THE NATURAL HISTORY BURNET MOTHS - Axel F. Hofmann & W. Gerald Tremewan PART III. 11 - ResearchGate
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Axel F. Hofmann & W. Gerald Tremewan
THE NATURAL HISTORY
OF
BURNET MOTHS
PART III. 1
Mu
en
se h
um nc
Witt Mü
Museum Witt Munich
2020
iii
e Natural History of
Burnet Moths (Zygaena Fabricius, 1775)
(Lepidoptera: Zygaenidae)
Part 3
Species section
Axel F. Hofmann & W. Gerald Tremewan (†)
Premium Edition, limited ( /100)
Proceedings of the Museum Wi Munich
2020
Part 3_Premium Edition AHrevised.indd iii 11.02.2021 12:41:26
iv
Text: Axel Hofmann, W. Gerald Tremewan (†)
Text editors: Adrian Spalding, W. Gerald Tremewan (†), Gregor Markl
Technical editor: Axel Hofmann
Layout: Axel Hofmann
Cover foto (Zygaena fausta suevica) by Axel Hofmann
Cover layout by Harry Sulak
Gedruckt mit Unterstützung durch die
omas-Wi-Stiung
zur Förderung der Wissenscha und Forschung im
Bereich der Zoologischen Systematik
is publication should be cited as:
Hofmann, A. F. & W. G. Tremewan, 2020. e Natural History of Burnet Moths (Zygaena Fabricius, 1775) (Lepidoptera: Zygaenidae),
Part 3.1. – Proceedings of the Museum Wi Munich, Premium Edition 6 (3.1): i–xxvi, 1–508, Munich.
Part 3.2. – Proceedings of the Museum Wi Munich, Premium Edition 6 (3.2): 509–1097, Munich.
Date of publication: 2020-10-30
Published by Museum Wi Munich, Tengstr. 33, D-80796 Munich, Germany.
Proceedings of the Museum Wi Munich
– a book series prepared and published by the Museum Wi Munich and the omas-Wi-Stiung, Munich
Contact address:
Axel Hofmann, Am Hochgestade 5, D-76351 Linkenheim-Hochsteen, Germany; mail: hofmann@abl-freiburg.de
© Axel Hofmann, 2020. Alle Rechte vorbehalten.
All rights reserved. No part of this book may be reproduced or translated, in any form, by print, photoprint, microfilm, microfiche,
disc, CD-ROM or any other means without wrien permission from the authors.
Bibliografische Information der Deutschen Bibliothek
Die Deutsche Bibliothek verzeichnet diese Publikation in der Deutschen Nationalbibliografie;
detaillierte bibliografische Angaben sind im Internet über hp://dnb.ddb.de abruar.
Akademischer Verlag München · Lindberghstraße 15 · 80939 München
Fon 089/51 61 61 51 · Fax 089/51 61 61 99 · avm@druckmedien.de
Gesamtherstellung: dm druckmedien gmbh, München · Fon 089/51616100 · dm@druckmedien.de
ISBN 978-3-940732-47-7
Mu
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Part 3_Premium Edition AHrevised.indd iv 11.02.2021 12:41:26
vii
Part 3.1
Introduction
Species section
Zygaena seitzi to Z. storaiae
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viii Contents
Contents
Parts 3.1 and 3.2 1–1097
Contents and determination overview of Part 3.1 and 3.2 viii
Foreword (by Axel Hausmann), Grußwort (by Jörg-Uwe Meineke) xviii
Grußwort (by Gerhard M. Tarmann) xx
Preface xxii
Acknowledgements xxiv
e life histories and bionomics of Zygaena species 1
Introduction, general advice, abbreviations, explanations 1
Coloured pathway to the species-groups (overview) 7
e species from Zygaena seitzi to Z. storaiae; Part 3.1 8
from Zygaena ferganae to Z. trifolii; Part 3.2 522
Varia, aberrations and rarities, support for identification 997
Index of images, picture credits 1010
References 1020
Index 1068
e manlia-group 8–147
Zygaena seitzi* 10
Zygaena nocturna* 20
Zygaena kermanensis 28
Zygaena turkmenica* 34
Zygaena cacuminum 44
Zygaena speciosa 50
Zygaena cuvieri 58
*) incl. special subject
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Contents ix
Zygaena tamara 68
Zygaena manlia 80
Zygaena araxis 84
Zygaena fredi 90
Zygaena mirzayansi 98
Zygaena rubricollis 102
Zygaena hindukuschi 108
Zygaena halima 114
Zygaena wyai 116
Zygaena aisha 120
Zygaena ginnereissi 124
Zygaena haematina 128
Zygaena fusca 138
Zygaena lydia 142
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x Contents
e purpuralis-group 148–213
Zygaena brizae 150
Zygaena rubicundus 160
Zygaena cambysea 164
Zygaena erythrus 172
Zygaena minos 178
Zygaena pseudorubicundus 190
Zygaena purpuralis 196
Zygaena alpherakyi 208
e graslini-group 214–221
Zygaena graslini 216
e cynarae-group 222–233
Zygaena cynarae 224
e centaureae-group 234–255
Zygaena centaureae 236
Zygaena laeta* 242
Zygaena huguenini 250
*) incl. special subject
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Contents xi
e corsica-group 256–263
Zygaena corsica 258
e zuleima-group 264–271
Zygaena zuleima 266
e favonia-group 272–329
Zygaena loyselis 274
Zygaena favonia* 284
Zygaena aurata 300
Zygaena sarpedon 308
Zygaena contaminei 318
Zygaena punctum 322
e fausta-group 330–381
Zygaena excelsa 332
Zygaena tremewani 338
Zygaena alluaudi* 342
Zygaena algira 352
Zygaena fausta 364
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xii Contents
e hilaris-group 382–411
Zygaena youngi 384
Zygaena maroccana 392
Zygaena marcuna 400
Zygaena hilaris 406
e cocandica-group 412–477
Zygaena kavrigini 414
Zygaena truchmena* 420
Zygaena esseni 426
Zygaena transpamirina* 432
Zygaena magiana 440
Zygaena cocandica 446
Zygaena pamira* 458
Zygaena sogdiana 466
Zygaena storaiae 474
Index (complete 3.1 and 3.2) 479
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Axel F. Hofmann & W. Gerald Tremewan
THE NATURAL HISTORY
OF
BURNET MOTHS
PART III. 2
Mu
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Witt Mü
Museum Witt Munich
2020Contents xiii
Part 3.2 509–1097
Contents and determination overview of Part 3.2 516
e species from Zygaena ferganae to Z. trifolii
e olivieri-group 520–555
Zygaena ferganae 522
Zygaena chirazica 524
Zygaena naumanni 530
Zygaena tenhagenova* 534
Zygaena haberhaueri 538
Zygaena olivieri 544
Zygaena sedi 550
e fraxini-group (p.p.) 556–611
Zygaena separata 558
Zygaena rosinae 564
Zygaena bakhtiyari 570
Zygaena sengana 574
Zygaena fraxini 578
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e fraxini-group (p.p.) 556–611
Zygaena escalerai 584
Zygaena formosa 592
Zygaena peschmerga 598
Zygaena afghana 602
e felix-group 612–635
Zygaena johannae 614
Zygaena felix 620
Zygaena beatrix 628
e orana-group 636–647
Zygaena orana 638
e carniolica-group 648–691
Zygaena carniolica* 650
Zygaena occitanica* 680
e exulans-group 692–707
Zygaena exulans* 694
e viciae-group 708–729
Zygaena viciae 710
Zygaena niphona 726
*) incl. special subject
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e loti-group 730–777
Zygaena christa 732
Zygaena loti 738
Zygaena armena 760
Zygaena ecki 766
Zygaena ignifera 772
e anthyllidis-group 778–787
Zygaena anthyllidis 780
e lavandulae-group 788–803
Zygaena lavandulae 790
Zygaena theryi 800
e rhadamanthus-group 804–827
Zygaena rhadamanthus 806
Zygaena oxytropis 816
Zygaena problematica 822
e persephone-group 828–835
Zygaena persephone 830
e nevadensis-group (p.p.) 836–871
Zygaena mana 838
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e nevadensis-group (p.p.) 836–871
Zygaena nevadensis 842
Zygaena romeo 852
Zygaena osterodensis* 860
e transalpina-group 872–935
Zygaena dorycnii 874
Zygaena ephialtes* 882
Zygaena transalpina 904
Zygaena angelicae* 924
e filipendulae-group 936–995
Zygaena filipendulae 938
Zygaena lonicerae 962
Zygaena trifolii* 976
Varia 996–1095
Selected rarities and scarce aberrations 998
Sympatric occurrences of similar taxa 1007
Z. storaiae/afghana; Z. sogdiana/separata 1007
Z. filipendulae/lonicerae/trifolii 1008
Z. brizae/mana/nevadensis 1009
Z. transalpina/filipendulae 1009
Index of images (I): Landscapes 1010
Index of images (II): Host-plants 1012
Picture credits 1016
References 1020
Index (complete 3.1 and 3.2) 1068
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Figs 1, 2. Zygaena cocandica hafiza and its photographer.
