Table Of ContentSE RIES
ENTOMOLOGICA
EDITOR
E. SCHIMITSCHEK
Wien
VOLUMEN 2
Springer-Science+Business Media, B. V. 1967
A
SYSTEMATIC CATALOGUE
OFTHE
GENUS ZYGAENA FABRICIUS
(LEPIDOPTERA: ZYGAENIDAE)
by
HUGO REISS
and
W. GERALD TREMEWAN
Springer-Science+Business Media, B.V. 1967
ISBN 978-94-017-5857-4 ISBN 978-94-017-6333-2 (eBook)
DOI 10.1007/978-94-017-6333-2
Copyright Springer Science+Business Media Dordrecht 1967
Urspriinglich erschienen bei Dr W. Junk Publisher, The Hague 1967.
Softcover reprint of the hardcover I st edition 1967
INTRODUCTION
It is forty years since Burgeff published, in 1926, the first comprehensive catalogue
of the genus Zygaena Fabricius, forming part 33 of the Lepidopterorum Catalogus.
Following the pattern and general Iayout of Burgeff's work, we have attempted
to produce a catalogue in which all names in the genus Zygaena are included,
with references to the Iiterature where these names were originally published.
Additional references are included when these refer to illustrations of a species,
subspecies or form, or to a taxonomic change, e.g., a change in status. References
to misidentifications are generally omitted unless a new species has been described
at a later date.
In compiling this work we have adhered to the International Code of Zoological
Nomenclature as adopted by the XV International Congress of Zoology. However,
although the International Commission on Zoological Nomenclature recognises
the necessity of names of lower rank than subspecies, they do not at present deal
with such names. The provisions of the Code do not apply to them and, therefore,
such names have no nomenclatural status.
Every subspecies is given equal status in the catalogue although their relative
value is not always the same. Certain authors have very often separated sub
species on minute differences and a subsequent examination of further material,
taken over a number of years, has shown that the differences are not always
constant. In many cases, however, we have been unable to verify the status of each
subspecies.
The names of forms and aberrations are included in the catalogue and are,
in our opinion, a necessity to geneticists and collectors. Long descriptions are
thereby avoided by the use of such names. This also applies to the names of
hybrids for the same reasons.
The hybrids which were described by Stauder from specimens captured in
the field are listed in the catalogue as aberrations or synonyms, since an examina
tion of many of his types has shown that the genitalia are identical with those
of normal specimens. It has been shown that the genitalia of hybrids, bred in
captivity from a cross-pairing between two different species, exhibit intermediate
characters.
It has often been customary to employ the same name when describing similar
aberrations occurring in two or more species of the genus Zygaena. For example,
the name flava has been applied by many authors to yellow aberrations occurring
in different species. This seems to be a more suitable method than employing
many different names when describing aberrations analogous for colour. The
name flava immediately suggests that the aberration is yellow. There should be
no confusion if the name is cited with its respective specific and subspecific
names. If the Law of Homonymy were applied to the names of aberrations, a
!arge percentage would immediately become junior primary homonyms. The
species of the genus Zygaena are extremely variable and many hundreds of names
of aberrations can be found in the existing literature. When describing an aber
ration, some authors have used a name that has already been employed for a
similar aberration occurring in the same species. For example, Rühl (1896, Ent.
VI
Z., 12: 117) applied the name grossmanni to a yellow aberration of Z. purpuralis
fatrensis Reiss. Later, Burgeff (1914, Mitt. münch. ent. Ges., 5: 43) employed
the same name for the yellow aberration occurring in Z. purpuralis nubigena
Lederer, as follows: "purpuralis nubigena ab. grossmanni Rühl (n. em.)". This
was quoted in later publications as ab. grossmanni (Rühl) Burgeff, 1914. This
method was originally introduced by Burgeff (loc. cit., 5: 37) who employed the
term "n. em." ( = nomen emendatum). This is not truly an emendation, which is
either the correction of an incorrect spelling of a name or an incorrect subsequent
spelling of a name. We do not condone Burgeff's method of naming aberrations
and, in the catalogue, those described in this manner are included and date from
the publication of the second author to whom the name is also attributed. Such
aberrations are considered separate entities and the descriptions are regarded as
new. This should not be confused with misidentifications.
