Table Of ContentF A U NA
NATURA
OPTIMA
M U N DI
DUX
Volume 3
Alexander G. Radchenko
Graham W. Elmes
Myrmica ants
(Hymenoptera: Formicidae)
of the Old World
Warszawa 2010
FAUNA
NATURA
OPTIMA
MUNDI DUX
Museum and Institute of Zoology
Polish Academy oí Sciences
EDITED BY
Dariusz Iwan (Editor in Chief)
Wojciech Czechowski, Wioletta Tomaszewska
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Natura optima dux Foundation
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ul. Wilcza 64, 00-679 Warszawa, Poland
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ABSTRACTED IN
Current Contents*/Agriculture, Biology, and Environmental Sciences;
Entomological Abstracts and Zoological Record.
The title partly sponsored by the Ministry of Science and Higher Education, Warsaw
Copyright by Natura optima dux Foundation, Warszawa 2010
ISSN 2081-4615
ISBN 978-83-930773-1-1
PRINTED IN POLAND BY
Warszawska Drukarnia Naukowa, Warszawa
Myrmica ants (Hymenoptera, Formicidae)
of the Old World
ALEXANDER G. RADCHENKO1 and GRAHAM W. ELMES2
1Museum and Institute of Zoology Polish Academy of Sciences, Wilcza 64,
00-679 Warszawa, Poland; e-mail: [email protected]
2CEH Research Fellow, Centre for Ecology and Hydrology, Crowmarsh Gifford,
Wallingford, Oxfordshire, OX10 8BB, United Kingdom; e-mail: [email protected]
ABSTRACT
The "Myrmica ants of the Old World" by A. G. Radchenko and G. W. Elmes,
is the first full taxonomic revision of this important Holarctic ant genus that is
abundant in many biotopes throughout the Palaearctic and parts of the Oriental
Regions. Myrmica ants have important functions in the dynamics of the plant and
animal communities of many temperate habitats, e.g. myrmechory, commensalism
and social parasitism. They are especially renowned for their role in the life cycle
of the endangered species of Large Blue Butterflies (the genus Phengaris, now
senior synonym of Maculinea). Latest European research shows that during the
larval stage most Phengaris populations are obligate social parasites of particular
species, or even particular populations of Myrmica, while in East Asia there are
many more species of Phengaris butterflies whose biology are poorly known, but
probably interact with Myrmica ants. Therefore besides being a rigorous taxonomic
study, this book is also aimed to support the many groups of scientists and nature
conservationists who need to identify Myrmica ants and to place species in their
context both within the genus as currently known and within the evolutionary
history of the genus.
The book opens with a preface section that briefly looks at the mention of Red
Ants (Myrmica rubra) in early literature and outlines the collaboration between
the authors: Alexander Radchenko is one of the leading ant taxonomists of his
generation and has developed a special knowledge of the Myrmica species of the
former Soviet Union; Graham Elmes is- a'well known social insect ecologist who
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has made special studies of the population dynamics of the genus Myrmica and
conservation problems of Phengaris butterflies. The preface is followed by 5 major
Chapters: it is not intended that these need be read sequentially, rather it is
expected that readers will dip in and out of the various chapters and sections as
required. Chapter 1 is an introduction to the genus, it starts by explaining the
scope and layout of the monograph and gives the reasons why the New World
fauna are not included, the main ones being that much primary taxonomy still
needs to be done on this fauna which shows no significant overlap with that of
the Old World. The main part of Chapter 1 comprises a well-referenced biological
review of the genus Myrmica sufficient to start a detailed literature search. It con
cludes with photographs and biographies of the 19 myrmecologists who have
described two or more "good" Myrmica species.
The core of the book is Chapter 3 entitled "Review of Species". Here the 426
names that have been ascribed to the genus Myrmica are listed and discussed in
various ways and the 142 extant and five extinct taxa considered to be good Old
World species are highlighted. These are then reviewed taxonomically in alpha
betical order and for each caste of each species a full set of accurate line drawings
are provided (using the type specimens where possible). The Keys to species are
provided in Chapter 5. Apart from the key to fossil species, the authors give
separate keys to the worker castes of extant species found within seven distinct
biogeographical regions. There is very little overlap between these regions and this
should make the keys less cumbersome for use by less specialist readers.
The remaining two chapters are more specialized and outline the authors'
latest views on the origins, evolution, spread and current distribution of the genus.