By many excellent photographs and targeted fieldwork on "missing
taxa" especially for these books (!) in France, Italy, Turkey, Iran,
Armenia, Georgia, Tajikistan, Kyrgyzstan, Algeria and Morocco,
Jean-Marie André was an essential contributor to the presentation
of this monograph. We know too well that we owe him surely
more than just a hearty 'merci beaucoup' for all these efforts over
all these years! 1 (Tajikistan: Dushanbe N., Ziddy vic., 2,250 m,
16.vii.2017), 2 (Kyrgyzstan: Korgon Saj W., Sjem Ozery, 2,350 m,
7.vii.2018). Photos: J.-M. André (1), A. Hofmann (2).
Part 3_Premium Edition AHrevised.indd xvii 11.02.2021 12:42:02xviii Foreword A. Hausmann
Foreword
Burnet moths (genus Zygaena F.), the subject of the herewith published
monograph, are a truly fascinating group, a real treasure in the realm of
biodiversity. With this book, the authors offer you the pleasure of discover-
ing the complete diversity of these beautiful, sun-loving, tame and gentle
creatures. Because of their beauty, easy traceability and ecological sensibil-
ity, burnet moths can be considered one of the research ‘flagships’ of Lepi-
doptera; they are most popular among collectors, photographers and/or
monitoring lepidopterists and frequently used for environmental analyses
and conservational decisions.
In the year of 2017, Axel F. Hofmann and W. Gerald Tremewan (‘Gerry’) pub-
lished the first part of their ‘trilogy’ on the natural history of burnet moths,
which doubtlessly is the best researched and most extensively illustrated
monograph ever published on a lepidopteran group. Aer volume 1, including
a general section and introductory chapters on historical observations on the
biology of burnet moths with detailed remarks on Afrotropical and Oriental
Zygaeninae, the herewith presented volume 3 (volume 2 will follow later)
deals with the variability, distribution, preimaginal stages, biology and habi-
tats of the 108 validated species of this genus. ough being emblematic, spe-
Fig. 3. Axel Hausmann (with wife) and Gerry Tre-
cies discrimination is not always easy and species delimitation was controver-
mewan; Symposium in Antakya, Turkey, 7.v.2010. sial in some cases. is volume is a valuable, modern tool for species
identification and it summarizes, on more than 1100 pages, a huge number of
facts and data concerning all species and species groups of the ‘red zygaenids’. e quality of data is warranted by
the profound knowledge of habitats and by many hundreds of rearings (aer countless expeditions to each corner
of the Palaearctic region where burnet moths occur) paired with the outstanding scientific precision of both authors.
Like in the first volume, Axel Hofmann demonstrates his great skills to invent innovative and reader-friendly ways
of presentation. As in the table of contents (‘coloured pathway’) with exemplary photographs of each species or the
double map presentations with one map for range and type localities and another for recorded sites. Each map has
its own defined coauthorship with different symbols suggesting and ensuring correct citation of all data.
We owe deep respect and thankfulness to Axel Hofmann who did not get discouraged when his co-author passed
away in 2016, with his publisher following in 2019, and who was fighting extremely hard to realize this monograph
which again (as Part I) has to be judged THE model and THE optimum how lepidopteran monographs can best be
published. is is an example of a book that can be expected to remain the standard reference forever. Many thanks
to the authors also for assembling and publishing thousands of photographs which make it a real pleasure to use
these books. Anyone who has ever published a book, even a small one, knows how time-consuming it is. However,
for preparing monographs like these of Axel and Gerry, ‘normal scientists’ would have to invest several life times.
Since that is not possible no monographs of comparable quality and comprehensiveness exist.
A final remark of mine: Please do not miss reading the last chapter of Axel Hofmann’s preface to Vol. 3. More and
more there is a common consensus among scientists that we are in the middle of the sixth mass extinction with
100fold species extinction rates compared with the ‘normal rates’ (Ceballos & Ehrlich, 2018). Extinction of 40% of
the world’s insect species over the next few decades is predicted by Sánchez-Bayo & Wyckhuys (2019) and the se-
vere loss of insect biomass has been shown in several other studies (e.g. Hallmann et al., 2017; Lister & Garcia,
2018). Similarly, the life-long experience of the authors with Zygaenidae, which are one of the best indicators for
environmental changes, clearly points to cases of extinction and to the loss of population sizes, which highlight the
urgent need of taking the right political decisions and of acting in favour of saving habitats, the biotic and abiotic
conditions of a global nature. Professor R. L. H. Dennis, in his wonderful foreword to Volume 1, draws a parallelism
between the German word for Burnet moths, ‘Blutströpfchen’ (blood drops) and the human-induced wounds that
we encounter everywhere on our planet. Let us get inspired from each ‘blood drop’ in this book to understand that
we cannot remain passive in front of the dramatic scenario of increasing extinction rates and habitat destruction.
Axel Hausmann
Munich, 1.xi.2019
Part 3_Premium Edition AHrevised.indd xviii 11.02.2021 12:43:23Grußwort J.-U. Meineke xix
"Probably new species…."
Lieber Axel,
Deine Bie um ein Grußwort zu Teil 3 – gedanklich auch in Gerrys Namen – ehrt mich sehr, vielleicht zu sehr! Ich
versuche es also in Briefform. Zuerst mein uneingeschränkter Glückwunsch: Du hast es tatsächlich gescha, die
monographische Bearbeitung aller Zygaena-Arten abzuschließen, in nie dagewesener fachlicher Tiefe und Detail-
liertheit, auauend auf den allgemeinen Grundlagen aus Teil 1 und präsentiert in dieser Form!
In den vergangenen etwa 10 Jahren hast Du – auch mit vielältiger Unterstützung – noch vorhandene Kenntnislük-
ken mit etlichen ergänzenden Reisen in Vorder- und Mielasien, Europa und Nordafrika geschlossen, und das ne-
ben der Arbeit am Manuskript, umfangreichen Zuchten, beruflichen und familiären Aufgaben.
Die zygaenologischen Vermächtnisse von Clas Naumann und Gerry Tremewan und Dein eigener Fundus sind zu
einem universalen Dreiklang zusammengeührt. Das konnte nur glücklich gelingen, wenn man ür seine Sache
brennt wie Du.