With the exception of Z. ephialtes Linne, little is known of the genetics of the
various forms and aberrations of the Zygaena species. Dryja (1959, Badania
nad Polimorfizmem Krasnika Zmiennego) has made an extensive study of ephialtes,
his work being the result of thirty-two years of breeding experiments. It has been
shown by Dryja that the two basic forms (peucedanoid and ephialtoid) each
depends on a pair of independently inherited allelomorphic genes and that peuce
danoid forms are dominant to ephialtoid forms. The peucedanoid and ephialtoid
forms can be either red or yellow, the former colour being dominant to the latter.
Each of these colours is dependent on a second pair of independently inherited
alleles. These characters showed, in the F2, F3 and back-crosses, typical Mendelian
segregation. The peucedanoid and ephialtoid forms can be five- or six-spotted.
It was found that five-spotted forms are dominant to six-spotted forms and that
each depends on a third pair of independently inherited alleles. The same results
have been obtained by Burgeff and Bovey and the references to their publications
may be found in this catalogue.
Some species of the genus Zygaena are closely allied and the genitalia do not
exhibit good characters. The concept of a species varies according to the author.
lf cross-breeding between closely related species were undertaken, it would show
whether the resulting hybrids (F1) could interbreed freely and produce a further
generation (F 2). If breeding experiments produced such results it would suggest
that the separation of these species is not justified. The closely allied species
trifo/ii Esper and lonicerae Scheven have almost identical genitalia and the minute
genital differences are not always constant. When these two species are crossed
the resulting hybrids can interbreed quite freely and produce fertile progeny.
However, the larvae of the two species are different in coloration and setal structure
and, in our opinion, such differences are of considerable taxonomic value. Closely
allied species, such as trifolii and /onicerae, have a wide geographical distribution
which overlaps. A large nurober of weil defined subspecies have evolved but each
of these subspecies has, however, the basic characters of either trifolii or /onicerae.
The grouping of such species into one species would, in our opinion, be quite
unjustifiable.
Although twenty-two subgenera have been erected within the genus Zygaena,
the species are grouped in the catalogue under three subgenera only, viz., Mesem-
VII
brynus Hübner, Agrumenia Hübner and Zygaena Fabricius. These three subgenera
are divided into sections or species groups, many of which were formerly placed
as subgenera. The remaining subgeneric names are now placed in synonymy.
This arrangement follows, in general, the classification of the senior author
(Reiss, 1958, Z. wien. ent. Ges., 43: 140 et seq.) who based his conclusions on
the morphology, extemal characters, such as wing pattem and coloration, and the
biology of the species. The species in subgenus Mesembrynus are considered to
be the most primitive. This subgenus is followed by the subgenus Agrumenia which
in turn is followed by subgenus Zygaena, the latter containing what are considered
to be the most recently evolved species. The coloration and spot-formation of
the forewings give support to this theory. In what are considered to be the most
primitive species of Mesembrynus the forewing spots are unicoloraus cream,
yellow, orange, light red or carmine. Most of the species in Agrumenia have the
forewing spots encircled with whitish cream, cream or light yellow coloration.
It is thought that the red coloration in the forewing spots in Agrumenia began
from the centre in the more primitive species. The yellowish or whitish ring then
appeared as the remainder of the former colour of the forewing spots which,
as already stated above, are in the more primitive species of Mesembrynus, yellow,
orange, light red or carmine. The closely related species Z. banghaasi Burgeff
and cocandica Erschoff are good examples. Only in the more recently evolved
species of Agrumenia, e.g., exulans Reiner & Hohenwarth and loti Denis &
Schiffermüller, are the rings normally absent around the forewing spots. When
present, the rings are found only in aberrant specimens. It follows that the most
recently evolved species grouped in Zygaena have lost the cream or whitish rings
surrounding the forewing spots. It is interesting to note that occasionally, spe
cimens of filipendulae Linne and transalpina Esper have been captured with
traces of whitish rings around the forewing spots. Such forms occur only as
extremely rare aberrations in Italy and suggest a possible link with the more
primitive Agrumenia species.
A red collar and abdominal ring are present in most species of the subgenera
Mesembrynus and Agrumenia. The abdominal ring also occurs in the moreprimitive
species of subgenus Zygaena. In the more recently evolved species in Zygaena,
however, the abdominal ring occurs only in aberrant specimens and has not
been detected at all in ramburii Herrich-Schäffer, trifo!ii and lonicerae, which
are considered to be the most recently evolved species of the whole genus Zygaena.
The larvae of the species in Mesembrynus feed on plants of the families Um
belliferae, Compositae and Labiatae. The larvae of the species in Agrumenia feed
mostly on hard-leaved Papilionaceae while the larvae of the species in Zygaena
feed mostly on soft-leaved Papilionaceae. A study of the larvae, pupae and cocoons,
as far as they are known, has provided valuable characters for placing the species
into species-groups or sections.
