm of fact-gathering, mostly faunistic lists (though to
be fair to Ticehurst, the paper was also just over 150 pages
long).

In a letter to Mayr, Lack confessed that the younger or-
nithologists in Britain “are all very disturbed about the
state of the Ibis, but we are all too busy to do anything
about h”. In his reply Mayr wrote: “. . .If you can round
up sufficient members who are dissatisfied and say so, you
will not have any trouble instituting a new deal” (John-
son 2004). Lack adopted Mayr’s suggestion to start a cam-
paign for changes in British ornithology. He started to re-
view in Ibis biological papers on birds published in a wide
range of journals dealing with topics as diverse as phys-
iology, genetics, and endocrinology (20 years after Stre-
semann had started a similar program in his Ornitholo-
gische Monatsberichte). By October 1945, when Lack was
appointed Director of the Edward Grey Institute of Field
Ornithology (EGI) at Oxford University, the “revolution”
in the Ibis was well on its way, and was finally complet-
ed in 1947 when R.E. Moreau was made its editor. As
Moreau (1959: 29, 33) later stated: “A revolutionary
change occurred [in the late 1940s], as Ibis contributions
came to reflect the fact that ornithology was no longer in-
bred and isolated from the main currents of biological sci-
ence; the Ibis was increasingly filled, up to three quar-

ters, with biological papers”.

It should be emphasized that these conceptual changes
in British ornithology did not reflect a gradual and natu-
ral advance of a science into more complex biological
fields of life history, behavior, and evolution after the sys-

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tematic-faunistic basis of ornithology had been estab-
lished. Rather, the “revolution” in the Ibis (from which
modern British ornithology originated) occurred through
the infiltration of a small group of young, biologically-
minded ornithologists struggling against the conservative
attitude of the majority of older colleagues. British or-
nithology “had needed a reorientation away from collect-
ing, anatomy and classification to the study of living birds
in their natural surroundings” (Johnson 2004: 544). As in
Central Europe, this reorientation included a shift of cen-
ters of activity from the museums to zoological institutes
at universities or to independent research organizations.
Eack’s research program at the EGI emphasized life his-
tory, evolutionary ecology, and population biology of
birds, culminating in his classic of animal ecology The
Natural Regulation of Animal Numbers (1954), where he
placed ecological ideas in the context of natural selection.
The EGI became a major training center for British or-
nithologists of the next generation. Up to the early 1990s,
130 doctorates were granted to students at the Institute
(Johnson 2004: 545).

Eack’s significant contributions during the 1960s and
1970s were presented in his well-known books Popula-
tion Studies of Birds (1966), Ecological Adaptations for
Breeding in Birds (1968), Ecological Isolation in Birds
(1971), and Island Biology, illustrated by the Land Birds
of Jamaica (1976). In this last work he postulated that
“ecological impoverishment”, rather than a decrease in im-
migration rate with distance, explains the decrease in
species numbers on remote islands. He summarized his
ideas on population regulation in birds as follows. The re-
productive rate - evolved through natural selection - is
that which results in the greatest number of surviving off-
spring per pair, and popuation density is regulated by den-
sity-dependent mortality, in most species by food short-
age outside the breeding season. However research in re-
cent decades has resulted in the discovery that the limi-
tation of breeding density by territorial behavior is wide-
spread among birds. Based on the evidence available to
Eack at the time, he could not accept that territorial be-
havior limited density. He also greatly underestimated the
importance of nest sites in limiting bird breeding densi-
ties. There is also experimental confirmation available now
that the breeding densities of birds can be influenced by
all the potential limiting factors, including food supply,
nest sites, predators, parasites, and competitors (Bennett
& Owens 2002, Newton 2003). See Haffer (2008) for
more details.

These examples of the modernization of ornithology in
the United Kingdom and in Central Europe illustrate two
alternative modes or models of an ornithological paradigm
shift: starting from the “periphery” of a national group pro-
gressing inward (centripitally, in the UK and North Amer-
ica), or starting from the professional center of a nation-
al group progressing outward (centrifugally, in Central Eu-

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rope). In both cases the modernization of ornithology was
a “revolutionary” transformation occurring in a relative-
ly short period of time through the influence of mainly one
person or a small group of individuals, rather than through
a slow and gradual transition and the influence of many
people over a long period.

Ornithology became a unified and, at the same time
highly diversified biological discipline only during the
mid-20th century after having been subdivided into two
widely separated branches - systematics and natural his-
tory - for over two centuries. Modern ornithologists thus
returned to the broad holistic view of John Ray and oth-
er early masters (Haffer 2007a).

DISCUSSION

Conceptual and biographical histories of ornithology.
There are basically two different ways of looking at the
history of any science such as ornithology;

(1) A theoretical or conceptual history stresses the the-
oretical contributions, ideas, and interpretations in publi-
cations by ornithologists and follows the various research
traditions over time, their conception and their results, and
their final disappearance or amalgamation with other
branches of research. We define a research tradition loose-
ly as a research program that can be followed through time
over more than one generation of researchers; the devel-
opment of certain ideas or questions asked, or the direc-
tion taken by the investigation of important themes in time,
like “exotic ornithology”, “animal species and evolution”,
“breeding biology in general or as applied to a specific
group of birds”, “behavior of birds”, etc. Biographical de-
tails of individual ornithologists in the past are reduced
to a minimum in this approach to the science because they
are judged to be hardly relevant. Such biographical infor-
mation can, at best, serve to explain or better understand
the success or failure of ornithologists.

(2) A biographical history of ornithology on the other
hand emphasizes the lives of past ornithologists, and can,
by restricting itself to the life of a single person or small
group of people, occasionally lose sight of the significance
of wider connections within the historical development of
the field. Most of the important historical surveys of or-
nithology unite aspects of the conceptual and the biogra-
phical approaches by giving accounts of the theoretical
contributions of earlier ornithologists as well as at least
some information concerning their lives.

In this study we have stressed the theoretical achieve-
ments of the ornithologists covered but have neglected bi-
ographical detail. Obviously a most important factor for
the success of important ornithologists is their personal-
ity type, but external circumstances can also make an enor-
mous contribution to their career. Examples of this are
John Ray in the 1 7th and Erwin Stresemann in the 20th

Bonn zoological Bulletin - Supplementum 59: 1-116

century. Over many decades Ray received financial sup-
port from the family of his pupil and friend Francis
Willughby - who died young - which enabled him, since
he was unable for political reasons to practise as a cler-
gyman, to write substantial works on plants, mammals,
fish, and birds as well as natural history in general that
considerably influenced the development of these fields.
By contrast, Blasius Merrem recognized as early as the
1780s the necessity of combining systematics and field or-
nithology in a general ornithology, but fate denied him the
opportunity of realizing his plan so that in 1788 only the
introduction to his magnum opus appeared.

Facts and interpretations. Many early naturalists collect-
ed descriptive facts, e.g. on the geographical-ecological
occurrence of birds, on their song, their breeding behav-
ior, their molt, their diet, etc. Some researchers asked cer-
tain questions in order to establish a relationship between
particular observations, thus allowing a scientific/biolog-
ical interpretation or conclusion, e.g. the climbing abili-
ty and corresponding morphological adaptations in
nuthatches and woodpeckers (C.E. Brehm), or the length
of the autumnal migratory restlessness period in captive
birds relative to the distance to their winter quarters (J.A.
Naumann). In the 18th century, physico-theology provid-
ed an important theoretical framework and was a useful
stimulus for naturalists to look for examples of directed
design in the organic world, and hence to study the adap-
tations of birds in their structures, behavior (e.g. injury-
feigning in ground-nesters), or plumage colors (e.g. pro-
tective coloration). Adam von Pernau, and even more so
Pastor Zorn in the 18th century, as well as Faber, C.E.
Brehm, and Gloger in the 19th, not only described many
avian phenomena but also attempted to relate them to oth-
er features of avian life, and thus to interpret their biolog-
ical significance and draw comparisons. Other ornithol-
ogists like Bechstein or J.F. Naumann hardly made any
use of this scientific approach.

Stresemann (1926a: 227) recognized these points ear-
ly in his analyses, but over-generalized when he stated that
the ornithologists of the early and mid- 19th century “still
lacked intellectual leadership, the ability to ask the right
questions, and a path into a comparative methodology.
Ecology, sociology were unknown terms then and their ob-
servations were simply stated as facts without any attempt
to relate them to each other - with certain outstanding ex-
ceptions, such as Faber’s Uber das Leben der hochnor-
dischen Vogel [On the life of birds in the far north] (1825-
1826)”. Similar exceptions were also the works of A. von
Pernau, J.H. Zom, and J.A. Naumann.

Acknowledgments. At the start of our work was a rare copy of
the little book on the Vogelsteller [The Bird-Trapper] (1789) by
Johann Andreas Naumann, which J.H. received as a gift from
Frau Amelie Koehler (Freiburg). Tim Birkhead (Sheffield), Hans

®ZFMK

102

Jurgen Haffer (t) et al.

Englander (Koln/Cologne), Gerhard Hildebrand! (Gnetsch,
Sachsen- Anhalt), Christoph Gasser (Seis, South Tirol, Italy), Rolf
Schlenker (Radolfzell), and Karl Schulze-Hagen (Monchenglad-
bach) all helped with the literature and many other requests for
information.

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Wendland V (1952-1953) Populationsstudien an Raubvogeln.

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Wendland V (1984) The influence of prey fluctuations on the
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295

Wenzel K (1909) Bernhard Christian Otto, Pommems friihester
Omithologe. Zeitschrift filr Omithologie und praktische

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The Development of Ornithology and Species Knowledge in Central Europe

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Wiltschko R, Wiltschko W (1999) Das Orientierungssystem der
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Online Appendix 1

Species knowledge of some Central European ornithologists: 16th and 17th centuries
http://zoologicalbulletin.de/BzB_Volumes/BzM_59/Final%20Appendix%201.doc

Online Appendix 2

Species knowledge of some Central European ornithologists: 18th century
http://zoologicalbulletin.de/BzB_Volumes/BzM_59/Final%20Appendix%202.doc

Online Appendix 3

Species knowledge of some Central European ornithologists: 18th and 19th centuries
http://zoologicalbulletin.de/BzB_Volumes/BzM_59/Final%20Appendix%203.doc

Online Appendix 4

Modern German and English bird names

http://zoologicalbulletin.de/BzB_Volumes/BzM_59/Final%20Appendix%204.doc

Bonn zoological Bulletin - Supplementum 59; 1-116

®ZFMK

116

Jurgen Haffer (t) et al.

Endnotes

‘ For the spelling of the name Gessner (with double ‘s’) see Pyle (2000).

^ "Petino" is a latinized form derived from the Greek peteinds (= feathered, winged) and petomai = I fly. Petino-theology is therefore Winged-Theology or Ornitho-The-
ology.

^ Wenzel (1988: 99-100) quotes another long poem by Naumann, in which he described the very wet year of 1770.

A discussion and critical summary of Bechstein’s work on forest entomology is provided by Schwerdtfeger (1983).

^ This is a reference to the works of C.K. Sprengel, who in the context of physico-theological work in the surroundings of Berlin had discovered the pollination of many
plant species by insects, as well as their corresponding mutual adaptations.

In this book Bechstein stated that the author’s name had remained unknown to him. This was doubted by Schlencker (1982) because several ornithological authors in
the late 18th century were well acquainted with Pemau’s name (see e g. von Gochhausen above), and he presumed that Bechstein had personal reasons for his asser-
tion.

’’ When Bechstein discovered Collared Flycatcher in 1795 he called it Muscicapa collaris, a name that was preoccupied by an African species of the genus named by
an earlier author. Therefore it was a later name for Collared Flycatcher (= M. albicoUis Temminck, 1815) which became the valid one.

* Marsh Sandpiper does not occur m America, which is why Bechstein in his translation of Latham’s Synopsis (1796) could not have referred to the species, as Pfauch
(1989; 15) erroneously stated.

Though Bechstein too established some simple causal connections, such as between neck length and the up-ending feeding habits of swans (Mauersberger 1990: 37).

Taxa still currently valid are printed bold, valid complete species names are also underlined . The review of the Naumanns’ names in von Boetticher (1957) is in-
complete.

" See Dornbusch {200\, Anhang [Appendix] 3). Today there are still an adult and an immature specimen in the collection of the Naumann Museum, Kothen (G. Hilde-
brandt, pers. comm ).

The many Latin names of these forms, which he published with his “mihr, are invalid synonyms.

The term “species” has three separate meanings, which must be clearly distinguished from each other: (1 ) A particular species taxon, i.e. a particular bird species (e.g.
Eurasian Robin or Golden Oriole). (2) The taxonomic category of species, between genus and subspecies, whose boundaries can be broadly drawn to include several
geographical forms, or narrowly drawn to include only one or a few subspecies. (3) The theoretical concept of the species, i.e. what a species represents in general,
(e.g.) a reproductive community (within the framework of typological or evolutionary biological thinking), an evolutionary lineage in geological time, or a constant
type in the realm of organisms. See also Bock (1995).

“ At present we have not been able to discover any taxonomic names introduced by Gloger which are no longer valid.

“Providence” in the sense of the Christian teaching of a divine guidance of human fate.

Since 1830 Professor Steffens had been a personal friend of the future King Friedrich Wilhelm IV. It was probably at Steffens’s suggestion that Gloger wrote three
extremely lengthy letters to the Prince in which he presented his ideas in a most verbose manner and petitioned him (in vain) for financial support {Staatsarchiv).

Bonn zoological Bulletin - Supplementum 59: 1-116

®ZFMK

03 haffer BZB-emI 02.12.13 13:42 Seitell?

Appendix 1. Species knowledge of selected ornithologists in Central Europe: 16th and 17th centuries.

Sachs 1531

Marcus zum

Gessner 1555

Schwenckfeld 1603

Ray 1678

English & scientific names [largely based on

Lamm 1577-1606

Gill & Wright 2006]

-

-

-

-

Greater Loon, Diver

Great Northern Loon Gavia immer

-

-

-

-

Ash-coloured Diver

Black- throated Loon Gavia arctica

-

-

-

-

-

Red-throated Loon Gavia stellata

Schertz

Wasser Deiichrlein

Duecchel

Merch, Gr. Kobel Teucher

Loon, Crested Diver

Great Crested Grebe Podiceps cristatus

-

-

-

-

-

Red-necked Grebe Podiceps grisegena

-

-

-

-

-

Horned Grebe Podiceps auritiis

-

Deiichel, Fluder

-

-

-

Black-necked Grebe Podiceps nigricollis

Taucherlein

Klein Dunckentlin

Diicherle

Klein schwarz Teucherlin

Dobchik, Didapper

Little Grebe Tachybaptus nificollis

-

-

-

-

Puffin

Manx Shearwater Piifflnus puffiniis

-

-

-

-

-

European Storm-petrel Hydrobates pelagicus

-

Merer Schwalbe

-

-

-

Leach’s Storm-petrel Oceanodroma leucorhoa

-

-

-

-

-

Northern Fulmar Fulmanis glacialis

-

Baum Gans

Solend

-

Gannet, Soland Goose

Northern Gannet Moms bass anus

-

Scharb

Scharb

Seerabe

Cormorant

Great Cormorant Phalacrocorax carbo

Krahenscharbe

-

-

-

Shag

European Shag Phalacrocorax aristotelis

Pellican

Frembder Vogel

Onvogel

Kroffgans, Onvogel

Pelecane

Great White Pelican Pelecanus onocrotalus

Reyger

Rayer

Grawer Reiger

Grawer Reger

Common Heron

Grey Heron Ardea cinerea

-

Rott Rayer

Grosser Rorreigel

Rodter Reger

-

Purple Heron Ardea purpurea

-

-

WeyBer Reigel

Weisser Reger

White Heron

Great White Egret Ardea alba

-

-

WeyBer Reigel

-

-

Little Egret Egretta garzetta

-

-

-

-

-

Squacco Heron Ardeola ralloides

Nachtrab

-

Nachtram

Nachtrabe, Bunter Reger

Night-raven

Black-crowned Night Heron Nycticorax nycticorax

-

-

-

-

-

Little Bittern Ixobrychus minutus

-

Roter Reyer

MoBkuh

Moos Ochse, Moosreigel

Bittern

Eurasian Bittern Botaurus stellaris

Storck

Storck

Storck

Storck

Stork

White Stork Ciconia ciconia

-

Schwartz Storck

Onschwal

Schwai'zer Stork

Black Stork

Black Stork Ciconia nigra

Loffler

Loffel GanB

Loffler

Loffler, Loffelgans

Spoonbill

Eurasian Spoonbill Platalea leucorodia

-2-

Sachs 1531

Marcus zum
Lamm 1544-1606

Gessner 1555

Schwenckfeld 1603

Ray 1678

Gill & Wright 2006

-

-

Siehler

-

Ibis

Glossy Ibis Plegadis falcinellus

-

Waldtrapp

Waldtrapp

Waldrapp, Steinrabe

-

Northern Bald Ibis Geronticus eremita

Schwan

Schwan

Sell wan

Schwan

Swan

Mute Swan Cygmis olor

-

Schwan

Sell wan

-

Hooper, Elk

Whooper Swan Cygnus cygnus

-

-

-

-

-

Tundra Swan Cygnus columbianus

-

Hagelgans,

Schnehgans

Gans

-

-

Bean Goose Anser fabilis

-

-

-

-

-

Greater White-fronted Goose Anser albifrons

Wiltgantz

-

Grawgans

Grawe Gans, Wilde Gans

Wild Goose

Greylag Goose Anser anser

-

-

-

-

-

Snow Goose Anser caerulescens

-

Baumgans

BaumganB

Baum Endtele,

Bauerngans

Brentgoose

Brent Goose Branta bernicla

-

-

-

Baumgans

Bernacle

Barnaele Goose Branta leucopsis

-

“Indianische”