Seit den 1980er Jahren haen wir durch die gemeinsame Mitarbeit am Grundlagenwerk „Die Schmeerlinge Ba-
den-Würembergs“ und das daraus resultierende „Artenschutzprogramm Schmeerlinge“ der hiesigen Natur-
schutzverwaltung zu enger Zusammenarbeit gefunden. Der deutsche Südwesten hat ja ür viele Schmeerlings-
vorkommen nationale Bedeutung, u.a. auch ür die Kronwicken-Widderchen Zygaena angelicae elegans und Z.
fausta. Nach Deiner ersten Reise in den Iran mit Pit Kau 1997 konnte ich Dich dann ab 1998 auf zwölf Touren in
dieses wunderbare und widersprüchliche Land begleiten. Dabei wurde das Zitat unseres mehrmaligen Fahrers
Mustafa Salehi „…probably a new species…“, wenn er mit selbst gefangenen Zygaenen ankam, zum Geflügelten
Wort. Er hae das öers von Dir oder Gerry aufgeschnappt, und nicht selten folgte auf eure Vermutung im Feld
danach die Beschreibung eines neuen Taxons.
Bei diesen prägenden gemeinsamen Reisen erlebten wir dicht ausgeüllte Tage, auch wirkliche Abenteuer und kriti-
sche Situationen. Mir brachten sie exorbitante Ausbeuten an Macrolepidopteren aus wenig oder noch nicht erkunde-
ten Regionen. Dabei mussten wir leider auch geradezu im Zeitraffer erleben, wie die Landschaen des Iran innerhalb
von 20 Jahren atemberaubend katastrophal durch Entwässerung, Austrocknung, Abholzung, Überweidung, Erschlies-
sung und Zersiedlung weiter entwertet werden. Die Menschheit haut auch hier alles auf den Kopf, was der Planet zu
bieten hat…Wir fanden in manchen Gegenden ohnehin nur noch einen Abglanz ehemaliger Zustände vor, verglichen
mit früheren Berichten. Die zahlreichen Großschutzgebiete beherbergen zum Glück aber immer noch großartige
Landschaen mit einzigartiger Natur. So fand sich vieles zwar nicht mehr an den erwarteten Fundorten, konnte aber
in abgelegeneren Gegenden gefunden werden. Dabei half Dir Deine in Jahrzehnten gewachsene Intuition so o: Aus-
ührliches Absuchen des Gebirges vom Tal aus mit dem Fernglas und dann der Entschluss, eine verdächtige grüne
Klinge oder ein Hochtal zu ersteigen. Meistens kamst Du dann erfolgreich mit neuer Beute und ausgepowert zurück,
und unser jeweiliger Guide, der um die Lizenz bangte, falls etwas passierte, war je nach Naturell vorwurfsvoll oder
nur erleichtert. Und ich kannte solche Geühle auch, wenn das Treffen um Stunden überällig war und wir mit stei-
gender Besorgnis den Berg mit dem Fernglas nach Dir absuchten bei schon tief stehender Sonne… Und passiert ist ja
auch so einiges…. So konnten wir 1998 das historische Arge-Bam bewundern. Als wir 2004 dann von Balutschestan
kommend wieder hier eintrafen, lag seit 2003 alles in Trümmern, die überlebende Einwohnerscha der Neustadt war
in Zelten der Hilfsorganisationen untergebracht, Gerry hae Tage zuvor einen schweren Sturz erlien und Du muss-
test mit einer lebensbedrohlichen photo-phytotoxischen Reaktion mit Tennisball-großen Hautgeschwüren in statio-
näre Behandlung, Folge nächtlicher Larvensuche in den Horsten eines unbekannten Doldenblütlers am Kuh-e Taān.
Die guten Erlebnisse und Erfolge wiegen dieses und anderes glücklich überstandene Ungemach auf: Die Diskussi-
onen und Gespräche bei einsamen Hochgebirgs-Lichtängen unter prächtigem Sternenhimmel über die Mechanis-
men der Artbildung und interspezifische Beeinflussungen oder die eindeutige Unmöglichkeit der Existenz der
Welt…, der erste Blick morgens aus dem Zelt in den Sonnenaufgang, ein weiterer Tag Freiheit lag wieder vor uns…
Dabei begleiteten uns auch immer wieder einheimische und europäische Kollegen und Freunde, wie natürlich
Gerry Tremewan oder Bernard Mollet, Axel Kallies, Christoph Neumann, Wolfgang Kramer oder Hossein Rajaei.
Unsere Fahrer luden uns durchweg zu ihren Familien ein. Auch die immer herzliche Aufnahme bei der Familie
Deiner Tabassom in Teheran muss ich hier festhalten.
Danke ür diese Bereicherung auch meines Lebens und daür, dass ich die Entstehung Eures unvergleichlichen
Werkes miterleben dure!
Dein Jörg
Jörg-Uwe Meineke
Kippenheim, 5.xi.2019
Part 3_Premium Edition AHrevised.indd xix 11.02.2021 12:43:52xx Grußwort
"Glückliche Umstände"
Wenn sich zum richtigen Zeitpunkt und in der richtigen Stimmung zwei en-
thusiastische Freunde dazu entschließen, ihr umfangreiches Wissen gemein-
sam zusammenzufassen, diese Idee mit Begeisterung hartnäckig und konse-
quent bis zu einem großen Manuskript entwickeln und umsetzen und das
dann auch tatsächlich in ein prachtvolles, opulentes Buchprojekt mündet,
muss man wohl von „glücklichen Umständen“ sprechen. Dabei könnten die
beiden Autoren dieses einzigartigen Buchwerkes auf den ersten Blick nicht
unterschiedlicher sein.
Walter Gerald Tremewan, der Senior-Autor, „Grandseigneur“ der Rot-Wid-
derchenforschung (Lepidoptera, Zygaenidae, Zygaeninae), von seinen
Freunden liebevoll „Gerry“ genannt, Brite aus Cornwall, distinguierter Gen-
tleman und Forscher der „alten Schule“, die Entomologie „von der Pike au“
gelernt und schon als Kind enthusiastischer Schmeerlingszüchter und –
sammler, erfuhr seinen wissenschalichen Feinschliff ganz in der Tradition
des Britischen Empires im „Mekka“ der Naturforschung am Britischen Mu-
seum ür Naturkunde in London, wo er jahrzehntelang Sammlungen bear-
beitete und kuratierte, Kataloge verfasste, umfangreiche bibliophile Werke
Fig. 4. Gerhard Tarmann (right) with Gerry
erstellte und publizierte und schließlich als Chefredakteur des renommier-
Tremewan (Italy: Passo San Marco, 14.vii.2016). ten „Bulletin“, der Fachzeitschri dieses illustren Hauses, die Entomologie
der Welt entscheidend mitbestimmte.
Axel Hofmann, der Junior-Autor, nicht weniger enthusiastischer, dynamisch-temperamentvoller Züchter und
Sammler von Kindesbeinen an, Deutscher aus Baden-Würemberg, Visionär, exzellenter Beobachter und Ge-
ländearbeiter, ‚Kind‘ der Fernweh-Generation der 1970er und 1980er Jahre mit unstillbarem Drang und Auf-
bruchsgeist zu neuen Taten, fundierter Kenner der entomologischen Sammlerwelt, entwickelte sich nicht nur zu
einem der besten Forschungsreisenden und Kenner der Rot-Widderchen, sondern schae auch noch eine be-
achtliche berufliche Karriere als Naturschutzexperte.
