Myrmecological News 14 87-96 Vienna, January 2011
Worldwide spread of the ruby ant, Myrmica rubra (Hymenoptera: Formicidae)
James K. WETTERER & Alexander G. RADCHENKO
The ruby ant, Myrmica rubra (LINNAEUS, 1758) (formerly Myrmica laevinodis NYLANDER, 1846), an aggressive Eur-
asian species with a powerful sting, is now spreading through temperate North America. To document the worldwide
distribution of M. rubra and evaluate its potential for further spread, we compiled published and unpublished specimen
records from > 2000 sites. We report the earliest known M. rubra records for 71 geographic areas (countries, major is-
lands, US states, Canadian provinces, and Russian federal districts), including three areas with no previously published
records: Prince Edward Island, Washington State, and the Far Eastern Federal District of Russia. All earlier published
records of M. rubra from East Asia, including the Far East of Russia, Japan, and China, appear to be misidentifications
of Myrmica kotokui FOREL, 1911.
Myrmica rubra is native to an enormous expanse extending from Ireland and Portugal in westernmost Europe across
8000 km to central Asia and eastern Siberia, and from 39 to 70° N in latitude. Exotic populations of M. rubra were first
recorded in eastern North America more than 100 years ago. Myrmica rubra is now documented from five southeastern
Canadian provinces (New Brunswick, Nova Scotia, Ontario, Prince Edward Island, and Quebec), six northeastern US
states (Maine, Massachusetts, New Hampshire, New York, Rhode Island, and Vermont), and one northwestern state (Wash-
ington) ranging from 41.5 to 47.6° N.
Given the vast range of M. rubra in Eurasia, perhaps the most striking aspect about this species in North America is how
little it has spread over the past century. Most North American records of M. rubra, however, date from the last ten years,
suggesting these North American populations are expanding. There appear to be no geographic barriers that would pre-
vent M. rubra from spreading across the US and Canada, from coast to coast.
Key words: Biogeography, biological invasion, exotic species, invasive species, stinging ant.
Myrmecol. News 14: 87-96 (online 26 August 2010)
ISSN 1994-4136 (print), ISSN 1997-3500 (online)
Received 26 January 2010; revision received 29 April 2010; accepted 29 April 2010
Prof. Dr. James K. Wetterer (contact author), Wilkes Honors College, Florida Atlantic University, 5353 Parkside Drive,
Jupiter, FL 33458, USA. E-mail: firstname.lastname@example.org
Prof. Dr. Alexander G. Radchenko, Museum and Institute of Zoology of the Polish Academy of Science, Wilcza str. 64,
00-679 Warsaw, Poland. E-mail: email@example.com
Numerous ant species have spread around the world through
human commerce. Only a few of these are known to have
significant ecological and / or economic impacts. Most of
the destructive invasive ants have spread through tropical
and subtropical regions, often achieving extremely broad
distributions in both the Old and New World, e.g., Mono-
morium destructor (JERDON, 1851), Pheidole megacephala
(FABRICIUS, 1793) and Linepithema humile (MAYR, 1868)
(WETTERER 2007, 2009, WETTERER & al. 2009). One po-
tentially destructive Eurasian ant now spreading through
temperate North America is the ruby ant, Myrmica rubra
(LINNAEUS, 1758). Reporting this ant from Massachusetts,
WHEELER (1908) wrote that M. rubra (as its junior synonym
Myrmica laevinodis NYLANDER, 1846) "is the most dis-
agreeable of the palaearctic Myrmicas ... its workers are ag-
gressive and sting severely. It is very fond of attending aph-
ids and, unlike our timid native Myrmicas, which live in the
retirement of woods, bogs, heaths and waste places gen-
erally, it prefers to nest in cultivated soil. Hence it may be-
come a nuisance in lawns and dooryards." Here, we docu-
ment the worldwide distribution of M. rubra and specu-
late on its future spread.
Taxonomy and identification: Myrmica rubra has
reddish-brown monomorphic workers (4 - 5 mm total
length; Figs. 1 - 3). Most strikingly, "M. rubra is a very
aggressive species which stings freely" (SEIFERT 1988).
COLLINGWOOD (1987) wrote that M. rubra "has the most
vicious sting of all the British ants, comparable with that
from a stinging nettle."