Gans

-

-

-

Red-breasted Goose Branta ruficollis

Rostgans

-

-

-

Ruddy Shelduck Tadorna ferruginea

-

Trapgans

FuchBgans

-

Sheldrake

Common Shelduek Tadorna tadorna

-

—

Sehmeant

Sehmalendte, Pfeiff
Endtlin

Wigeon

Eurasian Wigeon penelope

-

Schnatterente

Eeiner

Schnatter Endte, Eeiner

Gadwall

Gadwall Anas strepera

-

Tressel

Kruekendtle

Kriiek Endtlein

Teal

Eurasian Teal Anas crecca

Antrich

Wilde Ente

Retschendt

Wild Endte, Merg Endte

Mallard

Mallard Anas platyrhynchos

-

Weisse Mereh

Spitzschwanz

Spies Endte,

Sea Pheasent

Northern Pintail Anas acuta

Spitzschwanz

-

Kernel

Loffelent

Kernell

Breitschnabel

Kernel, Graw Endtlein
Loffel Endte

-3 -

Sachs 1531

Marcus zum
Lamm 1544-1606

Gessner 1555

Schwenckfeld 1603

-

Wilde Murente
Murente, Masente
Straub Ente

RothalB

Braun Endte, Mittel Endte
Rotthals, Rottkopf

-

Tressel

Y sendt

Schwarzer Teucher

Eys Endte, Nonn Endte

-

KnauBente

WeiBe Nun

Dressel

Schellent

Rhynent

Roter Teucher

Schall Ente, StauBente
Elster Endtlin, Nonn

Endte

Fisch Treiber

-

Merch

Merg

Grosser Taucher, Rache

Fischgeyer

-

Fischar

Fisch Ahr

-

Huner Whey

Adler; illustrated

MaBwy

Roetelwy

Meeradler

Mens Ahr, Schwimmer
Huner Ahr

Beinbrecher

Common Teal
Shoveler

Garganey Anas querqnedula
Northern Shoveler clypeata

Ray 1678 Gill & Wright 2006

Red-erested Poehard Netta ruflna
Poehard Common Pochard Aythya ferina

Ferruginous Duck Aythya nyroca
Tufted Duck Aythya fuligiila
Greater Scaup Aythya mania
Common Eider Somateria mollissima
Harlequin Duck Histrionicus histrionicus
Long-tailed Duck Clangula hyemalis
Scoter Common Scoter Melanitta nigra

Black Duck Velvet Scoter Melanitta fusca

Barrow’s Goldeneye Biicephala islandica
Golden Eye Common Goldeneye Bucephala clangula

White Nun Smew Mergellus albellus

Lesser Toothbilled Red-breasted Merganser Mergus serrator
Diver

Gosander Goosander (Common Merganser) Mergus merganser

White-headed Duck Oxyura leucocephala
Sea Eagle Osprey Pandion haliaetus

Honey Buzzard European Honey Buzzard Pernis apivorus

Black Kite Milvus migrans
Kite Red Kite Milvus milvus

Bald Eagle White-tailed Eagle Haliaeetus albicilla

-4-

Sachs 1531

Marcus zum
Lamm 1544-1606

Gessner 1555

Schwenckfeld 1603

Beinbrecher

Grimmer

-

-

ViBggyr

-

Hasengyr

MaBwey

Rohr Falcke

-

Weisser Weihe

Hiihnerarn

-

Wiesenweihe

Sprintzlein

Sperber

Sperber

Sperber, Sprintz

Taubenfalck

-

Habich

Habicht

MeuBgeier

-

Bushard

Bushard

Hasengeyer Hasenstosser Rotlichter Mause Ahr

Adler

Aar

Schwarzer Adler

Rotelgeyer

Wannenwaher

Wannenwaher

Rotelwey, Wannenwaher

no name;

illustrated but

identification

uncertain

Schmierlein

Schmirl

Mirle

Smyrle, Smyrlin

Paumfalk

Baumfalck

Steinfalck

Stos Falcklin

Ray 1678

Gill & Wright 2006

Bearded Vulture (Lammergeier) Gypaetus barbatus
Egyptian Vulture Neophron percnopteriis
Griffon Vulture Gyps fiibiis
Cinereous (Black) Vulture Aegypius monachus
Short-toed Eagle Circaetns gall tens
Moor Buzzard Western Marsh Harrier Circus aeruginosus

Hen (Northern) Harrier Circus cyaneus
Pallid Harrier Circus macrourus
Montagu’s Harrier Circus pygargus
Sparrow Hawk Eurasian Sparrowhawk Accipiter nisus

Gos Hawk N orthern Goshawk Accipiter gentilis

Common Buzzard Common Buzzard Buteo buteo

Rough-legged Buzzard Buteo lagopus
Lesser Spotted Eagle Aqiiila pomarina
Greater Spotted Eagle Acjuila clanga
Imperial Eagle Aquila heliaca
Golden Eagle Golden Eagle Aquila chrysaetos

Kestrel Common Kestrel Falco tinnunculus

Red-footed Falcon Falco vespertinus

Merlin

Hobby

Merlin Falco columbarius
Eurasian Hobby Falco subbuteo

-5-

Sachs 1531

Marcus zum
Lamm 1544-1606

Gessner 1555

Schwenckfeld 1603

Ray 1678

Gill & Wright 2006

S acker

S acker

Saker Falcon Falco cherrug

-

Gerfalcke

Gerfalck

-

Gerfalcon

Gyrfalcon Falco rusticoliis

PlaufuB

Falcke

Birgfalck,

Frombdling

Wanderfalck

Peregrine Falcon

Peregrine Falcon Falco peregrinus

Haselhun

-

Haselhun

Haselhun

Haselhen

Hazel Grouse Tetrastes bonasia

-

-

Murhan

-

Red Game

Willow Ptarmigan Lagopus lagopus

—

Schneehun,

Steinhun

Rock Ptarmigan Lagopus muta

Pirckhan

Birkhan

Spilhan

Spilhan, Birkhan

Black Cock

Black Grouse Lyrurus tetrix

Averhan

Grugelhan

Orhan

Orhan, Auer Han

Cock of the Wood

Western Capercaillie Tetrao urogallus

Pernijsen, Waltsch
Rephuen

Rothun

Rock Partridge Alectoris graeca

-

Rothuhn

-

-

-

Red-legged Partridge Alectoris rufa

Rephun, Feldhun

Feldt Hunle

Rabhun

Rabhun, Feldhun

Common Partridge

Grey Partridge Perdix perdix

Wachtel

Wachtel

Wachtel

Wachter, Quacke

Quail

Common Quail Coturnix coturnix

FaBhan

Phasan

Fasan

Fasan

Pheasant

Common Pheasant Phasiamis colchicus

-

Wasser Hiinlein

Aesch Hiinlin

Samet Hiinse, Moff
Hiinlin

Rail

Water Rail Rallus aquations

-

Glottis, Glutt

Heggeschar

-

-

Spotted Crake Porzana porzana

-

-

Mattkern

-

-

Little Crake Porzana parva

-

Kleines Wasser
Hunlen

Wynkernel

-

-

Baillon’s Crake Porzana pusilla

-

Rotzel

GroBwasserhtinle

Wachtelkonig

Corncrake

Corn Crake Crex crex

-

Wasser Flunlein

-

-

Water Hen, More Hen

Common Moorhen Gallinula chloropus

PleBlein,

Wasserhun

Blessentel

Boelch

Wasserhun, Thum Herr

Coot

Eurasian Coot Fulica atra

Kranich

Kranch

Kranich

Kranich

Crane

Common Crane Grus grus

- 6 -

Sachs 1531

Marcus zum

Lamm 1544-1606

Gessner 1555

Schwenckfeld 1603

Ray 1678

Gill & Wright 2006

Little Bustard Tetrax tetrax

-

-

-

-

-

Macqueen’s Bustard Chlamydotismacqueenii

Trap

Trap

Trapp

Trappe, Ackertrappe

Bustard

GroBtrappe Great Bustard Otis tarda

-

-

Wasserap

-

Sea Pie

Eurasian Oystercatcher Haematopus ostralegus

"

Koppriegerle

Sandregerlein,

Sandvogel

Sea Lark

Common Ringed Plover Charadrius hiaticula

Dulfist

-

Riegerle

Riegerlin, Tulfiss

-

Little Ringed Plover Charadrius dubius

-

-

-

-

-

Kentish Plover Charadrius alexandrinus

-

Stein Meis

Dotterelle

-

Dotterel

Eurasian Dotterel Charadrius morinellus

PiilroB

-

PulroB

Pulvier, Brachvogel

Green Plover

European Golden Plover Pluvialis apricaria

-

-

-

Grawer Kiebitz

Grey Plover

Grey Plover Pluvialis squatarola

Geubitz

-

Gyfitz

Gybitz, Gywitt

Lapwing

Northern Lapwing Vanellus vanellus

-

-

Mattknyllis

-

-

Dunlin Calidris alpina

-

-

-

-

-

Purple Sandpiper Calidris maritima

-

-

OckerftiB

-

Knot

Red Knot Calidris camitus

-

-

-

-

-

Little Stint Calidris minuta

-

-

Rotknyllis

-

-

Curlew Sandpiper Calidris ferriiginea

-

-

-

-

-

Sanderling Calidris alba

-

-

Brachhun

-

-

Broad-billed Sandpiper Limicola falcinellus

-

Teuffel

Sammethunlin

Gelfiissel

Ruff, Reeve

Ruff Philomachus pugnax

-

-

Herrschnepf

-

Jack-Snipe

Jack Snipe Lymnocryptes minimus

-

Wasserschnepff

Rietschnepff

-

Snipe

Common Snipe Gallinago gallinago

-

Wasserschnepff

-

-

-

Great Snipe Gallinago media

Schnepfe

Waldtschnepff

Schnepff

Wald Schneppe,
Schnepffe

Woodcock

Eurasian Woodcock Scolopax rusticola

-

Goduvitta, Deffyt

-

Godwit

Black- tailed Godwit Limosa limosa

-

Polschnep

-

-

Bar-tailed Godwit Limosa lapponica

- 7 -

Sachs 1531

- 8 -

Sachs 1531

Marcus zum

Lamm 1544-1606

Gessner 1555

-

Regenvogel

-

Brachvogel

Rottbein

Steingallyl

Wasser Schnepff

Rotbein

Wassergall

Glutt

LeiBklicker

Fysterlin

-

Koppriegerlin

-

Samethiile

Avosseta

Uberschnabel

-

Schnirring

Marcus zum

Gessner 1555

Schwenckfeld 1603

Gr. Brachvogel,
GeiBvogel

Rottbein

Griinfussel

Schwenckfeld 1603

Ray 1678

Gill & Wright 2006

Whrimbell

Curlew

Whimbrel Numenius phaeopus

Eurasian Curlew Numenius arquata

Redshank

Spotted Redshank Tringa erythropus

Rotschenkel Common Redshank Tringa totanus
Marsh Sandpiper Tringa stagnatilis

Common Greenshank Tringa nebularia

Green Sandpiper Tringa ochrupos

Wood Sandpiper Tringa glareola

Common Sandpiper Actitis hypoleucos

Turn- Stone

Ruddy Turnstone Arenaria interpres

Red-necked Phalarope Phalaropus lobatus

Grey Phalarope Phalaropus fulicarius

Black-winged Stilt Himantopus himantopus

Avocet

Stone Curlew

Pied Avocet Recurvirostra avosetta

Eurasian Stone-curlew Burhinus oedicnemus

Gannet, Skua

Collared Pratincole Glareola pratincola

Black-bellied Sandgrouse Pterocles orientalis

Great Skua Stercorarius skua

Arctic Skua (Parasitic Jaeger) Stercorarius parasiticus
Black-legged Kittiwake Rissa tridactyla

Eittle Gull Larus minutus

Mediterranean Gull Larus melanocephalus

Ray 1678

Gill & Wright 2006

Lamm 1544-1606

"

Grosse

Rheinschwalb

Meb

Black-headed Gull Chroicocephalus ridibindus

-

-

-

-

Common Sea Mai

Common (Mew) Gull Larus canus

-

-

Meermeb

-

Herring Gull

European Herring Gull Larus argentatus

-

-

-

-

-

Eismdwe Glaucous Gull Larus hyperboreus

-

-

-

-

Blaek and White Gull

Lesser Black-backed Gull Larus fuscus

-

-

-

-

-

Great Black-backed Gull Larus marinus

-

Fischerlin

Fiseherlin

Fiseherlin

Lesser Sea Swallow

Little Tern Sternula albifrons

-

-

-

-

Greater Sea Swallow

Arctic Tern Sterna paradisaea

Meerschwalb

Spyrer, Schnirring

Sehnirring

-

Sea Swallow

Common Tern Sterna hirundo

-

-

-

-

-

Sandwich Tern Sterna sandvicensis

-

-

-

-

-

Caspian Tern Hydroprogne caspia

-

-

-

-

-

Gull-billed Tern Gelochelidon nilotica

-

Grosse Rhein
Schwalb

-

-

-

White- winged Tern Chlidonias lecopterus

-

Meyvogelein

Meyvogelin

Kl. Sehwarze
Seesehwalbe

Seare Crow

Black Tern Chlidonias niger

-

-

-

-

Sea Turtle

Black Guillemot Cepphus grylle

-

no name, but
illustrated

-

-

-

Common Guillemot (Murre) Uria aalge

-

-

-

-

Auk, Razor-Bill

Razorbill Alca tor da

-

-

-

-

-

Little Auk Alle alle

-

Islendisehe Ent

Meerkray

-

Razor-Bill

Atlantic Puffin Fratercida arctica

Holtaub

Holztaub, Loehtaub

Loehtaub

Hoi Taube

Stock Dove

Stock Dove Columba oenas

Tauber

Wilde Daub

Ringeltaub

Ringel Taube

Ring-Dove

Common Wood Pigeon Columba palumbus

-

-

-

-

-

Collared Dove Streptopelia decaocto

Turteltaub

-

Turteltaub

Turtel Taube

Turtle Dove

European Turtle Dove Streptopelia turtur

Gutzgauch

Gutzgaueh

Gueker, Guggaueh

Gueh Huch, Kuekuk

Cuckoo

Common Cuckoo Cuculus canoriis

Schleiereule

-

Sehleyeriil

Sehleiereule

Barn Owl

Barn Owl Tyto alba

-9-

Sachs 1531

Marcus zum

Lamm 1544-1606

Gessner 1555

Schwenckfeld 1603

Ray 1678

Gill & Wright 2006

Schaffickel

_

Klein Keuzlin

Eittle Horn Owl

Eurasian Scops Owl Otus scops

-

Huw

Berghu, Puhuy

Great Eagle Owl

Eurasian Eagle Owl Bubo bubo

-

-

-

-

-

Snowy Owl Bubo scandiacus

-

-

-

Ohreule, Ohrkauz

Horn Owl

Long-eared Owl Asio otus

-

-

-

-

-

Short-eared Owl Asio flammeus

-

-

-

-

-

Northern Hawk Owl Surnia ulula

-

-

-

-

-

Eurasian Pygmy Owl Glaucidium passerinum

Keutzlein,

Stainewl

—

Steinkutz, Kutz

Steineule, Nachteule

—

Little Owl Athene noctua

-

-

-

-

-

Tengmalm’s O^XAegolius funereus

Stockeul

-

Stoekeul

Graw Eule, Pusch Eule

Brown Owl

Tawny Owl Strix aluco

-

-

-

-

-

Ural Owl Strix iiralensis

-

Nachtschadt

GeiBmaleker

Nachtschade,

Tern Owl, Goat-

European Nightjar Caprimulgus europaeus

Tagschlafer

Sucker

-

Spyr

Spyr

Mauer-, Reinschwalbe

Swift

Common Swift Apus apus

-

-

-

-

-

Alpine Swift melba

Eisvogel

Eisvogel

Eisvogel

Eysvogel

King Fisher

Common Kingfisher Alcedo atthis

-

-

Imbenwolff

-

-

European Bee-eater Merops apiaster

Welscher Haher,
Roller

Roller

Mandelkrahe

European Roller Coracias garrulus

Widhopff

-

Wydhopff

Wiede-Hopffe, Kot-Han

Hoopoe

Eurasian Hoopoe Upupa epops

Windhals

Windthals

WindthalB

Windhals, Wendehals

Wryneck

Eurasian Wiyneck Jynx torquilla

-

Kleiner Grtinspeeht

-

-

-

Grey-headed Woodpecker Picus canus

Griinspecht

Grunspecht

Grunspecht

Griin Specht

Green Woodpecker

Green Woodpecker Picus viridis

Holkro

Krayeii Specht

Kraespecht

Hoi Krahe

Great Black
Woodpecker

Black Woodpecker Dry oc opus martius

Paumheckel

“ein kleiner

Bunter Specht

Bundter Specht

Greater Spotted

Great Spotted Woodpecker Dendrocopos major

gesprockelter

Specht”

Woodpecker

Middle Spotted Woodpecker Dendrocopos medius

-

-

-

-

-

White-backed Woodpecker Dendrocopos leucotos

-

-

-

-

Lesser Spotted

Lesser Spotted Woodpecker Dendrocopos minor

.

.

.

.

Woodpecker

Eurasian Three-toed Woodpecker Picoides tridactylus

- 10-

Sachs 1531

Marcus zum

Gessner 1555

Schwenckfeld 1603

Ray 1678

Gill & Wright 2006

Heubel-Lerch

Lamm 1544-1606

Kobellerche,

Heubellerch

Heubel Lerche

Crested Lark

Crested Fark Galerida cristata

Heydel-Lerch

Haubenlerche

Waldt Lerche

Waldlerch

Heide Lerche

Wood lark

Woodlark LuUida arborea

Lorch, Stainlerch

Lerch

Sanglerch

Sanglerche, Feld Lerche

Field Fark

Eurasian Skylark Alaiida arvensis

-

Ohrenlerch

-

-

-

Horned Fark Eremophila alpestris

Reinschwalb

-

Rhynschwalm

Uferschwalbe

Sand Martin

Sand Martin Riparia riparia

-

-

-

-

-

Eurasian Crag Martin Ptyonoprogne rupestris

Hausschwalb

Schwalb

HauBschwalm

Haus Schwalbe

House Swallow

Barn Swallow Hirundo rustic a

Plicksterz

Gemeine Schwalbe

Murspyr

Leimschwalbe

Martin

Common House Martin Delichon urbicum

-

-

Guckerlin

-

-

Tawny PipxtAnthus campestris

Greinerlein

no name, but

Grienvogelin

Spies Lorche

Tit Fark

Tree Pipit Anthus trivialis

Krautvogel

illustrated;
“Waldt Lerche”

ditto

Meadow Pipit Anthus pratensis

-

-

-

-

-

Water Vv^ii Anthus spinoletta

Kiischeisen

Gelbe Bachsteltz,

Ryserle

Kuh-Bachstelze

Yellow Wagtail

Yellow Wagtail Motacilla /lava

Wasser Steltz
Gemeine blaue

Galb Wassersteltz

Grey Wagtail Motacilla cinerea

Bachsteltz

Pachstelz

Weibe, Graue
Bachsteltz

Bemlein

Seidenschwantz,

Bohmerl

-

Wasser Amsel

Kiiniglein

-

Praunellen

-

-

Vogel von Kyburg

Rotkropff

Rotbrustlin

- 11 -

Sachs 1531 Marcus zum

Lamm 1544-1606

Nachtigal Nachtegal

Rotschwentzlin

Prandvogel

Rotschertz Haus Rottele

Steinschmatz

Steinrotlin Steintroschel

Pirgamschel Ring Amsel

Amschel Stein Amsel

Krametvogel Krametsvogel

Troschel Troschl

Ziemer
Sangdroschel

Wysse Wassersteltz Bachstelze

Behemle

Seideschwanz

Wasseramsel

Wasser Amsel

Zaunschlipflin

Zaunkonig

Prunelle

-

Waldtrdtele

Rott Kalichen

Gessner 1555

Schwenckfeld 1603

.