Beide Autoren sind verbunden durch die Liebe zur Feldarbeit, zu exakter Eigenbeobachtung, zu zeitraubenden
Zuchtbeobachtungen im Labor und zu konsequenter, exakter Beobachtungs-Dokumentation. Perfekt präparierte
Belegexemplare in umfangreichen Sammlungen zur Beweissicherung waren und sind bei beiden eine Selbstver-
ständlichkeit und untrennbarer Bestandteil ihres gemeinsamen Erfolges.
Der Verfasser dieser Zeilen schätzt sich glücklich, viele Jahre seines Lebens mit beiden Autoren eng verbunden
zu sein. Beide waren und sind Menschen, bei denen man „Energie auanken“ kann. Ein persönliches Treffen war
und ist immer ein Highlight, menschlich wie wissenschalich. Dazu gehört neben intensivem Austausch von
Forschungsergebnissen auch das gemeinsame Erzählen über interessante, o kuriose Beobachtungen und Erleb-
nisse und vor allem das Träumen, wie man das immer noch endlose bisher Unbekannte vielleicht doch noch
entdecken und enträtseln könnte. Das ührt zu einem euphorischen Zustand, in dem man die Zeit vergisst. Wie
viele gemeinsame Pläne und Projekte sind so entstanden, bei gemeinsamer Inspiration in hitzigen, begeisterten
Diskussionen und beglückendem Geplauder bei Speis und Trank!
Es gibt viele Dinge, die man von beiden Autoren lernen kann. Eines davon ist der tiefe Respekt vor den Leistun-
gen früherer Forscher und Autoren und ihren Werken. Aus den Erzählungen über Expeditionen früherer Zei-
ten, Visionen, Hartnäckigkeit und Fantasie ein angestrebtes Ziel trotz widrigster Umstände zu erreichen und
aus den Erfahrungen dieser Forscher, ihren Erfolgen aber auch aus Misserfolgen, haben beide Autoren ihre
eigene Feldforschungen geplant und durchgeührt, mit fantastischen Ergebnissen aber auch mit vielen Strapa-
zen und Rückschlägen.
Ein besonderes Charakteristikum beider Autoren war bei allen Reisen und Expeditionen stets das Ziel, aus jeder
Beobachtung maximalen Informationsinhalt zu schöpfen. Ein Falter wurde nicht einfach gesammelt, genadelt und
in die Schachtel gesteckt. Er wurde im Gelände beobachtet, wie er fliegt, was er macht, ob er saugt und woran, ob er
auf Partnersuche sein könnte oder auf der Suche nach einer geeigneten Fuerpflanze ür die Eiablage, die dann als
Fuer ür die Raupen dienen kann. War es ein Weibchen, wurde die Eiablage möglichst im Gelände beobachtet und
dokumentiert. War das nicht möglich, wurde es zur Eiablage eingesperrt und, wenn diese erfolgte versucht, die
Raupen zu züchten, damit man Informationen über alle Lebensstadien bekommt. Diese Zuchten zogen sich bei man-
chen Arten über Jahre. Herbarien der Fuerpflanzen wurden angelegt, eventuell auretende Parasiten genauso ge-
Part 3_Premium Edition AHrevised.indd xx 11.02.2021 12:43:52G. M. Tarmann xxi
wissenha dokumentiert und bestimmt, wie die Falter. Das alles waren me-
thodisch keine neuen Erfindungen, aber wie bei einem guten Essen sind es die
Details und die Erfahrung, die die alität bestimmen und das lernt man nur
durch stetes Tun, was bei Studien im Gelände von besonders hohem Wert ist.
Die Beobachtungsnotizen in den Tagebüchern muss man gleich schreiben,
noch am selben Tag. Nur dann sind sie authentisch. Beide Autoren haben das
von den großen Vorbildern übernommen und selbst nach anstrengendsten
Sammeltagen konsequent beherzigt.
Das Großartigste ist jedoch, dass beide Autoren sich zum Ziel gesetzt haben,
ihr gesamtes Wissen in einem monumentalen Werk zu vereinen. Kaum je-
mand, der nicht selbst ein umfangreiches wissenschaliches Werk verfasst
hat, kann sich vorstellen, wie lange und steinig der Weg, von der ersten Idee
so ein Werk zu schaffen, bis zum abschließenden Druckwerk ist. Um so etwas
überhaupt zu schaffen, muss man ein ganz besonderes Persönlichkeitsprofil
haben. Man braucht eine exakte Planung der Arbeit, endloses Durchhaltever-
mögen, eine immer auch selbstkritische Betrachtungsweise des Projektes und
die Bereitscha zur Kooperation mit dem Koautor und anderen Forschern.
Daneben muss man auch ähig sein, konstruktive Kritik in die eigenen Vor-
stellungen einzubeziehen und diese als Bereicherung zu erkennen. Organisa-
Fig. 5. Jörg Meineke (left) with Axel
torisches Talent ist ür so ein großes Projekt natürlich besonders wichtig. Hofmann (Iran: Shāh-Kuh, 21.vii.1999).
Das vorliegende große Opus entstand zum richtigen Zeitpunkt. Die umfangreiche Information, die ihm zugrunde
liegt, konnte nur in den letzten 50 Jahren in der benötigten Form zusammengetragen und bearbeitet werden, in
einer Zeit des Friedens und des Auruchs in ein neues, wenigstens zum Teil vereinigtes Europa. Noch bis vor 75
Jahren standen sich die Heimatländer der beiden Autoren als Feinde gegenüber und waren maßgeblich in die bei-
den schrecklichsten Kriege der Menschheitsgeschichte involviert. Dann gab es Frieden und den Wunsch, nie wie-
der Krieg zu ühren. Ein enormer wirtschalicher Aufschwung brachte Wohlstand, Sicherheit und Toleranz. Reisen
in alle Länder der Welt wurden möglich und Wissenscha konnte fast unbehindert blühen. Die Digitalisierung
schließlich brachte neue Möglichkeiten der Dokumentation, Archivierung und Präsentation. Die beiden Autoren
nutzten ihre Chancen und unternahmen ausgedehnte Forschungsreisen in die wichtigsten Länder, wo Rot-Zygae-
nen zu finden sind. Viele davon sind heute nicht mehr im früheren Sinne zu bereisen, sei es wegen Krieg (z. B.
Syrien und Afghanistan), schwieriger oder unsicherer politischer Umstände (z. B. Algerien, Libyen, Turkmenistan),
oder wegen Arbeits- und Sammelverboten ür „Ausländer“ aus kurzsichtigem Nationalismus oder falsch verstan-
denem „Naturschutz“ (z. B. Türkei, Indien). Die Welt hat sich in den letzten 50 Jahren grundlegend verändert. In-
dustrie und industrialisierte Landwirtscha, exzessive Landnutzung ür Bauten und Verkehrswege und der unre-
flektierte Einsatz von Umweltgien haben zwar wirtschalichen Aufschwung gebracht, aber zur größten
Umweltzerstörung aller Zeiten geührt. Die Widderchen, als sensible Offenlandarten und hochempfindlich gegen-
über Umweltgien, waren hier besonders betroffen. Zahllose Populationen, die früher in großer Individuendichte
gefunden wurden, sind heute verschwunden. Beide Autoren mussten persönlich zusehen, wie in ihren Heimatlän-
dern lokale Unterarten ausstarben (z. B. Zygaena purpuralis segontii Tremewan, 1958 und Z. viciae ytenensis Briggs,
1888 in England und Zygaena cynarae franconica Holik, 1936 in Deutschland).