Chapter 2 starts by examining the taxonomic position and diagnosis of the genus
Myrmica in relation to other ants. The authors describe their concept of species
groups within the genus and list the main characteristics of the 17 groups that they
currently recognise, and the species therein. Recent molecular-genetic studies sug
gest that most of these groups have some phylogenetic value. Chapter 4 addresses
the zoogeography and evolution among the species listed in Chapter 3; Radchenko
and Elmes recognise nine major Zoogeographie areas that generally are quite dis
tinct from each other, the faunas of the two areas within the Oriental Region are
quite distinct from those of the Palaearctic Region. The paucity of fossil Myrmica
ants makes discussion of the timing of the evolution and spread of the genus Myr
mica very subjective. However, the authors give reasons for supposing that the
genus evolved in ancient Eurasia more than 45 Ma and had spread throughout
Eurasia by the start of the Miocene Epoch, probably invading North America via
the Beringian land bridge several times in subsequent ages.
KEY WORDS
Systematics, ants, Myrmica, taxonomy, revision, identification-keys, biogeo-
graphy, evolution, biological-review, fossil, Palaearctic and Oriental Regions.
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CONTENTS
Abstract 3
Preface 7
Chapter 1. General background and biology of Myrmica 13
1.1. Scope of this monograph 13
1.2. The layout and use of this monograph 14
1.3. The biology of Myrmica 16
1.3.1. Myrmica in ecosystems 16
1.3.2. Anatomy and surface chemistry 17
1.3.3. Colony structure 18
1.3.4. Social interactions 20
1.3.5. Physiology 23
1.3.6. Social parasitism 25
1.4. A brief history of the taxonomic investigation of Myrmica in the Old
World ' 27
1.4.1. Pre 1950 27
1.4.2. Post 1950 30
1.5. Morphometries and Myrmica taxonomy 32
1.5.1. History of measurement of Myrmica 32
1.5.2. A brief critique of the value of morphometries 33
1.5.3. Measurements and indices used in this monograph 35
1.6. Biographies of the authors 37
1.6.1. Biographies of the authors of this monograph 37
1.6.2. Biographies of other authors (ordered chronologically) 38
Chapter 2. Taxonomic structure of the genus Myrmica 51
2.1. Taxonomic position and diagnosis of the genus Myrmica 51
2.2. Species group concept in the genus Myrmica 58
Chapter 3. Review of species 69
3.1. Material studied and its deposition 69
3.2. The 303 names definitely attributable to the genus Myrmica 71
3.3. Review of extant Myrmica species of the Old World 79
3.3.1. Review of species 79
3.3.2. Drawings of the details of structure of extant Myrmica species . . . 331
3.4. Fossil species, undoubtedly belonging to the genus Myrmica 653
3.5. Names incertae sedis in the genus Myrmica 657
3:6: Nomina nuda in the genus Myrmica 663
3.7. Unavailable names in the genus Myrmica 663
3.8. List of taxa originally described in the genus Myrmica, but afterward
transferred to other genera or excluded from Myrmica 664
Chapter 4. Zoogeography and evolution of the genus Myrmica 671
4.1. The centres of Myrmica species diversity 672
4.2. Comparison of the regional faunas of Myrmica 678
4.2.1. Diversity between the nine centres 678
4.2.2. Interpretation of the diversity between the centres 686
4.3. Origin and evolution of the genus Myrmica 689
4.3.1. Putative ancestor and estimated time of Myrmica origin 689
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4.3.2. Evolution within the genus Myrmica and origin of the species
groups 691
4.3.2.1. Ancestral Myrmica forms and place of their origin 691
4.3.2.2. Origin of the species groups and their probable relation
ships 696
Chapter 5. Keys for the identification of the Myrmica species of the Old
World 711
5.1. A key for the identification of the Myrmica species of West Europe
and North Africa 713
5.2. A key for the identification of the Myrmica species of East Europe,
West Siberia, northern Kazakhstan, Caucasus, Asia Minor, Turkmeni
stan and Iran 721
5.3. A key to the identification of the Myrmica species of East Siberia,
Russian Far East, Mongolia, Korean Peninsula, northern China, and
Japan 729
5.4. A key for the identification of the Myrmica species of the Middle
Asian Mountains 734
5.5. A key for the identification of the Myrmica species of the Tibetan
Plateau 735
5.6. A key to the identification of the Myrmica species of the Himalaya and
Afghanistan 736
5.7. A key for the identification of the Myrmica species of South and
South-East Asia 740
5.8. A key for the identification of the fossil Myrmica species 744
References 745
Index of the all Latin names 781
Myrmica names recognized here as good species 781
Other names 785
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PREFACE
Red Ants - Myrmica
The name Red Ant as a description of reddish myrmicines, especially Myrmica ants,
does not appear in natural philosophical literature until the 17th century. However, it was
probably in common use much earlier because most modern European languages have some
version of the name. The ants that comprise the modern genus Myrmica must surely have
been recognized as a distinct form of animal life throughout most of the human cultural
history of the temperate Northern Hemisphere. They come to people's notice through their
readiness to administer irritating stings, especially in hot weather when it is easy to sit or
lie on an unnoticed nest. When one does so, the angry Myrmica workers have an uncanny
knack of getting under one's clothes where each can sting several times before exhausting
its supply of venom. For most people the result is a short-lived, localized stinging sensation
and, in some cases, small itchy bumps develop that can persist for several days. A few un
fortunate people can develop severe allergic reactions to the stings. The average Roman sol
dier, who served in northern Europe pitching camp in meadows and woodland clearings,
must have had some colourful names for Myrmica. During the European "Dark Age" the
tribal peoples who lived close to nature recognized many common insects, and although
we know of no name that refers directly to Myrmica they probably had one, the Saxon
peoples had a variety of words for ant, anthill, ant-nest, sting etc. Even today in England,
Myrmica ants are often called "Picnic Ants" or "Red Stinging Ants" while the name
"European Fire Ant" has been coined recently for the pest populations of Myrmica rubra
(L.) introduced into the USA.