Grosse Nachtigall

Nachtgall

Kl. Nachtigall, Dorling

Wegflecklin

-

HuBrotele

Haus Rotele, Wiistling

Rotsterz

Rotschwenzel

-

Steinfletsche

Biirstner

Steinrotele

Stein Rotele

Blawvogel

-

Ringamsel

Ringdrossel

Amsel

Amsel

Reckoltervogel

Ki'ammetsvogel

Trostel

Sang Drossel

Winsel

Klein Ziemer

Mistier

Mistier, Schnarre

Mistier

White Wagtail

White/Pied Wagtail Motacilla alba

Bohemian Waxwing Bombycilla garrulus

Water-Ouzell White- throated Dipper Cinclus cinclus

Wren Winter Wren Troglodytes troglodytes

Hedge Sparrow Dunnock Prunella modular is

Alpine Accentor Prunella collaris
Robin European Robin Erithacus riibecula

Ray 1678 Gill & Wright 2006

Thrush Nightingale Luscinia luscinia
Nightingale Common Nightingale Luscinia megarhynchos

Bluethroat Luscinia svecica
Black Redstart Phoenicurus ochruros
Redstart Common Redstart Phonicurus phoenicurus

Whin-chat Whinchat Saxicola rubetra

Stone-chaffer Common Stonechat Saxicola rubicola

Stone-chatter Northern Wheatear Oenanthe oenanthe

Greater Redstart Rufous-tailed Rock Thrush Monticola saxatilis

Solitary Sparrow Blue Rock Thrush Monticola solitarius
Ring-Ouzel Ring Ouzel Turdus torqiiatus

Blackbird Common Blackbird Turdus merula

Dusky Thrush Turdus eunomus
Fieldfare Fieldfare Turdus pilaris

Song Thrush Song Thrush Turdus philomelos

Redwing Redwing Turdus iliacus

Misselbird Mistle Thrush Turdus viscivorus

-

GraBmitch

-

-

- 12-

Sachs 1531

Marcus zum

Lamm 1544-1606

Gessner 1555

Schwenckfeld 1603

WeiBgrasmiicke

Guckerle

Biirstner

Grasmuck

Grasemiicke

Munchlein

Graue Grasmuck
Munchlein

Schwartzkopf

Munchlein, Grase Spatz

Flinderling

Weidengiickerle

Wydenguckerlin

Gelber Stichling

Goldhan

Golthanle, Konigle

Goldhendlin

Gold Hanlin

-

-

Thannmeislin

-

-

Baumfleckle

Todtenvdgele

-

Lesser erested Lark Grasshopper Warbler Locustella naevia

River Warbler Locustella fluviatilis
Savi’s Warbler Locustella luscinioides
Aquatic Acrocephalus paludicola

Ray 1678 Gill & Wright 2006

Sedge Warbler Acrocephalus schoenobaenus
Marsh Warbler Acrocephalus palustris
Eurasian Reed Warbler Acrocephalus scirpaceus
Great Reed Acrocephalus arundinaceus

Icterine Warbler Hippolais icterina
Barred Warbler Sylvia nisoria
Lesser Whitethroat Sylvia curruca
White Throat Common Whitethroat Sylvia communis

Garden Warbler Sylvia horin
Black-Cap Blackcap Sylvia atricapilla

Greenish Warbler Phylloscopus trochiloides
Western Bonelli’s Warbler Phylloscopus honelli
Wood Warbler Phylloscopus sibilatrix
Common Chiffchaff Phylloscopus collybita
Willow Warbler Phylloscopus trochilus
Golden-crowned Goldcrest Regulus regulus

Wren

Firecrest Regulus ignicapilla
Spotted Flycatcher Miiscicapa striata
Red-breasted Flycatcher Ficedula parva
Collared Flycatcher Ficedula albicollis
Pied Flycatcher Ficedula hypoleuca

Bearded Reedling Panurus biarmicus

no name, but
illustrated

ZaglmaiB

Sehwantz MeiBlein

Schwantzmeislin

Schwanzmeise

Longtailed Titmouse

Long-tailed Tit Aegithalos caudatus

- 13 -

Sachs 1531

Marcus zum

Lamm 1544-1606

Gessner 1557

Schwenckfeld 1603

Ray 1678

Gill & Wright 2006

KotmaiB

Koel, Mur, or Kat
Meis; Schwartz
Meise

MurmeiB

Mur Meise

Marsh Titmouse

Marsh Tit Poecile palustris

-

ditto

-

-

-

Willow Tit Poecile montana

HeubelmaiB

Kobelmaise

KobelmeiB

Heubel Meise, Kobel
Meise

Crested Titmouse

Crested Tit Lophophanes cristatus

ThonmaiB

Waldmeisle,

Tannenmeisle

Kleine Kol Meise

Black-headed

Coal Tit Periparus ater

Tannenmeisle

Titmouse

PynmaiB

Mehl Meisle

BlawmeiB

Blaw Meislin

Blue Titmouce

Blue Tit Cyanistes caeruleus

-

-

-

-

-

Azure Tit Cyanistes cyanus

KolmaiB

Ross Meis, Spiegel
Meis

SpiegelmeiB

Great Titmouse

Great Tit Pants major

Klayber

NuBpicker

Klaber, Blauspechtle

Blaw Specht

Nuthatch

Eurasian Nuthatch Sitta europaea

-

-

Murspecht

Mauer Specht

Wall-Creeper

Wallcreeper Tichodroma muraria

Baumheekel

Baumkletterle

Baumkletterlin

Baumkletterlin

Creeper

Eurasian Treecreeper Certhia familiaris

-

-

-

-

-

Short-toed Treecreeper Certhia brachydactyla

-

-

-

-

-

Eurasian Penduline Tit Remiz pendulinus

Tyrolt

Widtwalch, Bierolt

Witte wal

Witte wale, Bierholt

-

Eurasian Golden Oriole Oriohis oriolus

Dorndreer

Thorn Kretzer

Warkengel

Wankrengel

Lesser Butcher Bird

Red-backed Shrike Lanius collurio

-

-

-

Kl. Bunter Wankrengel

-

Eesser Grey Shrike Lanins minor

Wolgerhod

Warkengel

Nuenmoerder

-

Greater Butcher Bird

Great Grey Shrike Lanius excubitor

-

-

-

Roter Wankrengel

-

Woodchat Shrike Lanius senator

Heher

Gutheher, Hetzler

Haeher

NuB Har, Marcolfus

Jay

Eurasian Jay Garrulus glandarius

Agerlaster

Atzel

Aglaster

Aglaster, Elster

Magpie

Eurasian Magpie Pica pica

NuBheher

Nussbrecher

NuBbrecher

NuBbicker, NuBkrahe

-

Spotted Nuteracker Nucifraga caryocatactes

Pirgkra

-

Alprapp

-

-

Alpine Chough Pyrrhocorax gracuhis

-

-

Tulla

-

Cornish Chough

Red-billed Chough Pyrrhocorax pyrrhocorax

Tahe

Tul

Talke

Thale, tule

Jaek-Daw

Western Jackdaw Corviis monedula

Ruchen

-

Roerk

Rouch, Roocke

Rook

Rook Cojx^us fi^ugilegus

-

-

Kray

-

Carrion Crow

Carrion Crow Corvus corone

-

Nebelkrehe

-

-

-

Hooded Crow Corvus cornix

Rabe

Rhab

Rapp

Rappe, Kol Rabe

Raven

Northern Raven Corvus corax

- 14-

Sachs 1531

Marcus zum

Lamm 1544-1606

Gessner 1555

Schwenckfeld 1603

Ray 1678

Gill & Wright 2006

Veldstar

Star

Staar

Staar

Starling

Common Starling Sturnus vulgaris

-

-

-

-

-

Rose-coloured Starling Pastor roseus

Sperck

Spatz

Spatz

Spatz, Sperrling

House Sparrow

House Sparrow Passer domesticus

Pirgspatzen

Rohrspatz

Ringelspatz

Wold-, Weidensperling

Mountain Sparrow

Tree SpaiTow Passer montanus

-

Steintlnck

-

-

-

Rock Sparrow Petronia petronia

-

Schneefinck

-

-

-

White-winged Snowfinch Montifringilla nivalis

Einck, Kotfinck

Buchfmck

Einck

Buch Eincke

Chaffinch

Common Chaffinch Fringilla coelebs

Bemlein, Gegler

Danfinck, Thanfmck

Waldfinck

Queck, Goldfmcke

Brambling

Brambling Fringilla montifringilla

Hirngrill

Zerin

Hirngrylle

-

-

European Serin Serinus serinus

Wonitz

Grtinling

Grtinling

Grtiner Henffling

Greenfinch

European Greenfinch Chi oris chloris

Zitrinlein

ZeiBlein

Zitrinle

-

-

Citril Finch Carduelis citrinella

Stiglitz

Disteltlnck

Distelfinck

Stieglitz

Gouldfinch

European Goldfinch Carduelis carduelis

ZeiBlein

-

Zeisel, ZinBle

Zeisich

Siskin

Eurasian Siskin Carduelis spinus

Henfling

Henffling

Eynfinck

Henffling

Einnet

Common Einnet Carduelis cannabina

-

-

-

Stein Henffling

-

Twite Carduelis flavirostris

Zotscherlin

Plaetlin

Schosserle

Zotscherlin

Redheaded Einnet

Common Redpoll Carduelis flammea

Krumbschnabel

Kreutzvogel

Krinitz

KL Gemeiner Krinitz

Crossbill

Common Crossbill Loxia curvirostra

-

-

-

Gr. Krinitz

-

Parrot Crossbill Loxia pytyopsittacus

-

Gyntel

-

-

-

Common Rosefinch Carpodacus erythrinus

-

-

-

-

-

Pine Grosbeak Pinicola enucleator

Giimpel, Quetsch

Blutfinck

Blutfinck

Blutfinck, Giimpel

Bull-Finch

Eurasian Bullfinch Pyrrhula pyrrhiila

Kernbeisser

Kirschfinck

Steinbeisser

Steinbeisser

Grosbeak, Hawfinch

Hawfinch Coccothraustes coccothraustes

- 15-

Sachs 1531

Marcus zum

Lamm 1544-1606

Gessner 1555

Schwenckfeld 1603

Ray 1678

Gill & Wright 2006

-

-

-

-

-

Eapland Bunting (Eongspur) Calcarius lapponicus

Wysse Emmeritz

Winterling,

Schneevogel

Bunting

Snow Bunting Plectrophenax nivalis

Emerling

Goldtheimer

Emmeritz

Embritz

Yellowhammer

Yellowhammer Emberiza citrinella

-

-

Gaulammer,

Goldammer

-

-

Cirl Bunting Emberiza cirlus

-

-

Wisemmertz

-

-

Roek Bunting Emberiza cia

-

Gilbling

Gilbling

-

-

Ortolan Bunting Emberiza hortulana

-

-

Rohrsperling

-

-

Reed Bunting Emberiza schoeniclus

WeiBemerling

Emmeritz

Gersthammer

Gersthammer

-

Corn Bunting Emberiza calandra

Appendix 2. Species knowledge of selected ornithologists in Central Europe: 18th century

- 1 -

Gochhausen 1710,
1732

Taucher, Wasser-Huhn

Fischreiher

Kleiner Reiher
Rohr-Dommel

Frisch 1733-1743

Zorn 1742, 1743

Klein 1760

Gill & Wright 2006

Embergans, Zweite Halbente

Great Northern Eoon Gavia immer

Sehwarzkehliger Seetaueher

-

Duntergans, Vierte Flalbente

Black-throated Loon Gavia arctica

-

-

Erste Flalbente, Dritte Flalbente

Red-throated Loon Gavia stellata

Gehornter Seehahn

Taueher

Bekappter u. gehornter Taucher

Great Crested Grebe Podiceps cristatus

-

-

-

Red-necked Grebe Podiceps grisegena

-

-

-

Horned Grebe Podiceps aiiritus

-

-

-

Black-necked Grebe Podiceps nigricollis

Kleiner Seehahn

-

Schwarzes Teucherlein

Little Grebe Tachybaptiis ruficollis

-

-

-

Manx Shearwater Puffinus puffinus

-

-

-

European Storm-petrel Hydrobates pelagicus

-

-

-

Leach’s Storm-petrel Oceanodroma leucorhoa

-

-

Mallemugge

Northern Fulmar Fulmarus glacialis

-

-

Bassaner, Schottische Gans

Northern Gannet Moms bassamis

Seharb

-

Wasserrabe

Great Cormorant Phalacrocorax carbo

-

-

Seekrahe

European Shag Phalacrocorax aristotelis

-

-

Kropfgans

Great White Pelican Pelecamis onocrotalus

Gemeiner Reiger

Fiseh-reiger

Graiier Reyger

Grey Heron Ardea cinerea

-

-

-

Purple Heron Ardea purpurea

-

-

WeiBer Reyger

Great White Egret Ardea alba

WeiBer Reiger

-

-

Little Egret Egretta garzetta

-

-

-

Squacco Heron Ardeola ralloides

Asehgrauer Reiger

-

Bunter Reyger

Black-crowned Night Heron Nycticorax nycticorax

Kleine Rohrdomme

-

Kleine Rohrdommel

Little Bittern Ixobrychus minutus

Rohrdommel

-

-

Eurasian Bittern Botaurus stellaris

Storch

Schwarzstorch

Storch

Schwarzer Storch
Loffelreiger

Storch

-2-

Gdchhausen 1710,

Frisch 1733-1763

Zorn 1742, 1743

1732

Schwane

Schwaane

Schwan

Wilde Gans

Wilde Gans

Wilde Gans

-

WeiBwangige Gans

-

-

Baumgans

-

-

Moppelgans

-

_

Krachtente

_

Pfeiffente, Speckente

Blassente

-

Kriekente

_

Wilde Ente

Wilde Ente

Gemeine wilde Ente

-

Fasanente

-

-

Knackente

-

-

Eoffelente

-

WeiBer Storch White Stork Ciconia ciconia

Schwarzer Storch Black Stork Ciconia nigra

WeiBer Loftier Eurasian Spoonbill Platalea leucorodia

Waldhoff Northern Bald Ibis Geronticus eremita

Glossy Ibis Plegadis falcinellus

Klein 1760

Gill & Wright 2006

Zahmer Schwan

Mute Swan Cygnus olor

Wilder Schwan

Whooper Swan Cygnus cygnus

-

Tundra Swan Cygnus columbianus

Wilde Gans

Bean Goose Anser fabilis

Eachgans

Greater White-fronted Goose Anser albifrons

Graugans

Greylag Goose Anser anser

-

Snow Goose Anser caerulescens

-

Barnacle Goose Branta leucopsis

Brentgans

Brent Goose Branta bernicla

-

Red-breasted Goose Branta rufficollis

-

Ruddy Shelduck Tadorna ferruginea

Fuchsgans

Common Shelduck Tadorna tadorna

Pfeifente

Eurasian Wigeon Anas penelope

Mittelente, Schnarrente

Gadwall Anas strepera

Kriekente

Eurasian Teal Anas crecca

Gemeine wilde Ente

Mallard Anas platyrhynchos

SpieBente

Northern Pintail Anas acuta

Franzente

Garganey Anas querquedula

-

Northern Shoveler ^^<25 clypeata

Tafelente

StrauBente

-3 -

Gochhausen 1710, Frisch 1733-1763 Zorn 1742, 1743

1732

Nordische schwarze Ente

Schwarzkopf, Ententaucher

Kreutzente Scheckente, Mergente

Schmielente Mittelente, MoBkolben

See-Rachen Meerrach Schnarrgans

Fisch-Geyer - Fich-Ar, Fisch-Geyer

Wye

Schwalben-Schwantz - Milan, Gabler

Meeradler Schwarzbrauner Adler

Gyr, Steinbracher

Hasengyr

Braune Ente
Moorente

Schwarze Sehopfente

Red-erested Poehard Netta rufina
Common Pochard Aythya ferina
Ferruginous T>\\ck Aythya nyroca
Tufted Duck Aythya fuligula
Greater Seaup Aythya marila

Klein 1760 Gill & Wright 2006

Eidergans Common Eider Somateria mollissi

Harlequin Duck Histrionicus histrionicus
Islandische Spiesente Long-tailed Duck Clangida hyemalis

Common Scoter Melanitta nigra
Schwarze Ente mit rotem und Velvet Scoter Melanitta fusca
gelbem Schnabel

Barrow’s Goldeneye Encephala islandica
Golden Auglein Common Goldeneye Bucephala clangula

Kleiner weiBkopfiger Sager Smew Mergellus alhellus
Gezopfter Sager Red-breasted Merganser Mergus serrator

Gemeiner Sager Goosander (Common Merganser) Mergus merganser

White-headed Duck Oxyura leucocephala
Osprey Pandion haliaetiis
European Honey Buzzard Pernis apivorus
Black Kite Milvus migrans