Beide Autoren, Meister ihres Faches und der Präsentation ihrer Ergebnisse, lehren uns, was möglich ist, wenn Neu-
gier, Ausdauer, Erfahrung, Wissen, Freude und Freundscha zum Fundament eines großen Werkes geworden sind
und man im richtigen Augenblick die „Gunst der Stunde“ erkennt. Ein Werk, das in dieser Form vorher nie häe
entstehen können und das auch in Zukun nie mehr entstehen kann, ist der verdiente Lohn ür die Autoren und
eine Bereicherung nicht nur ür die Wissenscha, sondern eine fundamentale Basis ür Studien über den ständigen
Wandel unserer Umwelt und damit ein wichtiges Grundlagenwerk ür die gesamte Menschheit.
Gerhard M. Tarmann
Innsbruck, 21.x.2019
Part 3_Premium Edition AHrevised.indd xxi 11.02.2021 12:43:53xxii Preface
Fig. 6. During their last common fieldtrip the authors visited again this beautiful landscape with snowfields on Kuh-e Kameran, a herb-rich
nature with pinkish Salvia cf. hydrangea and a mixture of traditional landuse and less intensive grazing in the vicinity of Sibak, understandably
one of the most beloved biotopes of the senior author (Iran: Zāgros range, Fereydun Shahr S., 2,650 m, 8.vi.2013). Photo: A. Hofmann.
Preface to Part 3
'I have the feeling I will never have these books in my hands' was an oen spoken sentence of the senior author during
the last ten years of our work on this project; especially when the necessity of conceptional changes arose and the dim
light at the end of the tunnel grew weaker again. Originally thought as an ecological addition to the intended monograph
handbook which Clas M. Naumann had announced to write aer retirement, we could focus our research and concept
until the end of the 1990s to aspects mainly concerning the life-histories of burnet moths. 'Die Schmeerlinge Baden-
Würembergs' (G. Ebert, ed.) was the conceptional model at that time. Only photographs of living moths (preferable
taken in the wild) and the preimaginal stages should be figured and the text should mainly deal with these stages.
However, when it became more and more probable that this handbook would never appear - definitely so aer
15.ii.2004 when Clas passed away - we realised that our species chapters could not be added to an already existing
general frame. Now we had to develop a completely new concept. e title mutated from Life History to Natural
History and several new chapters had to be envisaged. But this was just one of the main reasons for the lengthen-
ing project until at last publication of Part 1 in 2017. e decision to figure the important genitalia structures of all
the species and to generate georeferenced maps and plates of set specimens, as well as personal fate and changes
to our family circumstances and the PhD of W. Gerald Tremewan (2000–2003) were other events which forced us
time and time again to extend our schedules.
Finally on 28th of April 2017 Part 1 of 'e Natural History of Burnet Moths' was published and Gerry's pessimistic
apprehension had changed to fact. Aer more than a quarter century of work on these books a few months were
missing to let him have at least the satisfaction of holding the first part in his hands. On 1 October 2016 the senior
author, W. G. Tremewan, passed away (Hofmann, A. 2017; Hofmann A., Spalding, A. Tarmann G. & Wi T., (eds.),
2017; Spalding A., 2017; Morton, G., 2017; Carter, D., 2017; Young, M., 2017; Saler, K., & Vane-Wright, R. I., 2017).
Our common friend Adrian Spalding immediately and very kindly offered help in this project-critical moment.
Without hesitation he took over the responsibility for the final editing of the english text of the forthcoming parts.
Of course, there can be no doubt that this cannot be an equivalent substitute, as Gerry did not only master the
language but also the subject.
Part 3_Premium Edition AHrevised.indd xxii 11.02.2021 12:43:55A. Hofmann xxiii
e basic text for Part 3 was wrien between 2005 and 2013 but many data, records of current fieldwork, new
publications and personal observations of colleagues had to be incorporated. e 200 plates figuring 5841 set specimens
and the transcription of labels took up a considerable amount of time, as did the gathering of geo-information, the
endless feeding of a database and the generating of maps, pictures and captions. e greatest problem now, however,
was missing feedback from and discussion with Gerry, and the improvement by common ideas.
On 28.i.2019 our publisher Professor Dr h.c. omas J. Wi passed away quite unexpectedly (Schwarz & Gusenleit-
ner 2019; Hausmann 2019, 2020), a tragedy not only for these books but for the science of lepidoptera diversity and
especially for palaearctic Bombycoidea. Nevertheless, it was clear that our project will go on. is was not a question
of duty, pledged word and promises; it was and still is the vested interest of the junior author to bring this monograph
to an end. Moreover, an encouragement to go on came from the numerous benevolent reviews (vide p. 1020) and many
positive feedbacks of colleagues on Part 1.
e now presented Part 3, the species section, grew to a voluminous amount of pages. Instead of 400–500 pages, as
originally intended, it expanded to more than twice the size. Weight and binding made a division into two books,
3.1 and 3.2, necessary. However, both books form one unit and consequently exhibit continuous pagination and
one index (at the end of each book), while the bibliography is placed at the end of Part 3.2. anks to Verena Wi,
eldest daughter of omas Wi, the family of omas Wi and the omas-Wi- Foundation and its supporters
(vide page 1097) it was possible to keep the price for these two books at the same level as for Part 1!
A set of three typical specimens as an overview for determining the species and for quickly finding the right pages
for all 108 species is provided twice, i.e. at the beginning of Part 3.1 (pp. viii–xvi), and at the beginning of Part 3.2.
e taxonomic division of higher categories (species-groups, subgenera) as dealt within these books is listed on
double-page 6 and 7 and optical marks on the le and right side of each page lead the reader to the species-groups
with blue colours for Mesembrynus, green to orange for Agrumenia and orange to red for Zygaena.
Now that Part 1 and 3 are published the work will go on with Part 2 which will comprise 14 more chapters (vide 1/5,
6) dealing with hibernating strategies and reproductive biology, phenology and voltinism, dispersal and colonization,
aposematism and mimicry, parasitoids and Mendelian genetics, rearing burnets in captivity and - last not least - conser-
vation and national responsibilities. Two more years are calculated, thus making a date of publication in 2023 realistic.
Aer completion of these two books I finally found the reason for such a lifelong deal with this subject. Now I
definitively know that it was not the unquestionable beauty of the moths, signalling their toxicity with their unfor-
geable colours, or the wonderful co-operation with unforgeable friends. It was (and still is) the beauty of their
habitats. When I regard the herein presented pictures of biotopes I feel great yearning for these places of silence
with the smell of Artemisia, thyme and other herbs, to the spring flowering Mediterranean maquis and the scabious-
rich landscapes in Anatolia, to the Stipa-steppe of North Africa, the light clearings in Transcaucasian forests and the
cushion zone in the vicinity of snow-fields in high mountains of Iran and Afghanistan. During 50 years of fieldwork
everything has changed but nothing for the beer. Many of my habitats are destroyed, sometimes not even retriev-
able anymore. On most pages I cannot hide a feeling of nostalgia, or even more, a melange of melancholy and sad-
ness. e many pictures in these books may let the reader participate in my naive longings for retrotopia.
e substantial decline in insect populations, including burnet moths, is an indisputable fact. All our personal
observations, taken together with information and data collected by authorities over time, unambiguously con-
firm what we have become aware of more and more during recent decades. e abundances of populations have
decreased, in some areas they have dramatically collapsed; many habitats have changed or have even become
completely destroyed and with them the populations1. Now that this subject is being more openly discussed, we
have become increasingly aware of these trends. However, except for limited small scale efforts, we cannot see any
current serious global action being taken that encourages us to feel optimistic about prospects for the future. If we
were to begin working on these books today, we would be unable to bring them to completion in the same way.