Ants were well known to the classical Greek writers and although they are not discussed
at any length by Aristotle (384-322 С)В in his "Historia Animalium" it is clear from his
knowledge and treatment of other insect groups that he recognized different sorts of ants.
The Greek understanding of natural history was barely advanced by European writers until
the 17th century. The Roman Pliny the Elder (20-79) re-hashed a lot of Aristotle's work
and compounded it with various myths and legends, while Claudius Aelianus (ca. 175-235)
simply reproduced much of Pliny's work with added errors. There is no description that
can be linked to the genus Myrmica which is hardly surprising given that classical study was
centred on the Mediterranean region where species of Myrmica are rare.
In northern Europe some of the early writers showed considerable knowledge of plants
and animals, including ants, but they were usually more interested in their medicinal
properties rather than their natural history, e.g. the 12th century Abbess Hildegard of
Bingham. The Dutchman, Albertus Magnus (1205-1282) was instrumental in reviving the
classical knowledge of natural philosophy by reiterating much of Aristotle's, Pliny's and
Aelian's works with some additions from Arabian studies. By the 16th century, there was
a sudden surge of interest in natural history with numerous works based on the classic
writers: e.g. the Swiss, Conrad Gesner (1516-1565), and the Bolognian, Ulysses Aldrovandi
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(1527-1605). Aldrovandi devoted about 30 manuscript pages to ants, including a section
on types of ants, although it added little to the work of his predecessors it nicely
illustrates how by the mid 16th century natural historians were beginning to recognize the
great diversity of insects from different regions of the world. We have no information on the
extent of oriental knowledge of types of ants, but given the highly realistic pictorial depic
tion of plants and animals that had developed in China by the 16th century it was probably
considerable.
So, it was not until the 17th century that the first formal descriptions of native Euro
pean ants were made. The British natural historian, philosopher and theologian John Ray
(1628-1705), who some consider to have influenced Linnaeus' work, recognized five
species of ant in Britain (Wheeler 1926, p. 223). The posthumously published "Historia In-
sectorum" (Ray 1710) as Ray always acknowledged, owed a considerable amount to his
pre-deceased friend and colleague Francis Willughby; whichever of them contributed the
section on ants they recognized the following names: Formica maxima hippomyrmex,
Formica media ruja, Formica media nigro colore splendens, Formica minima rubra,
Formica minor ex fusca nigricans, that Wheeler equated to Camponotus herculeanus (L.),
Formica rufa L., Formica fusca L., Myrmica rubra (L.) and Lasius niger (L.) respectively
(though the first can not be C. herculeanus because Camponotus species are not present in
Britain). Some thirty years later (ca. 1743) this classification was well known in Europe.
For example, in an unpublished text the 18th century French natural philosopher Reaumur,
acknowledged great diversity among ants: "Ray n'en observe en angleterre que cinque
espèces; les descriptions qu'il en a faites m'ont mis en etat de juger que les mêmes sont
aussi en france, ou il est facile d'en trouver beaucoup plus d'espèces" (Wheeler 1926).