Scheerschwanzel Red Kite Milvus milvus

Beinbrecher White-tailed Eagle Haliaeetus albicilla

BMiger Geyer Bearded Vulture (Lammergeier) Gypaetus barbatus

Egyptian Vulture Neophron percnopterus
Hasengeyer Griffon Vulture Gyps fulvus

Schwanadler

Fischer, MaBwy

Fisehgeyer

WeiBer Geyer

Rostweye

-

GrauweiBer Geyer

-

-4-

Gochhausen 1710,
1732

Frisch 1733-1763

Zorn 1742, 1743

Sperber, Schmerl
Habicht

Mause-Geyer

Sperber

Habicht Hiihnergeyer
Schwarzbrauner Habicht
RauhfuB

Sperber, Sprinz
Habicht

MauB Ar

Adler

Rittel-Geyer

Rotelgeyer

Rotel-Geyerlein

Schmerl

Baum-Falcke

Zwergfalke

Steinfalck

-

Blau-FuB

Haselhun

Schwarzbrauner Falke
Haselhuhn

WeiBback, Schmerl
Hasselhuhn

Birek-Wildpret

Auerhahn

Birkhahn

Auerhahn

Auerhuhn

Kuttengyer

Cinereous (Blaek) Vulture Aegypius monachus

-

Short-toed Eagle Circaetiis gaUiciis

Buntrostiger Falek

Western Marsh Harrier Circus aeniginosus

-

Hen (Northern) Harrier Circus cyaneus

-

Pallid Harrier Circus macrourus

Montagu’s Harrier Circus pygargus

Klein 1760

Gill & Wright 2006

Sperber, Finkenfalk

Eurasian Sparrowhawk Accipiter nisus

Taubenfalk

Northern Goshawk Accipiter gentilis

BuBhart

Common Buzzard Buteo buteo

-

Rough-legged Buzzard Buteo lagopus

-

Eesser Spotted Eagle Aquila pomarina

Sehelladler

Greater Spotted Eagle Aquila clanga

-

Imperial Eagle Aquila heliaca

Goldadler

Golden Eagle Aquila clvysaetos

Mauerfalk, Wannenwaher

Common Kestrel Falco tinnunculus

-

Red-footed Faleon Falco vespertinus

Sperber

Merlin Falco columbariiis

Lerchenfalk

Eurasian Hobby Falco subbuteo

Saerefalk, BlaufuB

Saker Falcon Falco cherrug

Gyrfalk

Gyrfalcon Falco rusticolus

Edler Falk

Peregrine Falcon Falco peregrinus

Haselhuhn

Hazel Grouse Tetrastes bonasia

-

Willow Ptarmigan (Grouse) Lagopus lagopus

Schneehuhn

Rock Ptarmigan Lagopus muta

Birkhahn

Black Grouse Lyrurus tetrix

Auerhahn

Western Capereaillie Tetrao urogallus

-

Rothes Rebhun

-

Rebhun

Rebhuhn

Feldhuhn

Wachtel

Waehtel

Waehtel

-

Fasan

Fasan

-

Langschnabliges

Wasserhuhn

-5-

Gochhausen 1710,

Wasserhuhn

Frisch 1733-1763

Zorn 1742, 1743

1732

Klein gesprenkeltes

-

Wasserhuhn

Sammethuhnehen

-

Wachtelkonig

Sehnarre

-

Rotblassiges kl. Meerhuhn

Grosses Wasserhuhn

BlaBlein, Horbel

WeiBblassiges gr.

BlaBling

Wasserhuhn

Kranich

Kranieh

Keilhacken

-

-

Trapp, Saathun

Trapp

-

-

Kleines Riegerlein

Kleines Riegerlein

Sandlaufer, GrieB-Hun

Kleinster Kiewit

Braun-u.weiBscheekiges

Ditgen

-

Riegerlein

Saat-Hun

Reehter Braehvogel

Braehvogel

Grieehiseh Rebhuhn

Gemeines Rebhuhn

Gemeine Waehtel

Gemeiner Fasan

Schwarze Ralle

Rock Partridge Alectoris graeca

Red-legged Partridge Alectoris rufa

Grey Partridge Perdix perdix

Common Quail Cotiirnix cotiirnix

Common Pheasant Phasianus colchicus

Water Rail Rallus aquaticus

Klein 1760

Gill & Wright 2006

Kleiner Braehvogel

Spotted Crake Porzana porzana

Aschhtihnehen

Wachtelkonig

Meerteutbl

Schwarzes Blasshuhn

Little Crake Porzana parva

Baillon’s Crake Porzana pusilla

Corn Crake Crex crex

Common Moorhen Gallinula chloropus
Eurasian Coot Fulica atra

Grauer Kranich

Kl. Trappe, Trieltrappe

Ackertrappe, Trapp-Gans
Austermann, Strandheher
Sandregerlein, Seelerche

Common Crane Grus grus

Little Bustard Tetrax tetrax

Macqueen’s Bustard Chlamydotismacqueenii
Great Bustai'd Otis tarda

Eurasian Oystercatcher Haematopus ostralegus
Common Ringed Plover Charadrius hiaticula
Eittle Ringed Plover Charadrius dubius

Mornell

Griiner Kybitz

Kentish Plover Charadrius alexandrinus
Eurasian Dotterel Charadrius morinellus
European Golden Plover Pluvialis apricaria

Kibitz Kiewit Kybitz

Kl. Schnepfenstrandlaufer Duiserlein

Canutsvogel

- 6 -

Gochhausen 1710, Frisch 1733-1763 Zorn 1742, 1743

1732

Streitschnepfe

Haarschnepffe

Himmelsziege Himmelsziege MoB-Schnepfe

Ried-Schnepffe Doppelschnepfe

Keilhacke

Saathun GroBe Keilhacke Brachvogel

Sandlaufer

Rothbein

Kastanienbrauner
Strand laufer

Duiserlein

Grauer Kybitz Grey Plover Pluvialis squatarola

Sociable Lapwing Vanelliis gregarius
Kybitz Northern Lapwing Vanellus vanellus

Dunlin Calidris alpina
Purple Sandpiper Calidris maritima
Red Knot Calidris caniitus
Little Stint Calidris minuta

Braunroter Bracher Curlew Sandpiper Calidris ferruginea

Sanderling Calidris alba

Klein 1760 Gill & Wright 2006

Broad-billed Sandpiper Limicola falcinellus
Kampfhahnlein PaxfiPhilomachus piignax

Haarschnepfe Jack Snipe Lymnocryptes minimus

Heerschnepfe Common Snipe Gallinago gallinago

Doppelschnepfe Great Snipe Gallinago media

Eurasian Woodcock Scolopax rusticola
Black-tailed Godwit Limosa limosa
Bar-tailed Godwit Limosa lapponica
Kleiner Braacher Whimbrel Numenius phaeopus

Teutscher Braacher Brachvogel Eurasian Curlew Numenius arquata

Spotted Redshank Tringa erythropus
Rotbeinlein Common Redshank Tringa totanus

Marsh Sandpiper Tringa stagnatilis
Common Greenshank Tringa nebularia
Buntes Motthiihnchen Green Sandpiper Tringa ochrupos

Gelbbeinlein

Wood Sandpiper Tringa glare ola
Common Sandpiper Actitis hypoleucos

Steinwalzer

Steinpardel

GroBer Brachvogel

- 7 -

Gochhausen 1710,1732 Frisch 1733-1763 Zorn 1742, 1743

Kleinste Mowe

Rheinseeschwalbe

GroBe Haffmowe

See-Schwalbe

Schwarzplattige

Schwalbenmowe

Kleinere Meve

Ruddy Turnstone Arenaria interpres
Red-necked Phalarope Phalaropus lobatus
Grey Phalarope Phalaropus fulicarius

Riemenbein Black-winged Stilt Himantopus himantopus

Schabbelsclmabel Pied Avocet Recurvirostra avosetta

Steinpardel Eurasian Stone-curlew Biirhinus oedicnemus

Rothkniissel Collared Pratincole Glareola pratincola

Black-bellied Sandgrouse Pterocles orientalis
Skua Great Skua Stercorarius skua

Polmeve Arctic Skua (Parasitic Jaeger) Stercorarius parasiticus

Klein 1760

Gill & Wright 2006

Wei Be Meve, Strontjager
Fischerlein

GroBeste graue Meve
Graubraune groBe Meve
GrdBere bunte Meve

Kirrmeve

Rohrmeve

Black-legged Kittiwake Rissa tridactyla
Little Gull Larus minutus
Mediterranean Gull Larus melanocephalus
Black-headed Gull Chroicocephalus ridibindus
Common (Mew) Gull Larus canus
European Herring Gull Larus argentatus
Glaucous Gull Larus hyperboreiis
Lesser Black-backed Gull Larus fiiscus
Great Black-backed Gull Larus marinus
Little Tern Stermda albifrons
Arctic Tern Sterna paradisaea
Common Tern Sterna hirundo

- 8 -

Gochhausen 1710,
1732

Hohltaube

Ringeltaube

Turteltaube

Kuckuck

Steineule

Schuhu

Horneule

Gronlandische Taube

Seetaube

Tordalk

Seepapagey

Frisch 1733-1763

Holztaube

Ringeltaube

Turteltaube

Kuckuck

Rothe Schleyereule
Uhu

Rothes Kauzlein mit
Federohren

Zorn 1742, 1743

Hohltaube

Ringeltaube

Tiirken -Taube

Turtel-Taube

Kuckuck

Schleyer-Eule

Uhu

Ohr-Eule

Kleinstes Kauzlein

Schwarzkopf Sandwich Tern Sterna sandvicensis

Caspian Tern Hydroprogne caspia
Gull-billed Tern Gelochelidon nilotica
White-winged Tern Chlidonias lecopterus
Schwarze Meve Black Tern Chlidonias niger

Seetaube Black Guillemot Cepphus grylle

Lomme Common Guillemot (Murre) Uria aalge

Scheermesserschnabler Razorbill Alca torda

Rottchen Little Auk Alle alle

WeiBback, Pup in Atlantic Puffin Fratercula arctica

Klein 1760 Gill & Wright 2006

Holztaube Stock Dove Cohimba oenas

Ringel-Taube Common Wood Pigeon Colnmba palumbus

Collared Dove Streptopelia decaocto

Turteltaube European Turtle Dove Streptopelia turtur

Guguck Common Cuckoo Ciiculus canorus

Schleyereule Barn Owl Tyto alba

Gehorntes Kauzchen Eurasian Scops Owl Otiis scops

Schubuteule Eurasian Eagle Owl Bubo bubo

WeiBbunte schlichte Eule Snowy Owl Bubo scandiacus

Kleiner Schubus, Gehdrnter Long-eared Owl Asio otus
Kauz

Short-eared Owl Asio flammeus

Northern Hawk Owl Surnia ulula

Eurasian Pygmy Owl Glaucidium passerinum

Stein-Kautz

Kleiner Kauz

Kautzlein

Kautz

Brandeule

Andere Eule

Tagesschlaffe

Nachtschwalbe

-

-

Thurmschwalbe

Steinschalbe

EiBvogel

Eisvogel

EiBvogel

-

Bienenfresser

-

Mantel-Krahe, Blarack

Blauracke

Birkheher

Wiedehopff

Wiedehopf

Wiedehopf

WendehalB

Drehhals

Natterwindel

-9-

Gochhausen 1710,

Frisch 1733-1763

Zorn 1742, 1743

1732

Griin- Specht

Griinspecht

Griinspecht

Schwartzspecht,

Schwartzspecht

Hohlkrahe

Hohlkrahe
Bund-Specht, Roth-

Buntspecht

Baumheckel

Specht

Bund-Specht, mittel

-

-

Bund-Specht, klein

Kleiner Buntspecht

Kleinster Specht

Heide-Lerche

Heide-Lerche

Heide-Lerche

Lerche

Feldlerche

Feld-Lerche

Kautzlein
Gemeine Eule
Hexe

Mauerschwalbe

Gibraltarsehwalbe

Europaischer Eisvogel

Immenwolf

Mandelkrahe

Gemeiner Widhopf

Drehhalz

Eittle Owl Athene noctua
Tengmalm’s Owl Aegolins funereus
Tawny Owl Strix aliico
Ural Owl Strix uralensis
European Nightjar Caprimulgus europaeus
Common Swift Apus apus
Alpine Swift Apus melba
Common Kingfisher Alcedo atthis
European Bee-eater Merops apiaster
European Roller Coracias garnilus
Eurasian Hoopoe Upupa epops
Eurasian Wryneck Jy/Tx torquilla
Grey-headed Woodpecker Picus canus

Klein 1760

Gill & Wright 2006

Griinspecht Green Woodpecker Picus viridis

Schwarzer groBer Specht Black Woodpecker Dryocopus martius

GroBer Buntspecht Great Spotted Woodpecker Dendrocopos major

Middle Spotted Woodpecker Dendrocopos medius
White-backed Woodpecker Dendrocopos leucotos
Kleiner Buntspecht Eesser Spotted Woodpecker Dendrocopos minor

Specht mit auBerordentl. EiiBen Eurasian Three-toed Woodpecker Picoides tridactylus
Kobellerche Crested Lark Galerida cristata

Woodlark Lullula arbor ea

Himmelslerche Eurasian Skylark Alaiida arvensis

Gelbe Bachstelze

Graue u. weiBe

Bachstelze

Seidenschwantz

Wasser-AmBel

Zaunkdnig

Sehnee Lerehe
Erdsehwalbe

Rauehschwalbe

Hausschwalbe

Brachlerche

Pieplerche, Baumpieper
Wiesenpieper

Gelbbriistige Baehstelze

WeiBe Baehstelze

Seidenschwantz

Zaunkonig

Uferschwalbe

Hausschwalbe

Spierschwalbe

Koth-Lerche

Gereuth-Lerche

Pip-Lerehe

Gelbe Baehstelze
Blauliehe Baehstelze

Seiden-Schwantzlein

Wasser-Amsel

Zaunkdnig

- 10 -

Gdchhausen 1710, Frisch 1733-1763 Zorn 1742, 1743

1732

Braunfedrige Grasmtieke Braunellein

Roth-Kehle Rotkehlein Rot-Kehlein

Nachtigall

Naehtigall Naehtigall Naehtigall

Blaukehlein Blaukehle

Haus-Rdthling

Rothsehwanz Sehwarzkehlein Garten-Rdthling

GraB-Miieke Grosser Fliegenlnnger Kleiner Steinschmatzer

-

Horned Eark Eremophila alpestris

-

Sand Martin Riparia riparia

-

Eurasian Crag Martin Ptyonoprogne rupestris

Rauehsehwalbe

Barn Swallow Hirundo rustic a

Hausschwalbe

Common House Martin Delichon urbicum

Wiesenlerche

Tawny Pv^xiAnthus campestris

Pfeiflerche

Tree PxyiXiAnthus trivialis

-

Meadow Pipit Anthus pratensis

-

Water Px\)\\ Anthus spinoletta

Kuhstelze

Yellow Wagtail Motacilla flava

-

Grey Wagtail Motacilla cinerea

Klosterfraulein

White/Pied Wagtail Motacilla alba

Seidensehwanz

Bohemian Waxwing Bornbycilla garrulus

-

White-throated Dipper Cinclus cinclus

Winterkdnig

Winter Wren Troglodytes troglodytes

Klein 1760 Gill & Wright 2006

Braunellchen Dunnock Prunella modularis

Alpine Accentor Prunella collaris

Rotkehlein European Robin Erithacus rubeciila

GroBe Nachtigall Thrush Nightingale Luscinia luscinia

Kleine Nachtigall Common Nightingale Luscinia megarhynchos

Blaukehlein Bluethroat Luscinia svecica

Blaek Redstart Phoenicurus ochruros

Schwarzkehlchen Common Redstart Phonicuriis phoenicurus

Steinfletsehe Whinchat Saxicola rubetra

Common Stoneehat Saxicola rubicola

Stein-Klatsche

WeiBschwanz

GroBer Steinsehmatzer

Gelbsehwarzkehlehen

Northern Wheatear Oenanthe oenanthe

Stein- AmBel

Blaukopfige Amsel

-

Blaukopfige rothe Drossel

Rufous-tailed Rock Thrush Monticola saxatilis

Meer-AmBel

Ringel Amsel

Berg- Amsel

Ringdrossel

Blue Rock Thrush Monticola solitarius

Schwartz-Amsel

S ehwartz- AmBel

Schwartze Drossel

Gemeine Amsel

Ring Ouzel Turdus torquatus

-

-

-

-

Common Blackbird Turdus merula

-

-

-

-

Dusky Thrush Turdus eunomus

Krammetsvogel

Ziemer

Krammetsvogel

Krametsvogel

Fieldfare Turdus pilaris

Zipp-DroBel

Singdrossel

Sangdrossel

Zipdrossel

Song Thrush Turdus philomelos

WeindroBel

Rothdrossel, Weindrosel

Rothdrossel

Wein drossel

Redwing Turdus iliacus

Schnerr

Sehnarre

Mistier

Sehnarre

Mistle Thrush Turdus viscivorus

-

-

-

-

Grasshopper Warbler Locustella naevia

-

-

-

-

River Warbler Locustella fluviatilis

-

-

-

-

Savi’s Warbler Locustella luscinioides

-

-

-

-

Aquatic W'arhlQr Acrocephalus paludicola

-

-

-

-

Sedge ^dLxhlQx Acrocephalus schoenobaenus

-

-

Sehilfsehmatzer

-

Marsh WdxhXQX Acrocephalus palustris

- 11 -

Gdchhausen 1710,

Frisch 1733-1763

Zorn 1742, 1743

Klein 1760

Gill & Wright 2006

1732

-

-

-

-

Eurasian Reed ^ diXhlQx Acrocephalus scirpaceus

-

-

-

Bruchdrossel, Rohrdrossel

Great Reed ^ 2 ixh\QX Acrocephalus arimdinaceus

-

-

-

-

Icterine Warbler Hippolais icterina

-

-

-

-

Barred Warbler Sylvia nisoria

-

-

Witwerlein

Grasmiicke

Lesser Whitethroat Sylvia curruca

-

-

Heckenschmatzer

-

Common Whitethroat Sylvia communis

-

Fahle Grasmiicke

Dornreich

-

Garden Warbler Sylvia borin

-

Monch

Monch

Klosterwenzel

Blackcap Sylvia atricapilla

-

-

-

-

Greenish Warbler Phylloscopus trochiloides

-

-

-

-

Western Bonelli’s Warbler Phylloscopus bonelli

Weiden-Zeisig

Weidenzeisig

Weiden-Zeislein

Gold-Hahnichen

Gold-Hahnichen

Sommer-Zaunkonig
Braunfahle Grasmiicke

Goldhahnlein

Haus-Schmatzer

Schwantzmeise

Schwarzrtickiger und
Brauner Fliegenfunger

Indianischer Sperling
Langschwenzige Meise

Schwarz- und

WeiBscheckiger

Fliegenvogel

Schwanzmeise

- 12-

Gochhausen 1710,

Frisch 1733-1763

Zorn 1742, 1743

1732

-

Asch-Meise

Nonn-Meise

Kupp-MeiBe

Hauben-Maise

Schopf-Meise

Schwartz-MeiBe

Tannen Maise

Tann-Meise

Blau-MeiBe

Blau-Maise

Blau-Meise

Kohl-MeiBe

Kohl-Maise

Kohl-Meise

Blauspecht

Blau-Specht

Kleiber

Wood Warbler Phylloscopus sibilatrix

Weidenmiicke Common Chiffchaff Phylloscopus collybita

Willow Warbler Phylloscopus trochilus
Goldcrest Regulus regulus

Sommerkonig Fireerest Regulus ignicapilla

Todtenvogel Spotted Flycatcher Muscicapa striata

Red-breasted Flycatcher Ficedula parva
Collared Flycatcher Ficedula albicollis
Weibkehlein mit schwarzen Pied Flycatcher Ficedula hypoleuca
Backen