The habits of the species by day and their resting place at night are also worth
noting. The more recently evolved species, when not ftying or feeding, sit about
on ftowers and rest singly but more orten in groups during wet weather and at
night. The more primitive species fly for short periods in the sunshine and, after
feeding, rest singly on stems of plants and bushes.
vm
It is thought that the species in subgenus Mesembrynus, whose larvae feed on
Umbelliferae, Compositae and Labiatae, originated in the Tertiary (possibly the
middle Miocene) period. It is assumed that the subgenus Agrumenia originated
rather early, possibly at the end of the Miocene period and after the transition
of the larval feeding habits to the Papilionaceae bad taken place. The most re
cently evolved subgenus Zygaena probably originated from the middle of the
Pliocene period. It is not possible to conceive whether a second transition took
place of the larval feeding habits of the umbelliferous and composite feeders to
the Papilionaceae.
Many species of Mesembrynus, Agrumenia and Zygaena probably bad a far
wider distribution than they have in present times. The present, more localised
and discontinuous distribution was probably caused by the influence of the Iee
Age. It is thought that the more primitive species originated in and around the
Mediterranean region, spreading from here to the westem and northem parts
of Europe, to Mrica north of the Sahara, and north-east from Asia Minor to
Central Asia and Siberia. As far as it is known, only one species, niphona Butler,
has reached Japan. The distribution of the genus is palaearctic with the exception
of two species, viz., rubrico/lis Hampson and transpamirina Koch from Chitral.
In this catalogue the subspecies of each species are arranged according to the
geographical distribution, approximately from south to north and east to west.
The type locality of each subspecies is provided as it was originally quoted by
the author (to avoid errors in translation), and is placed in the righthand column
opposite the subspecific name. The names of forms and aberrations are arranged
under the subspecies to which they belong. The subspecific names are prefixed
with the term "ssp." and the names of aberrations are prefixed with the term
"ab.". We have refrained from using the term "f." ( = forma) for the aberrations,
since a form is considered tobe ofhigher status than an aberration as, for example,
seasonal forms (f.t. = forma tempestatis). The term "f." is used, however, to
denote the constantly recurring forms of the polymorphic species ephialtes. The
term "f. loc." ( = forma alicuius loci) is used to denote a local population which
differs sufficiently from the subspecies to merit a name. This terminology is based
on that of Rothschild and Jordan (1903, Novit. zool., 9, Supplement). The term
"var.", which has been used quite indiscriminately by many authors to denote
both subspecies or geographical races and aberrations, is here discarded as it has
no status in nomenclature. Junior Synonyms are placed in italics and are listed
under their respective senior synonyms. Junior homonyms are also given in italics
and are placed under an available, existing name.
We have experienced some difficulty in determining the dates of publication
of certain names of Esper, Hübner and Herrich-Schäffer. Esper's "Die Schmetter
linge" was issued in several parts. For the dates of publication of these various
parts we have referred to the paper by Sherbom & Woodward (1901, Ann. Mag.
nat. Hist., (7) 7: 137-140). The dates ofpublication ofthe various parts ofHübner's
"Sammlung europäischer Schmetterlinge" and "Verzeichniss bekannter Schmett
linge [sie]" are taken from Hemming's excellent work "Hübner", published in
two volumes by the Royal Entomological Society of London in 1937. The dates of
names published in Herrich-Schäffer's "Systematische Bearbeitung der Schmetter-
IX
Iinge von Europa" are also taken from Hemming's work. References to periodicals
are abbreviated and, where possible, follow the "World List of Scientific Periodi
cals, 1900-1950". Titles of books and other publications are quoted in full. All
references and the spelling of every name have been compared with the original
Iiterature for verification.
We are most grateful to Sylvia M. Tremewan, the wife of the junior author, for
her help in compiling the index of the catalogue, a formidable task, comprising
over three thousand four hundred names.
HUGO REISS, W. ÜERALD'fREMEWAN,
7 Stuttgart 1, 56, Hart Road,
Traubenstrasse 15B', Byfleet,
West Germany. Surrey, England.
10th September, 1966.
EIN
SYSTEMATISCHER KATALOG
DERGATTUNG
ZYGAENA FABRICIUS
(LEPIDOPTERA: ZYGAENIDAE)
von
HUGO REISS
und
W. GERALD TREMEWAN