Too much has been lost already, irretrievably lost, not only in Europe but also in North Africa, the Middle East and
Central Asia, wonderful areas with a wealth of natural beauty, which we have visited so oen. What we urgently
need now are political decisions, firm actions, based on a broad social consensus, and a belief that time has not
already run out to support the biodiversity that remains.
Axel F. Hofmann
1
In spite of intensive targeted search there have been Hochsteen, 8.i.2020
no records for Zygaena ferganae (last record 1940), Z.
problematica (xxiv Acknowledgements
Acknowledgements
is monograph has two authors but is the product of dozens of selfless, generous supporters, of providers
for data, pictures (vide p. 1016-1019) and soware assistance. We feel ashamed to be unable to distribute
the merits in an adequate way to these friends and colleagues. However, wherever it was appropriate in
the text and in the captions their names are cited, the least - as a minimum - that we could do.
Martin Albrecht (Bern), Sergei Andreev (Moscow), Dr Jean-Marie André (Caumont sur Durance),
Professor Dr Zeki Aytac (Ankara), Daniel Bartsch (Stugart), Dr Dubi Benjamini (Tel Aviv), Dr Christian Berg
(Graz), Stefan Birrer (Muenz), Rolf Bläsius (Eppelheim), Daniel Bolt (Domat/Ems), Marco Bonifacino (Vado
Ligure), Anne Bounias-Delacour (Cavaillon), Ulf Buchsbaum (München), Professor Dr Günter Buntebarth
(Clausthla-Zellerfeld), Constantin della Bruna (Roma), Berthold Clewing (München), J. F. Cardin (France),
Alexander Chuvilin (Tula), Dr Jerzy Dąbrowski (Kraków), Dr Alexander Danchenko (Moscow), Yann
Delmas (Robion), Mr Jaques Demange (Denis du Val), Professor Dr Dr Roger L. H. Dennis (Wilmslow), Éric
Drouet (Gap), Günter Ebert (Stutensee), Dr Wolfgang Eckweiler (Frankfurt), Professor Dr Dr Konstantin
Efetov (Simferopol), Dr Ulf Eitschberger (Marktleuthen), Rainer Ernst (Linkenheim-Hochsteen), Dr
Marianne Espeland (Bonn), Herrmann Falkenhahn (Ebersdorfergrund), Alessandro Floriani (Milano, Italy),
Josef J. de Freina (München), Michael Falkenberg (Karlsruhe), Evyator Feingold (Tel Aviv), Dr Riccardo
Galbiati (Pavia), Enrico Gallo (Genova, Italy), Professor Dr Oleg Gorbunov (Moscow), Professor Dr omas
Goschalk (Roenburg), Raymond Guenin (Wabern), the late Kenneth Guichard, Jean-Alain Guilloton
(Héric), Francisco Lencina Gutierrez (Jumilla), Helmut Haas (Aalen), Erik Haase (Luckenwald), Stefan
Hafner (Löffingen), Dr Ingmar Harry (Freiburg), Dr Axel Hausmann (München Foreword), Jeremy D.
Holloway (London), Dr Kiyoshi Horie (Tokio), Dr Herbert Huss (Stadl-Paura), Professor Dr Predrag Jakšić
(Niš), Dr Axel Kallies (Canberra), Oliver Karbiener (Freiburg), Pit Kau (Tübingen), omas Keil (Dresden),
omas Kissling (Zürich), Professor Dr Chiharu Koshio (Naruto), Dr Wolfgang Kramer (Freiburg), Jirí Klír
(Litomerice), Martin Kratochwil (Stugart), Albert Krebs (pictures via ETH Zürich), Igor Kostyuk (Kijev),
Lars-Inge Larsson (Stockholm), Professor Georgy Lazkov (Bishkek), Dr Hans-Georg Mark (Königstein),
Nora Magg (Freiburg), Professor Dr Gregor Markl (Tübingen), Dooressa Melania Massaro (Roma), Roman
Maximovin (Abakan), Marlies Mayr (Feldkirch), Dr Jörg-Uwe Meineke (Kippenheim), Paulin Mercier (Niort),
Alain Migeon (France), Dr Dimitry Milko (Bishkek), Bernard Mollet (Gif-sur-Yvee), Professor Dr Günther
Müller (Bamako), Alireza Naderi (Tehrān), Ana Nahirnić (Sofia), Alisher Nazarov (Dushanbe) Alexander
Naumann (Hamburg), the late Professor Dr Clas M. Naumann, Dr Elsa Nickel (Bonn), Marc Nicolle
(Angers), Klaus Nimmerfroh (Stugart), Dr Amirhossein Pahlevani (Tehrān), Dr Tom Presco (Kingussie),
Ralf Prosi (Aalen), Jean Raillot (Gap), Dr Hossein Rajaei (Stugart), Professor Dr Laszlo Rakosy (Cluj),
Dr Massoud Ranjbar (Hamadān), the late Dr Günter Reiss, Emmanuel Ruiz (Valensole), Evgeniy Rutjan
(Kijev), Professor Dr Nils Ryrholm (Gavle), the late Agustí Moliné Sala (1953–2018), José Luis Santa López
(Jumilla), Professor Dr Valerio Sbordoni (Roma), Professor Dr omas Schmi (Müncheberg), Walter
Schön (Kirchheim/Teck), the late Hans Seipel, Dr Mark R. Shaw (Edinburgh), Dr Adrian Spalding (Praze-an
Beeble), Hildegard Stalder (Hasliberg), Axel Steiner (Wöschbach), Dr Dieter Stüning (Bonn), Harald Sulak
(Weiden), Dr Wolfgang ten Hagen (Mömlingen), Valentin V. Tikhonov (Pjatigorsk), Dr Robert Trusch
(Karlsruhe), Vadim Tshikolovets (Kijev), Professor Dr Zoltan Varga (Debrecin), Professor Dr Richard Vane-
Wright (Canterbury), Pavel Vitek (Znojmo), Dr Wolfgang Wagner (Stugart), Jean-Claude Weiss (Metz),
Felicitas Werwie (Freiburg), Professor Dr Zoya A. Yefremova (Ulyanovsk), Professor Dr Mark Young
(Oldmeldrum), Dr Mika Zagrobelny (Copenhagen), Dr Heiner Ziegler (Chur), Dr Alberto Zilli (London).
None of these colleagues made his contribution dependent of any return services. id pro quo was never
asked and would be impossible for us to serve. All these above mentioned colleagues are thanked for both
their contributions and for the wonderful experience of how cooperation in this world can run!