Spitzbartiger Langschwanz Bearded Reedling Panurus biarmicus

Langgeschwanzte Meise Long-tailed Tit Aegithalos caudatus

Klein 1760 Gill & Wright 2006

Sumpfmeise Marsh Tit Poecile palustris

Willow Tit Poecile montana

Haubenmeise Crested Tit Lophophanes cristatus

Tannenmeise Coal Tit Peripanis ater

Blaumeise Blue Tit Cyanistes caeruleus

Azure Tit Cyanistes cyaniis

Kohhneise Great Tit Pants major

Spechtartige Meise Eurasian Nuthatch Sitta europaea

Wallcreeper Tichodroma muraria

Baum-Reiter

Kleiner Grauspecht

Braun-Laufferlein

Europaische Baumklette

Eurasian Treecreeper Certhia familiaris

-

-

-

-

Short-toed Treecreeper Certhia brachydactyla

-

-

-

-

Eurasian Penduline Tit Remiz pendulinus

Kirschvogel

Pyrol, Widewal

Gold-Amsel

Golddrossel

Eurasian Golden Oriole Oriolus oriolus

Neun-Todter

Neuntodter

Kleiner Neuntodter

Neuntodter

Red-backed Shrike Lanius collurio

-

Mittlerer Neuntodter

Mittlerer Neuntodter

Kleiner hunter Wankrengel

Eesser Grey Shrike Lanius minor

Kruck-Elster

GrdBerer Neuntodter

GroBer Neuntodter

GrdBester Neuntodter

Great Grey Shrike Lanius excubitor

-

Kleinerer Neuntodter

Dorndreher

Kleiner rostiger Neuntodter

Woodchat Slirike Lanius senator

NuB-Heher

Heher

Wald-Heher

Holzheher

Eurasian Jay Garrulus glandarius

Aglester

Aelster

Aelster

Gemeine Aelster

Eurasian Magpie Pica pica

Tannen-Heher

Tannen-Heher

NuB-Heher

NuBheher

Spotted Nutcracker Nucifraga caryocatactes

-

-

-

-

Alpine Chough Pyrrhocorax graculus

Schwarzer Geist m. feurigen
Augen

Red-billed Chough Pyrrhocorax pyrrhocorax

Dohle

Graue Dohle

Dohle

Dohle

Western Jackdaw Corvus monedula

Rucken

Schwarze Krahe

-

Karechel

Rook Corvus frugilegus

Kleiner Rabe

Bunte Rabenkrahe

Rabe

Rabenkrahe

Carrion Crow Corvus corone

Grauliehe Krahe

Nebelkrahe

Nebelkrahe

Graubunte Krahe

Hooded Crow Corvus cornix

Golck-Rabe

Kolck-Rabe

AaB-Rabe

Gemeiner Rabe

Northern Raven Corvus corax

- 13 -

Gochhausen 1710,

Frisch 1733-1763

Zorn 1742, 1743

Klein 1760

Gill & Wright 2006

1732

Staar

Staar

Star

Gemeiner Star

Common Starling Sturnus vulgaris

-

-

Rosenstar

-

Rose-coloured Starling Pastor roseus

-

Haussperling

HauB- Sperling

Spatz

House Sparrow Passer domesticus

Wilder Sperling

Baum Sperling

Feld-Sperling

-

Tree Sparrow Passer montanus

-

Graufmk

-

Graufink

Rock Sparrow Petronia petronia

-

-

-

-

White-winged Snowfinch Montifringilla nivalis

Einck

Buchfink

Gemeiner Fink

Buchfink

Common Chaffinch Fringilla coelebs

Quecker, Zahrling

Bergfink

Bergfmk, Quaker

Bergfink

Brambling Fringilla montifringilla

-

-

-

-

European Serin Serinus serinus

Griinling

Griinfink

Griinling

Griinling

European Greenfinch CM or is chi oris

-

-

-

-

Citril Finch Carcluelis citrinella

Stieglitz

Distelfink

Stieglitz

Distelfink

European Goldfinch Carduelis carduelis

ZeiBig

Zeislein

Zeisig

Zeisiehen

Eurasian Siskin Carduelis spinus

Hanffling

Blut-Hanfling

Hanfling

Bluthanfling

Common Einnet Carduelis cannabina

-

-

-

Gelber Quittenhanfling

T wite Carduelis flavirostris

Zitscherling

Zitscherlein

-

Schwarzbaitchen

Common Redpoll Carduelis flammea

Grienitz

Kreutzschnabel

Krummsehnabel

Ki*eutzschnabel

Common Crossbill Loxia curvirostra

-

-

-

-

Parrot Crossbill Loxia pytyopsittacus

-

-

-

-

Common Rosefinch Carpodacus erythrinus

-

-

-

Parisvogel

Pine Grosbeak Pinicola enucleator

Giimpel, Thumpfaffe

Blutfink

Blut-Fink

Thumpfaff

Eurasian Bullfinch Pyrrhula pyrrhula

Kern-Beisser

Kirschfink

Kirseh-Fink

Brauner SteinbeiBer

Flawfinch Coccothraustes coccothraustes

-

-

-

-

Eapland Bunting (Eongspur) Calcar ius lapponicus

-

Schneeammer

-

Schneeammer

Snow Bunting Plectrophenax nivalis

Aemmerling

Goldammer

Goldammer

Goldammer

Yellowhammer Emberiza citrinella

- 14-

Gdchhausen 1710,

Frisch 1733-1763

Zorn 1742, 1743

Klein 1760

Gill & Wright 2006

1732

_

_

Zirlammer

Cirl Bunting Emberiza cirlus

-

-

-

-

Rock Bunting Emberiza cia

-

Fettammer

-

Fettammer

Ortolan Bunting Emberiza hortulana

Rohrsperling

Rohrammer

-

-

Reed Bunting Emberiza schoeniclus

-

Grauammer

-

Grauer Ammer

Corn Bunting Emberiza calandra

Appendix 3. Species knowledge of selected ornithologists in Central Europe: 18th and 19th centuries.

- 1 -

J.A. Naumann 1789

Bechstein 1791-1795

J.A. & J.F. Naumann 1795-1817

J.F. Naumann 1820-1844

Gill & Wright 2006

_

Eistaueher

Schwarzkehliger Ententaueher

Great Northern Eoon Gavia imrner

-

Polartaueher

Grauer Ententaueher

Polar- Seetaucher

Black- throated Loon Gavia arctica

-

Rotkehliger Taueher

Rotkehliger Ententaueher

Nord-Seetaucher

Red-throated Loon Gavia stellata

Grosser Taucher

Gehaubter SteiBfuB

GroBer Haubentaucher

GroBer Lappentaucher

Great Crested Grebe Podiceps cristatus

-

Graukehliger SteiBfuB

Graukehliger Taucher

Rothalsiger Eappentaucher

Red-necked Grebe Podiceps grisegena

-

OhrensteiBfuB

Gehdmter SteiBfuB

Gehornter Eappentaucher

Horned Grebe Podiceps auritus

-

Dunkelbrauner SteiBfuB

Ohrentaucher

Geohrter Eappentaucher

Black-necked Grebe Podiceps nigricollis

Kleiner Taueher

Kleinster SteiBfuB

Kleiner Taucher

Kleiner Eappentaucher

Little Grebe Tachybaptus nificollis

-

-

-

-

Manx Shearwater Puffimis piiffmus

Gesehaekter Sturmvogel

Kleiner Schwalbensturmvogel

European Storm-petrel Hydrobates
pelagicus

Leach's Storm-petrel Oceanodroma
leucorhoa

-

-

-

-

Northern Fulmar Fulmarus glacialis

-

Bassaniseher Pelikan

WeiBer Tolpel

-

Northern Gannet Moms bass anus

-

Kormoran

Kormoran- Scharbe

Kormoran- Scharbe

Great Cormorant Phalacrocorca carbo

-

-

-

-

European Shag Phalacrocorax aristotelis

-

GroBer Pelikan

GroBer Pelikan

Gemeiner Pelikan

Great White Pelican Pelecanus onocrotalus

Reiger

Fisehreiher

Gemeiner Reiher

Fisch-Reiher

Grey Heron Ardea cinerea

-

Purpurreiher

Purpun*eiher

Purpur-Reiher

Purple Heron Ardea purpurea

-

WeiBer Reiher

Federbuschreiher

Silberreiher

Great White Egret Ardea alba

-

Federbuschreiher

StrauBreiher

-

Little Egret Egretta garzetta

-

Rallenreiher

Rallenreiher

-

Squacco Heron Ardeola ralloides

-

Nachtreiher

Quackreiher

Nachtliche Rohrdommel

Black-crowned Night Heron Nycticorax
nycticorax

Kleine Rohrdommel

Kleine Rohrdommel

Kleine Rohrdommel

Zwergrohrdommel

Little Bittern Ixobrychus minutus

GroBe Rohrdommel

GroBe Rohrdommel

GroBe Rohrdommel

Storeh

Storch

WeiBer Storch

-

Schwarzer Storeh

Schwarzer Storch

-

WeiBer Loftier

WeiBer Loftier

-

Siehelschnabliger Nimmersatt

Sichelschnabler

-2-

J.A. Naumann 1789

Bechstein 1791-1795

J.A. & J.F. Naumann 1795

Sehwan

Hockerschwan

Gemeiner Schwan

-

Singsehwan

Singsehwan

Saatgans Moorgans

-

Blassengans

Blassengans

Wilde Gans

Wilde Gans

GroBe Graugans

-

Schneegans

Schneegans

-

WeiBwangige Gans

WeiBwangige Gans

-

Ringelgans

Ringelgans

-

Rothalsgans

-

-

-

Rostfarbene Ente

-

Brandente

Brandente

Pfeifente

Pfeifente

Pfeifente

-

Schnatterente

Mittelente

Kleine Kriickente

Kriekente

Ki'iickente

Gemeine wilde Ente

Gemeine Ente

Gemeine wilde Ente

GroBe Ldffelente

SpieBente

Pfeilschwanz

GroBe Kriickente

Knackente

Knakente

[ Ldffelente ]

Ldffelente

Ldffelente

-

Kolbenente

Rothhaubige Ente

GroBe Rohrdommel
WeiBer Storch
Schwarzer Storch
WeiBer Loftier

1817 J.F. Naumann 1820-1844

Hocker-Schwan
Gelbnasiger Schwan

Acker-Gans, Mittel-Gans
Blassen-Gans

Grau-Gans

WeiBwangen-Gans

Ringel-Gans

Rothals-Gans

Rost-Ente

Brand-Ente

Pfeif-Ente

Mittel-Ente

Kriickente

Marz-Ente

Spitz-Ente

Knak-Ente

Loffel-Ente

Kolben-Ente

Eurasian Bittern Botaurus stellaris
White Stork Ciconia ciconia
Black Stork Ciconia nigra
Eurasian Spoonbill Platalea leucorodia
Northern Bald Ibis Geronticus eremita
Glossy Ibis Plegadis falcinellus

Gill & Wright 2006

Mute Swan Cygnus olor
Whooper Swan Cygnus cygnus
Tundra Swan Cygnus columbianus
Bean Goose Anser fabilis
Greater White- fronted Goose Anser
albifrons

Greylag Goose Anser anser
Snow Goose Anser caerulescens
Barnacle Goose Branta leiicopsis
Brent Goose Branta bernicla
Red-breasted Goose Branta ruficollis
Ruddy Shelduck Tadorna ferruginea
Common Shelduck Tadorna tadorna
Eurasian Wigeon penelope
Gadwall Anas strepera
Eurasian Teal Anas crecca
Mallard Anas platyrhynchos
Northern Pintail Anas acuta
Garganey Anas qiierquedula
Northern Shoveler clypeata
Red-erested Pochard Netta rufina

[ Tafelente ]

Tafelente

Tafelente

Tafel-Ente

Common Poehard Aythya ferina

-

WeiBaugige Ente

Moorente

Moor-Ente

Eerruginous Duck Aythya nyroca

Kleine Tauchente

Reiherente

Reiherente

Reiher-Ente

Tufted Duck Aythya fiiligula

-

Bergente

Bergente

Berg-Ente

Greater Scaup Aythya marila

-

Eidergans

Eidvogel

Eider-Ente

Common Eider Somateria mollissima

-

Kragenente

Kragenente

Kragen-Ente

Harlequin Duck Histrionicus histrionicus

-

Eisente

Kleiner Pfeilschwanz

Eis-Ente

Long-tailed Duck Clangula hyemalis

-3 -

J.A. Naumann 1789

Bechstein 1791-1795

J.A. & J.F. Naumann 1795-1818

J.F. Naumann 1820-1844

Gill & Wright 2006

_

Trauer-Ente

Trauer-, WeiBbackenente

Common Scoter Melanitta nigra

-

Sammet-Ente

Sammet-, Brillenente

Sammet-Ente

Velvet Scoter Melanitta fusca

-

Spatel-Ente

-

-

Barrow’s Goldeneye Bucephala islandica

WeiBe Tauchente

Schell-Ente

Schellente

Schell-Ente

Common Goldeneye Bucephala clangula

-

WeiBer Sager

Kreutzente

Kleiner Sager

Smew Mergellus albellus

GroBe Tauchente

Langschnabliger Sager

Gemeiner Seerachen

Mittlerer Sager

Red-breasted Merganser Mergus serrator

GroBe Tauchente

Gansesager

GroBer Seerachen

GroBer Sager

Goosander (Common Merganser) Mergus
merganser

-

Ruderente

WeiBkopfige Ente

-

White-headed Duck Oxyura leucocephala

Eischaar

EluBadler

Eischaar

EluBadler

Osprey Pandion haliaetus

-

Wespenbussard

Wespenfalke

Wespen-Bussard

European Honey Buzzard Pernis apivorus

Moosweyhe

Schwarzer Milan

Schvvarzbrauner Milan

Schwarzbrauner Milan

Black Kite Milviis rnigrans

Gabelweihe,

Gabelgeyer

Rother Milan

Gabelweyhe

Rother Milan

Red Kite Milvus milvus

Adler

Seeadler, Eischadler

Seeadler

See-Adler

White-tailed Eagle Haliaeetus albicilla

-

Bartadler

-

-

Bearded Vulture (Lammergeier) Gypaetus
barbatus

-

-

-

-

Egyptian Vulture Neophron percnopterus

-

WeiBkdpfiger Geyer

WeiBkopfiger Geyer

-

Griffon Vulture Gyps fulvus

-

Grauer Geyer

Grauer Geyer

-

Cinereous (Black) Vulture Aegypius

-

-

Kurzzehiger Adler

Moos-Weihe

Rohr-Weye, Sumpf-Weie

Rohrweyhe

WeiBe Weihe

Korn-Weye

Halbweyhe

-

Wasser-Weye

Halbweyhe

-

Kleine Weihe, Milane

Wiesen-Weye

Halbweyhe

-

Sperber

Finken-Habicht

Sperber

Habicht, Eichevogel

Huhner-Habicht

Habicht

Mausehabicht

Mausebussard

Mauseaar

-

RauhfuBiger Bussard

Schneeaar

-

-

Schreiadler

-

Schreyadler

-4-

J.A. Naumann 1789

Bechstein 1791-1795

J.A. & J.F. Naumann 1795-1817

Schwarzer Adler, Kaiseradler

Steinadler

Stein-, Goldadler

Steinadler

Rudelgeyer

Thurmfalke

Thurmfalke

-

RotfuBiger Falke

RothfuBiger Falke

-

Zwergfalke

Merlin

Kleiner Lerchenfalk

Baumfalke

Lerchenfalke

-

Sackerfalke

Wiirg-, Sakerfalke

-

Islandfalke

Gerfalke, Geyerfalke

BlaufuB

Wanderfalke

Wanderfalke, Blaufalke

-

Haselhuhn

Haselhun

-

Schneehuhn

-

WeiBes Waldhuhn

Schneehun

-

Birkhuhn

Birkhan

-

Auerhuhn

Auerhan

Nattern-Adler

Rohr-Weihe

Korn-Weihe

Wiesenweihe

Finken-Habicht

Hiihner-Habicht

Mause-Biissard

RauhfuB-Bussard

Schrei-Adler

GroBer Schreiadler

monachus

Short-toed Eagle Circaetus gallicus
Western Marsh Harrier Circus aeruginosus
Hen (Northern) Harrier Circus cyaneus
Pallid Harrier Circus macrourus
Montagu’s Harrier Circus pygargus
Eurasian Sparrowhawk Accipiter nisus
Northern Goshawk Accipiter gentilis
Coinnion Buzzard Buteo buteo
Rough-legged Buzzard Buteo lagopus
Lesser Spotted Eagle Aquila pomarina
Greater Spotted Eagle Aquila clanga

J.F. Naumann 1820-1844 Gill & Wright 2006

Imperial Eagle Aquila heliaca

Stein- Adler Golden Eagle Aquila chrysaetos

Thurmfalke Common Kestrel Falco tinmmculus

Red-footed Falcon Falco vespertinus
Merlin-Falke Merlin Falco columbarius

Lerchen-Falke Eurasian Hobby Falco subbuteo

Saker Falcon Falco cherrug
Gyrfalcon Falco rusticolus

Taubenfalke Peregrine Falcon Falco peregrinus

Europaisches Haselhuhn Hazel Grouse Tetrastes bonasia

Moorschneehuhn Willow Ptaiinigan (Grouse) Lagopus

lagopus

Alpenschneehuhn Rock Ptarmigan Lagopus miita

Birkwaldhuhn Black Grouse Lyrurus tetrix

Auer-Waldhuhn Western Capercaillie Tetrao urogallus

-

Griechisches Rothuhn

-

Steinfeldhuhn

Rock Partridge Alectoris graeca

-

Rothhuhn

-

Rothfeldhuhn

Red-legged Partridge Alectoris rufa

Rebhuhn, Feldhuhn

Rebhuhn

Rebhuhn

Rebfeldhuhn

Grey Partridge Perdix perdix

Wachtel

Wachtel

Wachtel

Schlagwachtel

Common Quail Coturnix cotiirnix

-

Fasan

Fasan

Edelfasan

Common Pheasant Phasianus colchicus

[ Wasserralle ]

Wasser-Ralle

Langschnabliges Wasserhuhn

Wasser-Ralle

Water Rail Rallus aquaticus

[ Tiipfelsumpfhuhn ]

Punktiertes Meerhuhn

Gesprenkeltes Wasserhuhn

Gesprenkeltes Sumpfhuhn

Spotted Crake Porzana porzana

-

Kleines Meerhuhn

Kleines Wasserhiihnchen

Kleines Sumpfhuhn

Little Crake Porzana parva

-

-

-

-

Bail Ion’s Crake Porzana pus ilia

Wachtelkonig

Wiesenknarrer

Wachtelkonig

Wiesen- Sumpfhuhn

Corn Crake Crex crex

Grosse Wasserhenne

GriinfuBiges Meerhuhn

Rothblassiges Wasserhuhn

Gemeines Teichhuhn

Common Moorhen Gall inula chi or opus

Hurbel, Blosse

Schwarzes Meerhuhn

WeiBblassiges Wasserhuhn

Gemeines Wasserhuhn

Eurasian Coot Fulica atra

Krannich

Grauer Kranich

Ki'anich

Gemeiner Kranich

Common Crane Grus grus

-5-

J.A. Naumann 1789

Bechstein 1791-1795

J.A. & J.F. Naumann 1795-1817

J.F. Naumann 1820-1844

Gill & Wright 2006

-

Kleine Trappe

-

Zwergtrappe

Little Bustard Tetrax tetrax

Kragentrappe

Macqueen’s Bustard Chlamydotis
macqiieenii

Trapp

GroBtrappe

Trappe

GroB-Trappe

Great Bustard Otis tarda

Geschackter Austernfischer

Europaischer Austernfischer

Eurasian Oystercatcher Haematopus
ostralegus

BuntschnabL Regenpfeifer

Strandpfeiffer

Sand-Regenpfeifer

Common Ringed Plover Charadrius
hiaticula

-

Schwarzbindiger Regenpf.