LINNAEUS (1758) first described Formica rubra (= M.
rubra) from Europe. LINNAEUS (1767) later added "pessi-
me nostratum pungit" (= "it stings the worst of our native
species") to its description. Unfortunately, LINNAEUS' (1758,
1767) written description matched more than one ant spe-
cies. It was generally presumed that M. rubra was the se-
nior synonym of either Myrmica ruginodis NYLANDER, 1846
or Myrmica laevinodis NYLANDER, 1846 (sometimes spel-
led levinodis), because these two European species have the
most powerful stings (EMERY 1908, DONISTHORPE 1913,
1915). Some authors considered these to be two subspecies
of M. rubra, with no nominal subspecies (i.e., M. rubra
rubra). SANTSCHI (1931) speculated (incorrectly) that M.
rubra was the senior synonym of M. ruginodis, and many
researchers accepted this judgment (e.g., WEBER 1947,
BRIAN & BRIAN 1949, CREIGHTON 1950). This error was
corrected when YARROW (1955) examined all Myrmica
specimens in the Linnaean type collection and concluded
that the type of M. rubra was actually the senior syno-
nym of M. laevinodis. Thus, all published records of M.
laevinodis refer to M. rubra. A few pre-1955 records of
M. rubra, however, actually refer to M. ruginodis, but these
can be readily recognized if an author refers to M. laevi-
nodis and M. rubra as distinct species (e.g., BRIAN &
Junior synonyms of M. rubra include M. laevinodis
(described from Finland and Sweden; synonymized with
M. rubra by YARROW 1955), Myrmica longiscapus CUR-
TIS, 1854 (described from Scotland and England; synony-
mized with M. laevinodis by MAYR 1863), Myrmica laevi-
nodis europaea FINZI, 1926 (first available use of Myrmica
rubra ssp. champlaini var. europaea FOREL, 1911) (de-
scribed from Norway; synonymized with M. rubra by
RADCHENKO & al. 1997), Myrmica laevinodis bruesi WE-
BER, 1947 (first available use of Myrmica rubra laevinodis
var. bruesi WHEELER, 1906) (described from Massachusetts;
synonymized with M. laevinodis by CREIGHTON 1950),
and Myrmica microrubra SEIFERT, 1993 (described from
Germany; synonymized with M. rubra by STEINER & al.
2006). One extant subspecies, Myrmica rubra neolaevi-
nodis FOREL, 1901, was described from specimens inter-
cepted in New York on flowers imported from Hamburg,
PROVANCHER (1887) synonymized Myrmica incompleta
PROVANCHER, 1881 (described from Canada) with M. lae-
vinodis (= M. rubra). FRANCOEUR & BEIQUE (1966), how-
ever, revived M. incompleta, recognizing it as the senior
synonym of Myrmica brevinodis EMERY, 1895, a species
known from across Canada and the northern US. SMITH
(1951) synonymized Myrmica rubra champlaini FOREL,
1901 (described from Quebec) with M. laevinodis, but GRO-
DEN & al. (2005) instead synonymized it with Myrmica
brevispinosa WHEELER, 1917.
RADCHENKO (1994) placed M. rubra with ten other spe-
cies in the M. rubra species-group. Within this group, M.
rubra and M. ruginodis formed a clade whose sister group
included the other nine species, all from Asia: Myrmica
dshungarica RUZSKY, 1905 (described from Kazakhstan),
Myrmica tibetana MAYR, 1889 (from Tibet), Myrmica chi-
nensis VIEHMEYER, 1922 (from China), Myrmica smythiesii
FOREL, 1901 (from India), Myrmica juglandeti ARNOL'DI,
1976 (from Kyrgyzstan), Myrmica kryzhanovskii ARNOL'DI,
1976 (from Tajikistan), Myrmica ferganensis FINZI, 1926
(from Kyrgyzstan), Myrmica dicaporiaccoi MENOZZI, 1939
(from India), and Myrmica everesti DONISTHORPE, 1929
(from near Mount Everest). After further analyses, how-
ever, A.G.R. now considers the three closest relatives of
M. rubra to be M. ruginodis, Myrmica arisana WHEELER,
1930, and Myrmica kotokui FOREL, 1911 (A.G. Radchenko,
unpubl.), and excludes the other species mentioned above
from the M. rubra species-group. Myrmica arisana is known
rom Taiwan (WHEELER 1930) and Myrmica kotokui is
known from Japan, Korea, and the Russian Far East (COL-
LINGWOOD 1976, RADCHENKO 2005).
Myrmica rubra can be distinguished from M. ruginodis
by the shape and sculpture of the petiolar node and the
propodeal spine length. The petiolar node is rounded in M.
rubra, but flat-topped in M. ruginodis; the sides of node
are striated or at most finely rugulose in M. rubra, but
coarsely rugose in M. ruginodis (laevinodis means "smooth
node," whereas ruginodis means "wrinkled node.") Also,
M. rubra has relatively shorter propodeal spines than does
M. ruginodis. In the past, many authors recognized inter-
mediate forms between M. rubra and M. ruginodis based
on propodeal spine length. BRIAN & BRIAN (1949), how-
ever, found that propodeal spine length is correlated with
worker size, such that large M. rubra workers have the
same spine length as small M. ruginodis workers. BRIAN &
BRIAN (1949) found that when comparing the spine length
to head width ratio, M. rubra and M. ruginodis are simple
Common names: In Eurasia, common names for M.
rubra include the red ant and common red ant (and their
equivalents in other languages), names that are also ap-
plied to several other species. Recently, North American
researchers have begun calling M. rubra the European red
ant or European fire ant. We consider all these names to be
unsatisfactory. The only distinctive name is European fire
ant, and this name is misleading because it falsely suggests
a kinship with "true" Solenopsis fire ants. In addition, M.
rubra is not strictly European; it is also native in much of
northern Asia as well, and most of its closest relatives are
strictly Asian species. In Latin, rubra means ruby, dark red,
ruddy, reddish, blushed, flushed, or shamed. We therefore
suggest "ruby ant" as a new English common name for M.