Part 3_Premium Edition AHrevised.indd xxiv 11.02.2021 12:43:59Acknowledgements xxv
Special thanks go to Dr Axel Hausmann Dr Jean-Marie André (Iran 2016, Morocco 2017,
(Munich) for his inspiring compliments and for Kyrgyzstan 2018, Algeria 2018, 2019), Shafiq Asad
providing the foreword and to Professor Dr Gerhard (Afghanistan 2007), Claus Becker (France 1971,
Tarmann (Innsbruck) and to Dr Jörg-Uwe Meineke 1972), Ulli Bense (Swabian Alb 1994, 1998, 2000),
(Kippenheim), our forbearing companions on many Rolf Bläsius (Hockenheim 1994), the late Philipp
tours. Moreover, both did not hesitate a second to Baenfeld (Morocco 1979), Franz Büschel (France
contribute with two personal memories wrien 1972, Italy 1972, 1977), Erich Beag (Hockenheim
down as a welcoming address aer the foreword. 1994), Dr Wolfgang Eckweiler (Tunisia-Libya 1981,
Very prompt and very colleagual was the South Africa 2016), Dieter Fritsch (Istein 1996),
support by Dr Wolfgang Wagner and Professor Dr Werner Fritsche (Kaiserstuhl 1996), Ernst Görgner
Gregor Markl. Both were essential to the success (Turkey 1981), Ulrike Haas (France 1973–1978),
of the photo documentation. Moreover, Gregor Stefan Hafner (Baden-Würemberg 1994–2012),
very carefully read a complete dra and provided René Herrmann (Baden-Würemberg 1993–1997,
substantial second editorship. France 1996–2000, Morocco 1994), Johanna Groß-
Hofmann (Turkey 1981, 1982), Hans-Dieter Herbst
We must not forget the sympathetic reviewers (Morocco 1978, Turkey 1981, 1982), Rüdiger
of our first volume (vide p. 1020). eir feedback Hofmann (Morocco 1977, 1978, 2008, 2015, Spain,
gave an important positive push to continue. 1977, 1978, Turkey 1981, 1982), Dr Axel Kallies (Iran
Many thanks to Professor Dr Lawrence Cook, 2002), Oliver Karbiener (Baden-Würemberg 2001,
Sylvain Delmas & Éric Drouet, Professor Dr Fidel 2012), the late Uli Kra (Swabian Alb 2007, 2008),
Fernandez-Rubio, Professor Dr Hartmut Greven, Peter Kau (Iran 1998, Swabian Alb 1992–1995),
Raymond Guenin, Dr Jörg-Uwe Meineke, Dr the late Franz Kirsch (Tauberland 1994–1996), Rudi
Wolfram Mey, Willy O. De Prins, Erwin Rennwald, Lang (France 1971–1975, Italy 1972, Morocco 1978,
Professor Dr omas Schmi, omas Sobczyk, 2008), Birgit Lehmann (Spain 1979, France 1980,
Professor Dr Gerhard M. Tarmann, Professor Dr Tunisia 1980), the late Erich Loser (Swabian Alb
Zoltan Varga, Professor Dr Dick Vane-Wright and 1992), Nora Magg (Swabian Alb 2017), Bernard
Dr Wolfgang Wagner. Mollet (Iran 1998, 1999, 2005), Dr Jörg-Uwe
A final reverent 'thanks for all' goes to Professor Meineke (Baden-Würemberg 1994–2012, France
Dr h.c. omas J. Wi (* 2.9.1947, † 28.1.2019). 1996, 2000, Iran 1998–2010), Sergio Mosca (Spain
Henceforward his philosophy, his friendship and 1996), Alireza Naderi (Iran 2006–2016), the late
always friendly manner and generosity will be Professor Dr Clas M. Naumann (Swabian Alb 1995,
sadly missed (Hausmann, 2019, 2020). e efforts 2002), Dr Hossein Rajayi (Iran 2004), the late Dr
of his daughters, Dr Alessa and Dr Verena Wi, Günter Reiss (Swabian Alb 1991–1998, Morocco
to save omas' Life Work deserve our greatest 1980, 1985, 1988, 1994, Tunisia 1981, 1982, Algeria
respect. 1982, 1984, Turkey 1986), Stefan Reiss (Morocco
1989, 1994, Algeria 1989, Turkey 1986), Roland
Dr Storai Naumann-Nawabi is thanked for Seitz (Morocco 1978, 2008), Professor Dr Klaus
continuous support, for her friendship and always Rose (Morocco 1979), Peter Strauss (Tunisia 1979,
positive vibrations. Turkey 1981 1982, Spain 1983, Morocco 1983), Dr
Both authors owe special thanks to their Karel Špatenka (Uzbekistan 1995), Michel Tarrier
partners and families for many tours during all of (Morocco 2017), Dr Robert Trusch, the late Dr
which Zygaenology was always in the main focus. Michael Witschel (Swabian Alb 1996, 1997) and Dr
Morover, the junior author would like to thank in Sergej Zonstein (Uzbekistan 1995).
particular his many companions on entomological As is known, 'Yes' is quickly said, but then?
tours, for their understanding and patience, for Dear Adrian Spalding, you very carefully did a
their help whilst on fieldwork and motivation to wonderful job over the past three years, aer we
go on. Every single tour remains unforgoen and had asked you to take over the editorship. I am sure
memories arise each time when reading the labels you could not guess what a longlasting word you
of set specimens in the drawers: gave then. anks for all!
Part 3_Premium Edition AHrevised.indd xxv 11.02.2021 12:43:59xxvi Introduction Part 3_Premium Edition AHrevised.indd xxvi 11.02.2021 12:43:59
General advices 1
e life histories and bionomics of
Zygaena species
Introduction and general advices Unless if mentioned otherwise all figured specimens
is part deals with the life histories and bionomics are stored in the collection of the junior author (AH).
of all currently recognized Zygaena species (check- In the legends to the plates with triplets of set speci-
list vide 1/52–57), when these are known. For all mens valid subspecies are cited in bold, framed by
information we have tried to provide an exact refer- a cartouche with cast shadow. When specimens of
ence, while accepting that this might interrupt the synonymic taxa or of unclear status (valid or not ?)
flow of the discursive text and that we risk receiving or of populations are figured, then the nominal taxon
criticism from the reader for such inconvenience. is cited as originally described. at does not neces-
In addition to so much data based on our own field sarily mean that we agree with this position or even
observations and cultures, authentic information rectify/revise them here, but want give the reader a
received from colleagues has been included as well chance of forming his own opinion about such avail-
as every important reference that we could find in able, subjective taxa. Only very obvious invalid taxa
the published literature. are placed in synonymy (distinguishable by citation
e main goal of this part is to describe, demon- with 'syn.' and in normal italic instead of bold italic).
strate and where possible to analyse infraspecific Unnamed but slightly to mediumly distinct popula-
diversity in a wide sense, not only what is mani- tions are occasionally highlighted as 'subsp. ?'; in no
fested in the morphological structures of adults way does this imply that nomenclatoric naming is
and pre-imaginal stages, but also in the spectrum urgently needed, but shows that even distinct popu-
of their specific and infraspecific habitats and eco- lations can remain unnamed.
logical adaptations.
e selection on the plates allows the reader to get
e plates of infraspecific differentiation a quick overview about distinct (or not distinct)
characters and geographical distances can be seen on
Each species chapter starts with a double page, the the distribution map. In this monograph taxonomy
geographical differentiation (preferably the valid has taken a back seat and only some relevant, recent
subspecies) figured by sets of three (or four) typical taxonomic changes are discussed.
specimens in one line on the le page, so that the
reader of the text on the right page can easily see Data cited here (e.g. altitude) are original citations
what the described infraspecific characters figured from the specimen pin-labels. When citing taxa in
on the le page. Oen the phenotypic spectrum of the legends of the pictures of landscapes and bio-
a subspecies is wider than can be shown with a few topes the authors and dates of publication are omit-
specimens. We have reduced the selection to 'popu- ted; this information is provided in the running text
lation typical specimens' accepting that aberrations - beside.
even regular aberrations - are oen not represented.
e species introductions
Additional information on subspecific descriptions
and further figures of specimens of western Palaearc- e text starts under the species heading with the
tic Zygaena species are to be found in de Freina & reference to the original description of the taxon,
Wi (2002). For an extented phenotypic comparison as cited by the author but rationalised to a certain
we recommend the virtual collection of the Museum extent. is is followed by a short introductory sec-
Wi Munich (hp://www.insecta-web.org/MWM/). tion providing brief biographical details of the author
Part 3_Premium Edition AHrevised.indd 27 11.02.2021 12:44:022 Introduction
or authors of the nominal taxon and the field workers targeted data from missing regions (especially United
who originally discovered the material, together with Kingdom) from the collection W. G. Tremewan and
the derivation of the name when either is known. e selected data from the Museum Wi (München),
geographical range, the vertical distribution and the Zoologische Staatssammlung (München), e
most significant variation within subareals of each Natural History Museum (London), Musée d'Histoire
species are briefly described and all noteworthy Naturelle (Paris), Museum Alexander Koenig (Bonn)
subspecies are characterised and figured. Here each and Staatliches Museum ür Naturkunde (Karlsruhe).