Strandpfeiffer

EluB-Regenpfeifer

Little Ringed Plover Charadrius dubius

-

Dunkelbrtistiger Regenpfeifer

-

See-Regenpfeifer

Kentish Plover Charadrius alexandrinus

Dtitgen

Dummer Regenpfeifer

Diitchen, Kleiner Brachvogel

Mornel l-Regenpfeifer

Eurasian Dotterel Charadrius morinellus

Saatvogel

Goldregenpfeifer

Saatvogel, Griiner Brachvogel

Gold-Regenpfeifer

European Golden Plover Pluvialis
apricaria

-

Schwarzbauchiger Kiebitz

Grauer Kibitz

Nordischer Kibitzregenpfeifer

Grey Plover Pluvialis squatarola

Kiebitz

Gehaubter Kiebitz

Kibitz

-

Alpenstrandlaufer

Brauner Sandlaufer

-

Meerstrandlaufer

-

-

Kanuts- Strandlaufer

GroBe rothbriistige Schnepfe

-

Kleiner Sandlaufer

Kleiner Sandlaufer

-

Rothbauchiger Brachvogel

Rothbrustige Schnepfe

-

Kleiner Strandlaufer

Dreizehiger Sandlaufer

-

-

Schnepfenstrandlaufer

-

Kampfender Strandlaufer

Streitschnepfe

[ Zwergsclmepfe ]

Haarschnepfe

Haarsclmepfe, Stumme Schnepfe

Ketschnepfe

Heerschnepfe

Becassine, Katschschnepfe

-

Mittelschnepfe

Mittel-, Doppelschnepfe

Waldschnepfe,

Waldschnepfe

Waldschnepfe

Bekasse

-

Gefleckter Wasserlaufer

Seeschnepfe

-

Dunkelbrauner Wasserlaufer

Kl. Keilhaken, Pfuhlschnepfe

Kleiner Brachvogel

Mittlerer Brachvogel

Regenvogel

Keulhaken, Gr.

GroBer Brachvogel

Keilhaken

Wasser-

Schnepfe

- 6 -

J.A. Naumann 1789

Bechstein 1791-1795

J.A.& J.F. Naumann 1795-1817

GroBer Rotschenkel

Stand schnepfe

GroBer Rotschenkel

Kleiner Rotschenkel

RotfuBiger Wasserlaufer

Kleiner Rotschenkel

-

Teich wasserlaufer

Kleine Pfuhlschnepfe

GroBe Pfuhlschnepfe

GriinfuBiger Wasserlaufer

Pfuhlschepfe, Griinbein

-

Punktirter Wasserlaufer

Schwarzer Sandlaufer

Kleine Pfuhlschnepfe

W aldstrandlaufer

Gefleckter Sandlaufer

Gemeiner Kibitz Northern Lapwing Vanellus vanellus

Alpen-Strandlaufer Dunlin Calidris alpina

Purple Sandpiper Calidris maritima
Islandischer Strandlaufer Red Knot Calidris canutus

Kleiner Strandlaufer Little Stint Calidris minuta

Bogenschnabliger Curlew Sandpiper Calidris ferruginea

Strandlaufer

Ufer-Sanderling Sanderling Calidris alba

Kleiner Sumpflaufer Broad-billed Sandpiper Limicola falcinellus

Vierfarbiger Strandlaufer Ruff Philomachus pugnax

Kleine Sumpfschnepfe Jaek Snipe Lymnocry^ptes minimus

Gemeine Sumpfschnepfe Common Snipe Gallinago gallinago

GroBe Sumpfschnepfe Great Snipe Gallinago media

Gemeine Waldschnepfe Eurasian Woodcock Scolopax rustic ola

Rostgelbe Llferschnepfe Black-tailed Godwit Limosa limosa

Schwarzschwanzige Bar-tailed Godwit Limosa lapponica

Uferschnepfe

Regen-Brachvogel Whimbrel Numenius phaeopus

GroBer Brachvogel Brachvogel Eurasian Curlew Numenius

arquata

J.F. Naumann 1820-1844 Gill & Wright 2006

Dunkler Wasserlaufer Spotted Redshank Tringa erythropus

Gambett-Wasserlaufer Common Redshank Tringa totanus

Teich-Wasserlaufer Marsh Sandpiper Tringa stagnatilis

Hellfarbiger Wasserlaufer Common Greenshank Tringa nebularia
Punktirter Wasserlaufer Green Sandpiper Tringa ochrupos

Bruch-Wasserlaufer Wood Sandpiper Tringa glareola

[ Flussuferlaufer ]

Trillernder Wasserlaufer

Grauer Sandlaufer

-

Steindrehender Wasserlaufer

Mornelkibitz

-

Rothalsiger Wassertreter

Schwimmschnepfe

-

RothfuBiger Strandreuter

Strantreuter

-

BlaufiiBiger Wassersabler

-

-

Lerchengrauer Regenpfeifer

Triel, Gr. Brachvogel

-

RothfuBiges Sandhuhn

Gemeines Sandhuhn

—

Braunringiges Sandhuhn

-

Struntmeve

Struntjager

-

Dreyzehige Meve

Wintermeve

-

-

Kleine Meve

[ Lachmowe ]

Schwarzkopfige Meve

Gemeine Meve

_

Sturmmeve

Sturmmeve

-

Graue Meve

-

-

WeiBschwingige Meve

GroBe Seemeve

-

Heringsmeve

Heeringsmeve

-

Mantelmeve

-

-

Kleine Seeschwalbe

Kleine Schwalbenmeve

- 7 -

J.A. Naumann 1789 Bechstein 1791-1795

J.A.& J.F. Naumann 1795-

FluB-Uferlaufer

Grauschwanziger
Stelzenlaufer
Avosett-Sabler
Europaischer Triel

Schmarotzer-Raubmeve

Dreizehen-Meve

Zwerg-Meve

Schwarzkopf-Meve

Lach-Meve

Silber-Meve

Herings-Meve

Zwerg-Meerschwalbe
Kiisten-Meerschwal be

Common Sandpiper Actitis hypoleucos
Ruddy Turnstone Arenaria interpres
Red-necked Phalarope Phalaropiis lobatus
Grey Phalarope Phalaropus fulicariiis
Black-winged Stilt Himantopus himantopus

Pied Avocet Recurvirostra avosetta
Eurasian Stone-curlew Burhinus
oe diene mils

Collared Pratincole Glareola pratincola
Black-bellied Sandgrouse Pterocles
oriental is

Great Skua Stercorarius skua

Ai'ctic Skua (Parasitic Jaeger) Stercorarius

parasiticus

Black-legged Kittiwake Rissa tridactyla
Eittle Gull Lams minutus
Mediterranean Gull Larus melanocephalus
Black-headed Gull Chroicocephalus
ridibindus

Common (Mew) Gull Larus canus
European Herring Gull Larus argentatus
Glaucous Gull Larus hyperboreus
Lesser Black-backed Gull Larus fuscus
Great Black- backed Gull Larus marinus
Little Tern Sternula albifrons
Arctic Tern Sterna paradisaea

J.F. Naumann 1820-1844

Gill & Wright 2006

GroBe Seekrahe

Gemeine Meersehwalbe

Gemeine Schwalbenmeve

FluB-Meerscwalbe

Common Tern Sterna hirundo

-

Stubbersche Meerschalbe

-

Brand-Meerschwalbe

Sandwich Tern Sterna sandvicensis

-

Kaspische Meersehwalbe

GroBe Schwalbenmeve

Raub-Meerschwalbe

Caspian Tern Hydroprogne caspia

-

-

-

Fach-Meerschwalbe

Gull-billed Tern Gelochelidon nilotica

-

-

-

-

White- winged Tern Chlidonias lecopterus

[ Trauerseeschwalbe ]

Sehwarze Seesehwalbe

Sehwarze Schwalbenmeve

Sehwarze Seesehwalbe

Black Tern Chlidonias niger

-

Schwarzer Lumme

Schwarzes Taucherhuhn

Gryll-Teiste

Black Guillemot Cepphiis grylle

-

Dummer Lumme

Dummes Taucherhuhn

Ringel-Fumme

Common Guillemot (Murre) Uria aalge

-

Tordalk

-

-

Razorbill Alca tor da

-

Kleiner Aik

Kleiner nordischer Aik

Kleiner Krabbentaucher

Little Auk Alle alle

-

Arktischer Aik

Graukehliger Aik

Arktischer Lund

Atlantic Puffin Fratercula arctica

Gemeine Holztaube

Holztaube

Holztaube

Hohltaube

Stock Dove Columba oenas

Ringeltaube

Ringeltaube

Ringeltaube

Ringeltaube

Common Wood Pigeon Columba palumbus

-

-

-

-

Collared Dove Streptopelia decaocto

Turteltaube

Turteltaube

Turteltaube

Turteltaube

European Turtle Dove Streptopelia turtur

Kuckuk, Guckguck

Kuekuek

Kukuk

Gemeiner Kuekuek

Coimnon Cuckoo Cuculus canorus

-

Schley erkauz

Schleiereule

Schleier Kauz

Barn Owl Tyto alba

-

Kleine Ohreule

Kleine Ohreule

-

Eurasian Scops Owl Otus scops

Schuhu

Uhu

Schuhu

Uhu-Ohreule

Eurasian Eagle Owl Bubo bubo

-

Schneekauz

Schnee-Eule

-

Snowy Owl Bubo scandiacus

Horneule, Ohreule

Mittlere Ohreule

Mittlere Ohreule

Wald-Ohreule

Long-eared Owl Asio otus

Kohleule

Sumpfohreule

Wieseneule

Sumpf-Ohreule

Short-eared Owl Asio flammeus

-

Habichtseule

Falkeneule

Sperber-Eule

Northern Hawk Owl Surnia ulula

-

Kleiner Kauz

Zwergkauz

Sperlings-Eule

Eurasian Pygmy Owl Glaucidium
passerinum

Kauz

Steinkauz

Gemeiner Kautz

Stein-Kauz

Little Owl Athene noctua

-

RauchfuBiger Kauz

RauchfuBiger Kautz

Tengmalms Kauz

Tengmalm’s O^XAegolius funereus

Knappeule

Nachtkauz

Waldeule

Wald-Kauz

Tawny Owl Strix aluco

-

Habichtseule

Habichteule

Habichts-Eule

Ural Owl Strix uralensis

- 8 -

J.A. Naumann 1789

Bechstein 1791-1795

J.A.& J.F. Naumann 1795-1817

J.F. Naumann 1820-1844

Gill & Wright 2006

Tageschlafer

Ziegenmelker

Tagesschlafer

Gemeiner Tagschlafer

European Nightjar Caprimulgus europaeus

Thurmschwalbe

Thurmschwalbe

Thurmschwalbe

Mauersegler

Common Swift Apus apus

-

-

-

Alpensegler

Alpine Swift Apus melba

Wasserspecht,

Eisvogel

Eisvogel

Eisvogel

Gemeiner Eisvogel

Common Kingfisher Alcedo atthis

-

Bienenfresser

Gemeiner Bienenfresser

Europaischer Bienenfresser

European Bee-eater Merops apiaster

Mandelkrahe

Blaue Racke

Mandelkrahe

Blaurake

European Roller Coracias garndus

Wiedehopp

Wiedehopf

Wiedehopf

Europaischer Wiedehopf

Eurasian Hoopoe Upiipa epops

Wendehals,

Grauspecht

Wendehals

Grauspecht

Grauer Wendehals

Eurasian Wryneck Jynx torqudla

-

Graukopfiger Specht

Graugriiner Specht

Grauspecht

Grey-headed Woodpecker Picus canus

Griinspecht

Griinspecht

Griinspecht

Griinspecht

Green Woodpecker viridis

Schwarzspecht

Schwarzspecht

Schwarzspecht

Schwarzspecht

Black Woodpecker Dry oc opus martius

GroBer Buntspecht

Buntspecht

GroBer Buntspecht

Rothspecht

Great Spotted Woodpecker
major

WeiBspecht

Mittelspecht

Mittelspecht

Middle Spotted Woodpecker Dendrocopos
mediiis

-

Elsterspecht

WeiBriickiger Specht

WeiBspecht

White-backed Woodpecker Dendrocopos
leiicotos

Kleiner Buntspeeht

Grasspecht

Kleiner Buntspecht

Kleinspecht

Lesser Spotted Woodpecker Dendrocopos

yi'i 1 VI

-

Dreizehiger Specht

Dreizehiger Specht

Dreizehenspecht

fnlHUt

Eurasian Three-toed Woodpecker Picoides
tridactylus

Haus-, Salatlerehe

Haupenlerche

Haubenlerche

Haubenlerche

Crested Lark Galerida cristata

Heidelerche

Baumlerche

Heidelerche

Haide lerehe

Woodlark Lidlula arborea

Lerehe

Feldlerche

Eerche

Feldlerche

Eurasian Skylark yt /a wzia arvensis

-

Berglerche

-

Berglerche

Horned Lark Eremophila alpestris

-

Uferschwalbe

Uferschwabe

Uferschwalbe

Sand Martin Riparia riparia

-

Felsenschwalbe

-

Felsenschwalbe

Eurasian Crag Martin Ptyonoprogne
rupestris

Haus-,

Stachel Schwalbe
Mehlschwalbe

SpieBlerche

Hiister

Rauchschwalbe

Hausschwalbe

Brachlerche

Pieplerche, Baumpieper
Wiesenpieper

Hausschwalbe

Mehlschwalbe

Brachlerche

Spieslerehe

Hiister

J.A. Naumann 1789 Bechstein 1791-1795

J.A.& J.F. Naumann 1795-1817

Gelbe Bachstelze

Aekermann, Blaue B.
Seidenschwanz

Zaunkonig

Spanier

Roth-Kehlgen

Naehtigall

Sehildnaehtigall

Wasserpieper
Gelbe Bachstelze
Graue Bachstelze
WeiBe Bachstelze
Europ. Seidensehwanz
Wasserschwatzer
Zaunkonig
Braunelle
Alpen-Fliievogel
Rotkehlehen
Sprosser
Naehtigall

Gelbe Bachstelze

Graue Bachstelze

Blaue Baehstelze

Seidenschwanz

Wasserstaar

Zaunkonig

Spanier

Rotkehlehen

Sprosser, Gr. Naehtigall

Naehtigall

Sehildnaehtigall Blaukehlchen Blaukehlchen

Haurothschwanzchen Schwarzer Rothschwanz

Gemeiner Rotschwanz Gartenrothschwanzchen Rothschwanz

Krautlerche Braunkehliger Steinschmatzer Krautvogel

Schwarzkehliger

Steinschmatzer

Schwarzkehliger Steinsehmatzer

Rauehschwalbe

Barn Swallow Hirundo rustica

Haussehwalbe Common House Martin Delichon urbicum

Brachpieper Tawny Vv^iX Anthus campestris

Baumpieper Tree Pipit Anthiis trivialis

Wiesenpieper Meadow Pipit Anthus pratensis

J.F. Naumann 1820-1844 Gill & Wright 2006

Wasserpieper

Gelbe Bachstelze

Graue Bachstelze

WeiBe Bachstelze

Seidenschwanz

Wasserschmatzer

Zaunschliipfer

Heckenbraunelle

Alpenbraunelle

Rotkehlchensanger

Sprossersanger

Nachtigalls^ger

Blaukehlchensanger

Hausrothling

Gartenrothling

Braunkehliger

Wiesenschmatzer

Schwarkehliger

Wiesenschmatzer

Water Pipit Anthus spinoletta
Yellow Wagtail Motacilla flava
Grey Wagtail Motacilla cinerea
White/Pied Wagtail Motacilla alba
Bohemian Waxwing Bombycilla garrulus
White-throated Dipper Cinchis cinclus
Winter Wren Troglodytes troglodytes
Dunnock Prunella modularis
Alpine Aecentor Prunella collaris
European Robin Erithacus rubecula
Thrush Nightingale Luscinia luscinia
Common Nightingale Luscinia
megarhynchos
Bluethroat Luscinia svecica
Black Redstart Phoenicurus ochruros
Common Redstart Phonicurus phoenicurus
Whinehat Saxicola rubetra