This then was the background knowledge available to 18th century writers such as the
Rev. William Gould (1747) who published the greatly under-rated volume "An Account of
English Ants", the first book on ants in the English language. Throughout, Gould compared
the biology of five species: the Hill Ant, the Jet Ant, the Red Ant, the common Yellow Ant
and the small Black Ant, that equate to F. rufa-like, F. fusca-like, M. rubra-like, Lasius
flavus-like and L. niger-like species. Gould used much the same morphological features
used earlier by Ray and later by Linnaeus, De Geer and others to define Red Ants. In his
first chapter on the various kinds of ants and their structure, Gould writes (ibid., p. 8)
"The ligament [petiole] that unites the breast [mesosoma] and body [gaster] in the Red
Ant consists of two lobes somewhat round and divided in the middle [petiole and
postpetiole]. A fine thread connects it with the breast, and another with the body. In other
ants their [there] appears but one lobe between the threads; which rises higher and is
broader than the lobes in Red Ants. This species has also at the end of its breast two sharp
prickles [propodeal spines]". This is followed (ibid., p. 9) by "The Red Species has also
a sting of the same contexture as a bee, only in miniature. With this it opens an orifice in
the flesh, and afterwards injects a venom which gives a smart but momentary pain". From
all the subsequent ecological description there is no doubt that in the main, Gould was
thinking of the genus Myrmica when discussing Red Ants.
The origin of our collaboration
Graham Elmes first studied Myrmica ants when he was recruited into the British Nature
Conservancy in 1961, to be an assistant to Michael V. Brian (1919-1990), a leading British
8
ant specialist at that time. Although employed as an ecologist, Brian was an "experimental
biologist" of the "Cambridge School" whose main interest was social biology, trying to un
derstand how queens and workers interact to regulate factors such as colony growth, worker
oviposition and sexual production. Myrmica rubra was the main "research model" species,
used because its colonies frequently had many queens. This enabled one to make many
small replicated experimental cultures of ants to test the effect of single variables (e.g.
presence or absence of a queen, high and low temperatures or different types of food). While
such experiments have their limitations, they were extremely informative and form a bench
mark against which many more recent studies fall short. The underlying rationale driving
Brian's team at that time could be summarized as "What we find for M. rubra almost cer
tainly holds true for all Myrmica ants and probably all myrmicine ants, if not all ant species";
a view that in hindsight seems hopelessly naive.
The fact that most other common Myrmica species had fewer queens than M. rubra in
trigued Elmes and he assumed that the difference might have an ecological basis. This re
sulted in his 1972-1974 PhD study of the comparative ecology of Myrmica (southern
English species). He developed the idea that polygyny is an adaptive trait within different
local populations. Costs to individual queens being mitigated by shared benefits accrued at
the colony level, and so the interaction between queens evolved various checks and
balances that ensure an "ideal density" of queens per colony. A single mechanism would
thus explain the observed variation in queen-density between conspecific populations and
between species. He proposed an ¡dea that met with some scepticism, that colonies
periodically recruited new, often unrelated queens and that "surplus" queens (beyond the
minimum number required to maintain the worker population) could be thought of as tem
porary social parasites (Elmes 1973a). Thus, while polygyny might be adaptive in some
circumstances, it always has the danger in some populations of degenerating into real
social parasitism; therefore Elmes regarded polygyny and microgyny as exhibited by
M. rubra as an "evolutionary precursor" to social parasitism (Elmes 1978a). The German
myrmecologist, Alfred Buschinger, explored similar ideas at about the same time to explain
social parasitism in Leptothorax Mayr species (Buschinger 1970).
Elmes' PhD study highlighted the fact that in many respects M. rubra does not provide
a model for all Myrmica species, rather it is the most unusual among the common British
species. Furthermore, as his post-doc studies developed, he found that population dynamics
and individual traits, e.g. basal physiology, larval ontogeny, varied between populations of the
same species. Consequently, he started retaining voucher specimens, working on the prin
ciple that if one is going to compare different populations of the same species then one needs
to be sure that the populations are not different species. Elmes continued the practice of
voucher specimens when in 1980 he started a more applied study, collaborating with Jeremy
Thomas, on the biology and conservation of Large Blue Butterflies of Europe (the genus
Phengaris Doherty = Maculinea Van Eecke; for synonymy see Fric et al. 2007), which still
continues (Settele et al. 2005). This study chimed with Elmes' interest in social parasitism
in the genus Myrmica, because the 4th instar larvae of Maculinea butterflies can be consid
ered as social parasites of Myrmica colonies. The details of the interaction vary between host
and parasite populations and may be related to variation between local races of Myrmica
throughout Europe. Examination of the voucher specimens led to Elmes' interest in Myrmica
taxonomy and since its modest beginnings of a few hundred British specimens it has grown
into probably the largest collection of a single ant-genus. It is now one of the major resources
that underpins this monograph.
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