nominal taxon (normally subspecies) is fully cited ese collections contain the famous collections of
and in bold when first mentioned, in the subsequent F. Daniel, F. Dujardin, K.-H. Wiegel, H. & G. Reiss
text and in the captions the authors(s) and the date of (all in Munich) and C. M. Naumann (in Bonn). More-
publication are omied. e subspecific taxonomy is over, an essential extension was the cooperation with
following Hofmann & Tremewan (1996) where fur- Laszlo Rakosy (Cluj, Romania), Igor Kostyuk (Kiev,
ther synonyma can be found. We have abstained from Ukraine) and Tiroler Landesmuseum Ferdinandeum
comprehensive descriptions and detailed discussions (Innsbruck). G. M. Tarmann has gathered data on
of subspecific characters. Only the most remarkable Palaearctic Zygaenindae since 2005 with main focus
characters in note form are mentioned. Otherwise it on the Apennnine Peninsula, the Alps and the Bal-
would lead to endless paragraphs with subsequent kans (s.l.). ese data of which the full sets are stored
debates about subspecific criteria, subjective deci- in the biooffice (Innsbruck) perfectly completed the
sions and transition zones. All of which is not the under-represented sets of eastern and south-eastern
goal of this monograph, as these but can be found in Europe. A very essential contribution for occurrences
standard works (Reiss in Seitz, 1930-1933; Naumann, in France was provided by the 'Groupe d'Information
Tarmann & Tremewan, 1999; de Freina & Wi, 2002). de Recherche et d'Animation sur les Zygaenidae'
(GIRAZ). Nevertheless a regionally more or less equal
For every species a selection of pictures of character-
presentation of distribution dots was not possible as
istic landscapes and habitats are provided to give the
the availablity of easily incorporated data sets was
reader the opportunity to imagine the species spe-
not given for many areas. Although well investigated
cific ecological plasticity. Here our aim and working
and with a large amount of data being stored in the
question was "Could one gets the specie's name just
collections for the Iberian Peninsula, central, north-
from seeing these pictures of its natural habitats?"
ern and eastern Europe, continuous digital data were
lacking and we contented ourselves with the incor-
e distribution maps
poration of the most important cornerstone data sets,
e specific distribution with subspecific type-local- being aware that much more data was available in
ities of each species are shown on simple maps with the collections. In contrast, for the more peripheral
few colours while exact sites are figured on separate regions (e.g. North Africa, Iran, Afghanistan), which
maps with distinct coloured altitudinal zones. Based were much less well explored, the majority of collec-
on georeferenced maps from U.S. Geological Supply tions and recorded data could here be incorporated.
we succeeded in transforming these data into maps
Every dot on the map is based on at least one veri-
of altitudinal zones for the whole palaearctic region.
fied set of data. e red coloured, framed areas
On such maps not only the distribution is exactly
represent ranges of potential occurrences, i.e. within
shown but also information on ecological limitations
these areas further records are known or to be
are provided.
expected and in no way do such areas indicate
For every species a distribution map is generated that the species is omnipresent here. When dots
which is based on authentic records that could be remain outside the specific range in red it means
georeferenced and incorporated in ESRI ArcView and that such records are either based on what are now
QGis projects (the laer an open source soware). most probably extinct populations (last records
ese maps in their present completeness could not more than 50 years ago), or they are unclear or
be worked out only based on personal data. ey doubtful and therefore require confirmation. For
are the product of intensive cooperation. Originally some species with very spare records (e. g. Z. sedi,
based on literature analysis (vide References, p. 1022– Z. nevadensis) or for some subareals with very few
1039) and on a complete data gathering of the collec- records, we understandably refrained from using
tion of A. Hofmann it was subsequently extended by framed ranges.
Part 3_Premium Edition AHrevised.indd 28 11.02.2021 12:44:03Distribution maps 3
Map basis Maps in co-authorship
e distribution maps in this volume were genera- Z. brizae (AFH, GMT, ED, IK)
Z. rubicundus (AFH, GMT)
ted using four different sources of information: Z. erythrus (AFH, GMT, ED)
• e observations and voucher material of the Z. minos (AFH, GMT, ED, AN)
authors. Z. purpuralis (AFH, GMT, ED, AN)
• Voucher material in other collections, both Z. cynarae (AFH, GMT, ED, IK)
private and public. Z. laeta (AFH, GMT, IK)
• Databases, both regional and national. Z. sarpedon (AFH, GMT, ED)
Z. punctum (AFH, GMT, AN, IK)
• Literature data (vide pp. 1022–1039). Z. fausta (AFH, GMT, ED)
Z. hilaris (AFH, GMT, ED)
Z. carniolica (AFH, GMT, ED, IK)
Twenty-eight maps were produced in close cooper- Z. occitanica (AFH, GMT, ED)
ation with colleagues, with different symbols (vide Z. exulans (AFH, GMT, ED)
legend below) used to allow reference to the origin Z. viciae (AFH, GMT, ED, IK)
Z. loti (AFH, GMT, ED, IK)
of the dots. When just citing or referring to these Z. anthyllidis (AFH, GMT, ED)
maps - oen including unpublished data - of the fol- Z. lavandulae (AFH, GMT, ED)
lowing species we therefore propose the citation as Z. rhadamanthus (AFH, GMT, ED)
follows: Z. nevadensis (AFH, GMT, ED)
Z. romeo (AFH, GMT, ED)
[insert authors] (2020) in Hofmann, A. F. & W. G. Z. osterodensis (AFH, GMT, ED, IK)
Tremewan. e Natural History of Burnet Moths, Z. ephialtes (AFH, GMT, ED, IK)
Part 3, [insert page]. Z. transalpina (AFH, GMT, ED)
Z. angelicae (AFH, GMT, IK)
(Abbreviations of authors: AFH, Axel F. Hofmann; Z. filipendulae (AFH, GMT, ED, IK)
GMT, Gerhard M. Tarmann; ED, Éric Drouet; AN, Z. lonicerae (AFH, GMT, ED, IK)
Ana Nahirnić; IK, Igor Kostyuk). Z. trifolii (AFH, GMT, ED).
Legend
Altitudinal levels Georeferenced sites
(as used in the distribution maps)
Data from coll. A. Hofmann [respectively data from
museums verified by A. Hofmann]
< sea level
Data from external databanks, viz. G. Tarmann (Italy
0 – 100 to the Balkans), (small dots)
101 – 200 Data (mainly France) from databank of GIRAZ via
201 – 500 É. Drouet, (grey dots)
501 – 750 Data from literature (white dots)
751 – 1000 (Bibliography p. 1022–1039)
1001 – 1300
1301 – 1600
Range of genus Zygaena F. (dark grey)
1601 – 1900
1901 – 2250 Range of species (red)
2251 – 2500
2501 – 3000 ① Type-locality [black dot with
# of taxon]
3001 – 3500
3501 – 4000 ①②③④.. Numbers (black) refer to valid taxa and type-
4001 – 5000 localities in maps, plates and text.
①②③④… Small numbers (white) refer to type-localities of
> 5000 synonymic taxa in maps; grey in text of Plate-Figs.
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