Common Stonechat Saxicola rubicola

Steinpieker,

Steinklitseh

Steinsehmatzer

Steinbieker

-

Steindrossel

Steindrossel

Ringamsel,

Sehildamsel

Ringdrossel

Sehildamsel

Amsel

Amsel

Schwarze Amsel

-

-

Bergdrossel

Krammetsvogel,

Ziemer

Waeholderdrossel

Ziemer

Zippdrossel

Singdrossel

Zippdrossel

Weindrossel

Rothdrossel

Weindrossel

Sehnarre, Mistier

Misteldrossel

Sehnarre

Rohrsperling

Spitzkopf

Lerchenfarbiger Spitzkopf

- 10-

J.A. Naumann 1789

Bechstein 1791-1795

J.A.& J.F. Naumann 1795-

-

-

Griinliehgrauer Spitzkopf

[ Seggenrohrsanger ]

Seggenrohrsanger

Gelbgestreifter Rohrsehirf

-

Sehilfsanger

Kleinster Rohrsehirf

-

Sumpfrohrsanger

Olivengrauer Rohrsehirf

Rohrsperling

Teiehrolirsanger

Brauner Rohrsehirf

GroBer Rohrsperling

Rohrdrossel

GroBer Rohrsehirf

Schackmthgen

Gelbbauchiger Laubvogel Schackeruthgen

Gesperberte Grasmiicke GroBe WeiBkehle

Grauer Steinschmatzer

Northern Wheatear Oenanthe oenanthe

Steinmerle

Rufous-tailed Roek Thrush Monticola
saxatilis

-

Blue Rock Thrush Monticola solitarius

Ringdrossel

Ring Ouzel Turdus torqiiatiis

Sehwarzdrossel

Common Blackbird Turdus merula

-

Dusky Thrush Turdus eunomus

Waeholderdrossel

Fieldfare Turdus pilaris

Singdrossel

Song Thrush Turdus philomelos

Rothdrossel

Redwing Turdus iliacus

Misteldrossel

Mistle Thrush Turdus viscivorus

Buschrohrsanger

Grasshopper Warbler Locustella naevia

J.F. Naumann 1820-1844 Gill & Wright 2006

FluBrohrsanger River Warbler Locustella fluviatilis

Naehtigall-Rohrsanger Savi’s Warbler Locustella luscinioides

Seggenrohrsanger Aquatie W a.rh\er Acrocephaliis paliidicola

Sehilfrohrsanger Sedge ^ Acrocephalus

schoenobaenus

Sumpfrohrsanger Marsh Warbler Acrocephalus palustris

Teiehrohrsanger Eurasian Reed Warbler Acrocephalus

scirpaceus

Drosselrohrsanger Great Reed Warbler Acrocephalus

arundinaceus

Gartenlaubvogel leterine Warbler Hippolais icterina

Sperbergrasmtieke Barred Warbler Sylvia nisoria

Kleine WeiBkehle

Klappergrasmiicke

Kleine WeiBkehle

Fliegenschnapper

Geschwatzige Grasmiicke

Braune WeiBkehle

Grasemticke

Graue Grasmiicke

Grasemticke

Plattmonch

Mdnch

Plattmonch

-

Griiner Laubvogel

-

-

Waldlaubsanger

Laubvogelchen

-

Weidensanger

Brauner Fitis

Fitis

Fitis

Gelber Fitis

Goldammerchen

Goldhahnchen

Goldhahnchen

Goldammerchen

Goldhahnchen

Goldhahnchen

Gr. Lohrfmke

Gefleckter Fliegenfanger

GroBer Fliegenschnapper

-

Kleiner Fliegenfanger

-

-

Fliegenfanger m. d. Halsband

-

Kl. Lohrfinke

Schwarzriick. Fliegenfanger

Kleiner Fliegenschnapper

-

Bartmeise

Bartmeise

Spiegelmeise

Sehwanzmeise

Spiegelmeise

Plattmeise

Sumpfmeise

Blechmeise

- 11 -

J.A. Naumann 1789

Bechstein 1791-1795

J.A.& J.F. Naumann 1795

Blechmeise

-

Haubenmeise

Haubenmeise

Tannenmeise

Tannenmeise

Tannenmeise

Blau-, Piimpelmeise

Blaumeise

Bumpelmeise

-

Lasur-Meise

Lasurblaue Meise

Kohlmeise

Kohlmeise

Kohlmeise

Zaungrasmiicke
Dorngrasmticke
Gartengrasmiicke
Monchsgrasmucke
Dickschnabliger Laubsanger

Berglaubvogel

Waldlaubvogel

Weidenlaubvogel

Fitislaubvogel

Gelbkopfiges Goldhahnchen
Feuerkopfiges Goldhahiichen
Gefleckter Fliegenschnapper
Kleiner Fliegenlnnger
WeiBhalsiger Fliegenfanger
Sehwarzgrauer Fliegenfanger
Bartrohrmeise
Sehwanzmeise
Sumpfmeise

Lesser Whitethroat Sylvia curruca
Common Whitethroat Sylvia communis
Garden Warbler Sylvia borin
Blackcap Sylvia atricapilla
Greenish Warbler Phylloscopus
trochiloides

Western Bonelli’s Warbler Phylloscopus
bonelli

Waldlaubsanger Wood Warbler
Phylloscopus sibilatrix
Common Chiffchaff Phylloscopus collybita
Willow Warbler Phylloscopus trochilus
Goldcrest Regulus regulus
Fhecrest Regulus ignicapilla
Spotted Flycatcher Muscicapa striata
Red-breasted Flycatcher Ficedula parva
Collared Flycatcher Ficedula albicollis
Pied Flycatcher Ficedula hypoleuca
Bearded Reedling Panurus biarmicus
Long- tailed Tit Aegithalos caudatus
Marsh Tit Poecile palustris

J.F. Naumann 1820-1844 Gill & Wright 2006

Haubenmeise

Tannenmeise

Blaumeise

Kohlmeise

Willow Tit Poecile montana
Crested Tit Lophophanes cristatus
Coal Tit Peripanis ater
Blue Tit Cycmistes caeruleus
Azure Tit Cyanistes cyanus
Kohlmeise Great Tit Par us major

Blauspecht, Kleiber

Kleiber

Blauspecht

-

Mauer-Baumlaufer

Mauerlaufer

Baumlaufer

Baumlaufer

Baumlaufer

-

-

Baumlaufer

Beutelmeise

Beutelmeise

Pfmgstvogel

Pirol

Pfmgstvogel

Neuntbdter,

Rotriickiger Wurger

Kleiner Neundtdter

Neintodter

-

Grauer Wurger

Gemeiner grauer Wurger

GroBe Kriickelelster

Gemeiner Wurger

GroBer grauer Wurger

Kleine Kilickelelster

Rothkopfiger Wurger

Rothkopfiger Wurger

Holzschi'eyer

Eichelhaher

Holzheher

Elster

Elster

Elster

NuBheher,

NuBkrahe

Tannenheher

Tannenheher

-

Schneekrahe

-

-

Steinkrahe

-

Dohle, Schneekrahe

Dohle

Dohle

Schwarze Feldkrahe

Saatkrahe

Saatkrahe

Schwarze Krahe

Rabenkrahe

Rabenkrahe

Schildkrahe

Nebelkrahe

Nebelkrahe

Rabe

Kolkrabe

Rabe

Staar

Staar

Staar

- 12-

J.A. Naumann 1789

Bechstein 1791-1795

J.A.& J.F. Naumann 1795

Haussperling

Rosenfarbige Drossel
Haussperling

Rosenfarbige Drossel
Sperling

Europaiseher Kleiber Eurasian Nuthateh Sitta europaea

Alpen-Mauerlaufer Wallcreeper Tichodroma miiraria

Grauer Baumlaufer Eurasian Treecreeper Certhia familiaris

Short-toed Treecreeper Certhia
brachydactyda

Beutelrohrmeise Eurasian Penduline Tit Remiz pendulinus

Kirschpirol Eurasian Golden Oriole Oriolus oriolus

Rotriickiger Wurger Red-backed Shrike Lanius colhirio

Grauer Wurger
GroBer Wurger
Rotkoptlger Wurger
Eichelheher
Elsterrabe
Tannenheher

Steinkrahe

Dohlenrabe

Saatrabe

Krahenrabe

Schildrabe

Kolkrabe

Gemeiner Star

Eesser Grey Shrike Lanins minor
Great Grey Shrike Lanius excuhitor
Woodchat Shrike Lanius senator
Eurasian Jay Garruliis glandarius
Eurasian Magpie Pica pica
Spotted Nutcracker Niicifraga
caryocatactes

Alpine Chough Pyrrhocorax graculus
Red-billed Chough Pyrrhocorax
pyrrhocorax

Western Jackdaw Corvus monedula
Rook Corvus frugilegus
CanJon Crow Connis corone
Hooded Crow Corvus cornix
Northern Raven Conms corax
Common Starling Sturnus vulgaris

J.F. Naumann 1820-1844 Gill & Wright 2006

Rose-coloured Starling Pastor roseus
Haussperling House Sparrow Passer domesticus

Holz-, Rohrsperling

Eeldsperling

Rohrsperling

-

Graufink

Graufink

-

Schneelink

Schneelink

Einke

Gemeiner Link

Fink

Quacker

Bergfink

Quaker

-

Girlitz

-

Schwunsch

Griinling

Schwunsch

-

Citronengirlitz

-

Stieglitz, Distellink

Distelzeisig

Stieglitz

Zeisig

Erlenzeisig

Zeisig

Hanfling

Bluthanfling

Rothbrtistiger Hanfling

Steinhanfiing

Arkti seller Fink

Berghanfling

Schittscherling

Bergzeisig, Zitscherlein

Tschettchen

Krinitz, Kreutzvogel

F ichten-Kreuzschnabel

Creutzvogel

-

Kiefem-Kreuzschnabel

GroBer Kreutzschnabel

-

-

Brandfmk

-

Haken-Kreuzschnabel

Fichtenkernbeisser

Gimpel, Dompfaff

Rothbrtistiger Gimpel

Dompfaffe

Kirschlincke

Kirsch-KernbeiBer

KernbeiBer

-

Grauer Sporner

Ammerlink

Schneelerche

Schneeammer

Schneelerche, Bergammer

Griinschling, Gehling

Goldammer

Grunschling

-

Zaunammer

-

-

Zippammer

-

- 13 -

J.A. Naumann 1789

Bechstein 1791-1795

J.A.& J.F. Naumann 1795

Gartenammer

Ortolan, Gartenammer

Feldsperling

Steinsperling

Sehneetlnk

Buehfink

Bergfmk

Girlitzhanfling

Griinhantling

Zitronenzeisig

Distelzeisig

Erlenzeisig

Bluthanfling

Berghanfling

Birkenzeisig

F iehtenkreuzsehnabel

Karmingimpel

Fiehtengimpel

Rothgimpel

KirsehkernbeiBer

Sehneespornammer

Goldammer

Zaunammer

Zippammer

Tree Sparrow Passer montanus
Rock Sparrow Petronia petronia
White-winged Snowfmch Montifringilla
nivalis

Common Chaffinch Fringilla coelebs
Brambling Fringilla montifringilla
European Serin Seriniis seriniis
European Greenfinch Chloris Moris
Citril Finch Carduelis citrinella
European Goldfinch Carduelis carduelis
Eurasian Siskin Carduelis spinus
Common Linnet Carduelis cannabina
Twite Carduelis flavirostris
Common Redpoll Carduelis flammea
Common Crossbill Loxia curvirostra
Parrot Crossbill Loxia pytyopsittacus
Common Rosefmch Carpodacus erythrinus
Pine Grosbeak Pinicola enucleator
Eurasian Bullfinch Pyrrhida pyrrhula
Hawfinch Coccothraiistes coccothraustes
Lapland Bunting (Longspur) Calcarius
lapponicus

Snow Bunting Plectrophenax nivalis
Yellowhammer Emberiza citrinella
Cirl Bunting Emberiza cirlus
Rock Bunting Emberiza cia

17 J.F. Naumann 1820-1844

Gill & Wright 2006

Gartenammer

Ortolan Bunting Emberiza hortulana

Schiebgen

Stmmpfweber

Rohrammer Schiebichen Rohrammer Reed Bunting Emberiza schoeniclus

Grauammer Stmmpfweber Grauammer Corn Bunting Emberiza calandra

1

Appendix 4. Modern German and English bird names.

Eistaueher Great Northern Loon Gavia immer

Prachttaucher Black-throated Loon Gavia arctica

Stemtaucher Red-throated Loon Gavia stellata

Haubentaucher Great Crested Grebe Podiceps cristatus

Rothalstaucher Red-necked Grebe Podiceps ^rise^ena

Ohrentaucher Horned Grebe Podiceps aiiritus

Schwarzhalstaucher Black-necked Grebe Podiceps ni^ricollis

Zwergtaucher Little Grebe Tachybaptus mficollis

Atlantiksturmtaiicher Manx Sheai^water Piiffmus piiffmiis

Sturmschwalbe European Storm-petrel Hydrobates pela^icus
Wellenlaufer Leach's Storm-petrel Oceanodroma leucorhoa

Eisstiinnvogel Northern Lulmar Fulmariis ^lacialis

Basstolpel Northern Gannet Moms bass anus

Konnoran Great Cormorant Phalacrocorax carbo

Krahenscharbe European Shag Phalacrocorax aristotelis

Rosapelikan Great White Pelican Pelecanus onocrotalus

Graureiher Grey Heron Ardea cinerea

Purpurreiher Purple Heron Ardea purpurea

Silberreiher Great White Egret Ardea alba

Seidenreiher Little Egret E^retta ^arietta

Rallenreiher Sqnacco Heron Ardeola ralloides

Nachtreiher Black-crowned Night Heron Nycticorax nycticorax

Zwergdomniel Little Bittern Ixobrychus minutus

Rohrdommel Eurasian Bittern Botaurus stellar is

Weifistorch White Stork Ciconia ciconia

Schwarzstorch Black Stork Ciconia ni^ra

Loftier Eurasian Spoonbill Platalea leucorodia

Waldrapp Northern Bald Ibis Geronticus eremita

Sichler Glossy Ibis Fle^adis falcine Hus

Hockerschwan Mute Swan Cygmis olor

Singschwan Whopper Swan Cy^nns cy^mis

Zwergschvvan Tundra Swan Cy^mis columbianus

Saatgans Bean Goosq Anser fabilis

Blassgans Greater White-fronted Goose Anser albifrons

Graugans Greylag Goose Anser anser

Schneegans Snow Goose Anser caenilescens

WeiBwangengans Barnacle Goose Branta leiicopsis

Ringelgans Brent Goose Branta bernicla

Rothalsgans Red-breasted Goose Branta ruficollis

Rostgans Ruddy Shelduck Tadorna ferru^inea

Brandgans Common Shelduck Tadorna tadorna

Pfeifente Eurasian Anas penelope

S chnatterente G adwal 1 A nas strepera

Kidckente Eurasian Teal Anas crecca

Stockente Mallard Anas platyrhynchos

SpieBente Northern Pintail Anas acuta

Knakente Garganey Anas querquedula

1

2

Loffelente Northern Shoveler Anas clypeata

Kolbenente Red-crested Pochard Netta rufina

Tafelente Common Pochard Aythva ferina

Moorente Ferruginous T>\xckAythva nyroca

Reiherente Tufted Duck Aythya fuli^ula

Bergente Greater Scaup Aythya marila

Eiderente Common Eider Somateria mollissirna

Kragenente Harlequin Duck Histrionicus histrionicus

Eisente Eong-tailed Duck Clan^da hyemalis

Trauerente Common Scoter Melanitta ni^ra

Samtente Velvet ^coXqx Melanitta fusca

Spatelente Barrow's Goldeneye Bucephala islandica

Schellente Common Goldeneye Bucephala clanQula

Zwergsager Smew Mer^ellus albellus

Mittelsager Red-breasted Merganser Mer^us s err at or

Gansesager Goosander (Common Merganser) Mergus merganser

WeiBkopf-Ruderente White-headed Duck Oxyura leucocephala

Fischadler Osprey Pandion haliaetiis

Wespenbussard European Honey Buzzard Pernis apivorus

Schwarzmilan Black Kite Milviis migrans

Rotmilan Red Kite Milvus milviis

Seeadler White-tailed Eagle Haliaeetus albicilla

Bartgeier Bearded Vulture (Lammergeier) Gypaetus barbatus

Schmutzgeier Egyptian Vulture Neophron percnopterus

Gansegeier Griffon Vulture Gyps fidviis

Monchsgeier Cinereous (Black) N ultnxQ Aegypius monachus

Schlangenadler Short-toed Eagle Circaetus gallicus

Rohrweihe Western Marsh Harrier Circus aeruginosus

Komweihe Hen (Northern) Harrier Circus cyaneus

Steppenweihe Pallid Harrier Circus macrourus

Wiesenweihe Montagu’s Hanier Circus pygargus

Sperber Eurasian SpaiTowhawkrtcc/rt/Y^r nisiis

Habicht Northern Goshawk Accipiter gentilis

Mausebussard Common Buzzard Buteo biiteo

RauhfuBbussard Rough-legged Buzzard Buteo lagopus

Schreiadler Lesser Spotted Eagle Aqiiila pomarina

Schelladler Greater Spotted Eagle Aquila clanga

Kaiseradler Imperial Eagle Aquila heliaca

Steinadler Golden Eagle Aquila chry’saetos

Turmfalke Common Kestrel Falco tinnuncidus

RotfuBfalke Red-footed Falcon Falco vespertinus

Merlin Merlin Falco columbarius

Baumfalke Eurasian Hobby Falco subbuteo

Wurgfalke Saker Falcon Falco cherru

Gerfalke Gyrfalcon Falco rusticolus

Wanderfalke Peregrine Falcon Falco peregrinus

Haselhuhn Hazel Grouse Tetrastes bonasia

Moorschneehuhn Willow Ptarmigan (Grouse) Lagopus lagopus

Alpensclineehuhn Rock Ptai'migan Lagopus miita

2

3

Birkhuhn Black Grouse Lyrurus tetrix

Auerhuhn Western Capercaillie Tetrao iiro^allus

Steinhuhn Rock Partridge Alectoris ^raeca

Rothuhn Red-legged Partridge Alectoris rufa

Rebhuhn Grey Partridge Perdix perdix

Wachtel Common Quail Cotiirnix coturnix

Fasan Common Pheasant Phasianns colchicus

Wasserralle Water Rail Rallus aquaticus

Tupfelsumpflruhii Spotted Crake Porzana porzana

Kleines Sumpfhuhn Little Crake Porzana parva

Zwergsumpfhuhn Baillon’s Crake Porzana pus ilia

Wachtelkonig Corn Crake Crex crex

Teichhuhn Common Moorhen Gallimda chloropus

Blasshului Eurasian Coot Fulica atm

Kranieh Common Crane Grus ^rus

Zwergtrappe Little Bustard Tetrax tetrax

Ki^agentrappe Macqueen’s Bustard Chlamydotis macqueenii

Grofitrappe GroBtrappe Great Bustard Otis tarda

Austernfischer Eurasian Oystercatcher Haematopus ostrale^us
Sandregenpfeifer Common Ringed Plover Charadrius hiaticula

Flussregenpfeifer Little Ringed Plover Charadriiis diibiiis

Seeregenpfeifer Kentish Plover Charadrius alexandriniis

Mornell Eurasian Dotterel Charadrius morinellus

Goldregenpfeifer European Golden Plover Pluvialis apricaria

Kiebitzregenpfeifer Grey Plover Pluvialis squatarola

Kiebitz Northern Lapwing Vane Hits vanellus

Alpenstrandlaufer Dunlin Calidris alpina

Meerstrandlaufer Purple Sandpiper Calidris maritima

Knutt Red Knot Calidris canutus

Zwergstrandlaufer Little Stint Calidris minuta

Sichelstrandlaufer Curlew Sandpiper Calidris ferru^inea

Sauderling Saiideiiing Calidris alba

Sumpflaufer Broad-billed Sandpiper Limicola falcinellus

Kampflaufer Ruff Philomachus pu^nax

Zwergschnepfe Jack Snipe Lymnocryptes minimus

Bekassine Common Snipe Gallina^o ^allina^o

Doppelschnepfe Great Snipe Gallina^o media

Waldschnepfe Eurasian Woodcock Scolopcix nisticola

Uferschnepfe Black-tailed Godvvit Limosa limosa

Pfuhlschnepfe Bar-tailed Godwit Limosa lapponica

Regenbrachvogel Whiinbrel Niimenius phaeopus

Grofier Brachvogel Eurasian Curlew Niimenius arquata

Dunkler Wasserlaufer Spotted Redshank Trin^a erythropus

Rotschenkel Common Redshank Trin^a totamis

Teichwasserlaufer Marsh Sandpiper Trin^a staj^natilis

Grunschenkel Common Greenshank Trin^a nehularia

Waldwasseiiaufer Green Sandpiper Trin^a ochrupos

Bruchwasserlaufer Wood Sandpiper Trin^a ^lareola

Flussuferlaufer Common Sandpiper Actitis hypoleucos

3

Steinwalzer Ruddy Turnstone Arenaria interpres

Odinshuhnchen Red-necked Phalarope Phalaropiis lobatiis

Thorshtihnchen Grey Phalarope Phalaropus fiilicarius

Stelzenlaufer Black-winged Stilt Himantopus himantopus

Sabelschnabler Pied Avocet Recurvirostra avosetta

Triel Eurasian Stone-curlew Burhimis oedicnemus

Rotflugelbrachschalbe Collared Pratincole Glareola pratincola

Sandfliighuhn Black-bellied Sandgrouse Pterocles orientalis

Skua Great Skua Stercorariiis skua

Schmarotzrerraubmowe Arctic Skua (Parasitic Jaeger) Stercorarius parasiticus

Dreizehenmowe Black-legged Kittiwake Rissa tridactyla

Zwergmowe Little Gull Lariis minutus

Schwarzkopfmowe Mediterranean Gull Lams melanocephalus

Lachmowe Black-headed Gull Chroicocephalus ridibindiis

Sturmmowe Common (Mew) Gull Lams caniis

SilbeiTubwe European Herring Gull Larus ar^entatus

Eismbwe Glaucous Gull Larus hyperboreus

Heringsinowe Lesser Black-backed Gull Larus fuscus

Mantelmowe Great Black-backed Gull Larus marinus

Zwergseeschwalbe Little Tern Stenmla albifrons

Kustenseeschwalbe Arctic Tern Sterna paradisaea

Llussseeschwalbe Common Tern Sterna hirimdo

Brandseeschwalbe Sandwich Tern Sterna sandvicensis

Raubseeschwalbe Caspian Tern Hydropro^ne caspia

Lachseeschwalbe Gull-billed Tern Gelochelidon nilotica

Weififlugelseeschwalbe White- winged Tern Chlidonias lecopterus

Trauerseeschwalbe Black Tern Chlidonias ni^er

Gryllteiste Black Guillemot Cepphiis ^rylle

Trottellumme Common Guillemot (Muitc) Uria aal^e

Tordalk KsiZOYhiW Alca torda

Krabbentaucher Little Auk yl//g alle

Papageitaucher Atlantic Puffin Fratercula arctica

Hohltaube Stock Dove Columba oenas

Ringeltaube Common Wood Pigeon Cohimba palumbus

Turkentaube Collared Dove Streptopelia decaocto

Turteltaube European Turtle Dove Streptopelia turtur

Kuckuck Common Cuckoo Cuculus canorus

Schleiereule Bai’n Owl Tyto alba

Zwergohreule Eurasian Scops Owl Otus scops

Uhu Eurasian Eagle Owl Bubo bubo

Schnee-Eule Snowy Owl Bubo scandiacus

Waldohreule Long-eared Owl Asio otus

Sumpfoln^eule Short-eared Osn\ Asio flammeus

Sperbereule Northern Hawk Owl Siirnia ulula

Sperlingskauz Eurasian Pygmy Owl Glaiicidium passerinum

Steinkauz Little Owl Athene noctiia

RauhfuBkauz Tengmalm's Owl Ae^olius funereus

Waldkauz Tawny Owl Strix aluco

Habichtskauz Ural Owl Strix uralensis

5

Ziegenmelker European Nightjar Caprimulgus europaeus

Mauersegler Common Swift apiis

Alpensegler Alpine Swift Apus melba

Eisvogel Common Kingfisher Alcedo atthis

Bienenfresser European Bee-eater Merops apiaster

Blauracke European Roller Coracias ^arnilus

Wiedehopf Eurasian Hoopoe Upiipa epops

Wendehals Eurasian Wryneck Jynx torquilla

Grauspecht Grey-headed Woodpecker Picus canus

Griinspecht Green Woodpecker Picus viridis

Schwarzspecht Black Woodpecker Dryocopus martins

Buntspecht Great Spotted Woodpecker Dendrocopos major

Mittelspecht Middle Spotted Woodpecker medius

Weifiriickenspecht White-backed WoodpQckQr Dendrocopos leucotos

Kleinspecht Eesser Spotted Woodpecker Dendrocopos minor

Dreizehenspecht Eurasian Three-toed Woodpecker Picoides tridactylus

Haiibenlerclie Crested Lark Galerida cristata

Heidelerche Woodlark Lullula arbor ea

Eeldlerche Eurasian Skylark A arvensis

Ohrenlerche ELomed Lark Ereniophila alpestris

Uferschwalbe Sand Martin Riparia riparia

Eelsenschwalbe Eurasian Crag Martin Ptyonopro^ne rupestris

Raiichschwalbe Bam Swallow Hirimdo nistica

Mehlschwalbe Common ELouse Martin Delichon urbiciim

Brachpieper Tawny Pipit Anthus campestris

Baumpieper Tree Pipit Anthiis trivialis

Wiesenpieper Meadow Pipit Anthus pratensis

Wasserpieper Water Pipit Anthus spinoletta

Schafstelze Yellow Wagtail Motacilla flava

Gebirgsstelze Grey Wagtail Motacilla cinerea

Bachstelze White/Pied Wagtail Motacilla alba

Seidenschwanz Bohemian Waxwiiig Bombycilla ^arrulus

Wasseramsel White-throated Dipper Cinclus cinclus

Zaunkonig Winter Wren Troglodytes troglodytes

Heckenbraunelle Dunnock Prunella modularis

Alpenbraunelle Alpine Accentor Prunella coUaris

Rotkehlchen European Robin Erithacus nibecula

Sprosser Thi’iish Nightingale Luscinia luscinia

Nachtigall Common Nightingale Luscinia me^arhynchos

Blaiikehlchen Bluethroat Luscinia svecica

Hausrotschwanz Black Redstart Phoenicunis ochriiros

Gartenrotschwanz Common Redstart Phoniciirus phoenicurus

Braunkehlchen Whinchat Saxicola rubetra

Schwarzkehlchen Common Stonechat Saxicola rubicola

Steinschmatzer Northern Wheatear Oenanthe oenanthe

Steinrotel Rufous-tailed Rock Thrush Monticola saxatilis

Blaumerle Blue Rock Thrush Monticola solitarius

Ringdrossel Ring Ouzel Turdiis torquatus

Amsel Common Blackbird Turdiis merula

5

6

Rostflugeldrossel Dusky Thrush Turdus eunomus

Wacholderdrossel Fieldfare Tiirdiis pilaris

Singdrossel Song Thrush Turdus philomelos

Rotdrossel Redwing Turdus iliaciis

Misteldrossel Mistle Thrush Turdus viscivoms

Feldschwiii Grasshopper Warbler Locustella naevia

Schlagschwirl River W Sirhlcr Locustella fluviatilis

Rohrschwirl Savi's Warbler Locustella luscinioides

Seggenrohrsanger Aquatic Wmbl^r Acrocephalus paludicola

Schilfrohrs^ger Sedge W 3xb\Qr Acrocephalus schoenobaemis

Sumpfrohi’sanger Marsh Acrocephaliis paliistris

Teichrohrsanger Eurasian Reed W dixh\QX Acrocephaliis scirpaceus
Drosselrohrsanger Great Reed ^ 2 ixh\Qr Acrocephalus arundinaceus

Gelbspotter Icterine V^ai'blQV Hippolais icterina

Sperbergrasmticke BaiTed Wai’bler Sylvia nisoria

Klappergrasmticke Lesser Whitethroat Sylvia curruca

Domgrasmucke Common Whitethroat Sylvia communis

Gartengrasmucke Garden Warbler Sylvia borin

Mdnchsgrasmiicke Blackcap Sylvia atricapilla

Griiner Laiibsanger Greenish Warbler Phylloscopus trochiloides
Berglaubsanger Western Bonelli's Warbler Phylloscopus honelli

Waldlaubsanger Wood Phylloscopus sibilatrix

Zilpzalp Common Chiffchaff Phylloscopus collybita

Fitis Willow ^?ixh\Qr Phylloscopus trochilus

Wintergoldhahnchen Goldcrest Re^uliis re^iiliis

Sommergoldhalinchen FIxqcyqsI Re^uliis i^niccipilla

Grauschnapper Spotted Flycatcher Muscicapa striata

Zwergschnapper Red-breasted Flycatcher Ficedula parva

Halsbandsclmapper Collai^ed Flycatcher Ficedula albicollis

Trauerschnapper Pied Flycatcher Ficedula hypoleuca

Bartmeise Bearded Reedling Paminis biarmicus

Schwanzmeise Long-tailed Tit Ae^ithalos caudatus

Sumpfmeise Marsh Tit Poecile palustris

Weidenmeise Willow Tit Poecile montana

Haubenmeise Crested Tit Lophophanes cristatus

Tannenmeise Coal Tit Periparus ater

Blaumeise Blue Tit Cyanistes caenileiis

Lasurmeise Azure Tit Cyanistes cyanus

Kohlmeise Great Tit Pams major

Kleiber Eurasian Nuthatch Sitta europaea

Mauerlaufer Wallcreeper Tichodroma muraria

Waldbaumlaufer Eurasian Treecreeper Certhia familiaris

Gartenbaumlaiifer Shoit-toed Treecreeper Certhia brachydactyla

Beutelmeise Eurasian Penduline Tit Remiz pendulinus

Pirol Eurasian Golden Oriole Oriolus oriolus

Neuntoter Red-backed Shrike Lanins collurio

Schwarzstirnwtirger Lesser Grey Shrike Lanins minor

Raubwurger Great Grey Shrike Lanius excubitor

RotkopBvurger Woodchat Shi'ike Lanins senator

6

7

Eichelhaher Eurasian Jay Garrulus ^landarius

Elster Eurasian Magpie Pica pica

Tannenhaher Spotted Nutcracker Niicifra^a caryocatactes

Alpendohle Alpine Chough Pyrrhocorax oraculus

Alpenkrahe Red-billed Chough Pyrrhocorax pyrrhocorax

Dohle Western Jackdaw Coi'vus monedula

Saatkrahe Rook Corvus fru^ile^us

Rabenkrahe Carrion Crow Corvus corone

Nebelkrahe Hooded Crow Corviis cornix

Kolkrabe Northern Raven Corvus corax

Star Common Starling Sturnus vulgaris

Rosenstar Rose-coloured Starling Pastor roseus

Haussperling House Sparrow Passer dornesticus

Feldsperling Tree Sparrow Passer montanus

Steinsperiing Rock Sparrow Petronia petronia

Schneesperling White-winged Snovvfmch Montifrin^illa nivalis

Buclifink Common Chaffinch Frin^illa coelebs

Bergfink Brambling Frin^illa montifr infill a

Girlitz European Serin Serinus serinus

Griiiifink European Greeatlnch Chi oris chloris

Zitronengirlitz Citril Finch Cardiielis citrinella

Stieglitz European Goldfinch Cardiielis carduelis

Erlenzeisig Eurasian Siskin Cardiielis spinus

Bluthanfling Common Einnet Carduelis cannabina

Berghanfling Twite Cardiielis f lav irostr is

Birkenzeisig Common Redpoll Carduelis flammea

Fichtenkreuzschnabel Common Crossbill Loxia curvirostra
Kiefernkreuzschnabel Parrot Crossbill Loxia pytyopsittacus

Kamiingimpel Common Rosefmch Carpodacus erythrimis

Hakengimpel Pine Grosbeak Pinicola enucleator

Gimpel Eurasian Bullfinch Pyrrhula pyrrhiila

Kembeifier Hawfmch Coccothraiistes coccothraiistes

Spornammer Lapland Bunting (Eongspur) Calcariiis lapponicus

Schneeammer Snow Bunting Plectrophenax nivalis

Goldainmer Yellowhainmer Ember iza citrinella

Zaunammer Girl Bunting Emberiza cirlus

Zippammer Rock Bunting Emheriza da

Ortolan Ortolan Bunting Emberiza hortiilana

Rolirammer Reed Bunting Emberiza schoeniclus

Grauammer Corn Bunting Emberiza calandra

1

7

Haffer et al. (2014^ Errata

1. R 26, left column, fourth line from bottom, for '2282' read '282'.

2. R 47, left column, eleventh line from bottom, for '1165' read '165'.

3. R. 102, References, in Bechstein JM (1793-1812) delete last two lines: 'Englischen

Niirnberg'.

4. R. 105, References, in Groebbels F (1910), for 'Vogelwelt' read 'Vogelwelf .

5. R. 106, References, between Haffer J (2001) and Haffer J (2006) insert 'Haffer J (2003) Christian
Ludwig Brehm (1787-1864) iiber Spezies und Subspezies von Vogeln. Journal fur Ornithologie
144: 120-147'.

6. Online Appendix 1, p. 14, column 'Ray 1678', for 'Gouldfmch' read 'Goldfinch'.

BLANK PAGE
INSERTED

Bonn zoological Bulletin - Supplementum 59: 1-116

©ZFMK

Jurgen Haffer and Hans Hudde have brought together a wealth of information in this survey of the deve-
lopment of ornithology in Central Europe, supplemented by a brief outline of the modernizing process in
the United Kingdom. This is quite fitting, since the monograph begins with perhaps the greatest of the
early pioneers, the 17th-century Englishmen John Ray and Erancis Willughby. Although Dr. Haffer has
written many papers and articles on the subject, this monograph pulls together all of his ideas and takes
into account the latest literature, hence creating a masterly and definitive overview of the history of orni-
thology in Central Europe. But the book is more than that: in addition to the ornithological history
reflected in the great works and discoveries of the past, Hans Hudde has added to the text an innovati-
ve set of tables that enhance the written history by showing the expansion of species knowledge as seen
in German bird names through the centuries. His tables (available as online appendices due to their size)
begin with Hans Sachs (1 531) and end with J.E. Naumann (1820-1844). These appendices will be an inva-
luable tool for non-German researchers since English, modern German, and scientific names are also
included.

The gradual refining of species recognition and identification is also made visible in the many illustra-
tions accompanying the text. A further aspect that was close to Dr. Haffer's heart was to bring to the
attention of the wider ornithological public J.A. Naumann's first book Der Vogelsteller [The Bird-Trapper]
of 1 789. The elder Naumann's encyclopedic knowledge of birds is well illustrated by the lengthy extracts
from this fascinating work.

This monograph will probably be the last published work by one of the great names in modern orni-
thological writing, Jurgen Haffer, and in a way it is sadly appropriate that it should be his final word about
the subject on which he was a world authority: the history of ornithology in Gentral Europe. However the
book remains tragically unfinished, in the sense of not being entirely from his hand; he died suddenly in
April 2010 unable to conclude the work. A detailed account is given in a preface by the translator.

Bonn zoological Bulletin - Supplementum - Vol. 59 (2014)
Managing Editor: Thomas Wesener

Zoologisches Eorschungsmuseum Alexander Koenig -
Leibniz-lnstitut fur Biodiversitat der Tiere (ZEMK)

Adenaueralllee 160, D-53113 Bonn, Germany

ISBN-13: 978-3-925382-63-5
ISSN: 0302-671X

Gover illustration:

Hawfinch Coccothraustes coccothraustes, male (top) and female
J.A & J.E. Naumann (1795-1817)

Bundesministerium
furBildung
und Forschung

Ministerium fur Innovation,
Wissenschaft und Forschung
des Landes Nordrhein-Westfalen

Bonn zoological Bulletin - Supplementum Volume 59 (2014)