We documented the worldwide range of Myrmica rubra
using both published and unpublished records. We ob-
tained unpublished site records from museum specimens
in the collections of the Museum of Comparative Zoology
(MCZ), Archbold Biological Station (ABS), Zoological
Museum of the Moscow State University (ZMMU), Zoo-
logical Institute of the Russian Academy of Sciences, St.
Petersburg (ZISP), Schmalhausen Institute of Zoology of
the Ukrainian National Academy of Sciences, Kiev (SIZK),
and the Museum and Institute of Zoology of the Polish
Academy of Sciences, Warsaw (MIZ). Stefan Cover iden-
tified all specimens at the MCZ and ABS; A.G.R. identi-
fied all specimens at the ZMMU, ZISP, SIZK, and MIZ.
In addition, we used on-line databases with collection in-
formation on specimens by Antweb (www.antweb.org) and
the Global Biodiversity Information Facility (www.gbif.
org). We also received unpublished records from Herbert
Zettel (Czech Republic, Serbia, Slovenia), S. Tschesno-
kova (Russia), R. Schultz (Kazakhstan, Kyrgyzstan, and
Russia), J. Huber (Canada), A. Francoeur (Canada), G. El-
mes (Kazakhstan, Kyrgyzstan, and Russia), and M. Bustos
Geographic coordinates for collection sites came from
published references, specimen labels, maps, or geography
web sites (e.g., earth.google.com, www.tageo.com, and
www.fallingrain.com). If a site record listed a geographic
region rather than a "point locale," and we had no other
Figs. 1 - 3: Myrmica rubra. (1) Head
of worker from Italy; (2) lateral
view of the same worker; (3) dorsal
view of the same worker (photos by
A. Nobile; copyright AntWeb.org).
record for this region, we usually used the coordinates of
the largest town within the region. If one source had many
sites less than 10 - 20 km apart (e.g., KOFLER 1995, SCHLICK-
STEINER & STEINER 1999), we often did not plot every site.
We did not map records of M. rubra intercepted in
shipments by quarantine inspectors, e.g., GRODEN & al.'s
(2005) records of M. rubra found in cargo arriving in
Massachusetts (from Poland), New Jersey (from Holland
and England), New York (from Germany), Pennsylvania
(from Ireland & Germany), and Washington DC (from Ger-
We compiled published and unpublished specimen records
from > 2000 sites. We documented the earliest known M.
rubra records for 71 geographic areas (countries, major
islands, US states, Canadian provinces, Russian federal
districts; Tabs. 1 - 4), including the first records of M.
rubra from Prince Edward Island (det. S. Cover), Wash-
ington (det. S. Cover), and the Far Eastern Federal Dis-
trict of Russia (det. A.G.R.). For five of these geographic
areas, we had no specific site record, so we mapped the
record to the largest city (Bosnia mapped to Sarajevo, Ka-
liningrad Oblast mapped to Kaliningrad), or in the case of
three southern areas, we mapped a northern mountain (Ko-
ula in Greece, Monte das Raposas in Portugal, and Mahya
Dagi in Thrace).
Myrmica rubra has records from a large area of the
Palaearctic stretching from Ireland and Portugal in West-
ern Europe across 8000 km to central Asia and Siberia,
and spread in latitude from 39 - 70° N (> 95% of records
from 40 - 65° N; Fig. 4; Tabs. 1 - 3). At lower latitudes, M.
rubra is usually found only at higher elevations. In conti-
nental Europe, M. rubra is known from all countries except
Albania, Monaco, and San Marino (Tabs. 1 - 3). Most M.
rubra records come from the western part of its native
range, but this appears to be largely due to greater sam-
pling and most accessible data in Western and Central
Fig. 4: Worldwide distribution records of Myrmica rubra. ? = questionable record (see text).
Europe. Similarly, the dates of the earliest records of M.
rubra in different parts of its native range have little sig-
nificance beyond indicating where early ant research was
conducted and the results published in accessible sources.
In North America, M. rubra has been reliably recorded
from five southeastern Canadian provinces (New Bruns-
wick, Nova Scotia, Ontario, Prince Edward Island, and
Quebec), six northeastern US states (Maine, Massachusetts,
New Hampshire, New York, Rhode Island, and Vermont),
and one northwestern state (Washington) (Tab. 4). Most M.
rubra records in North America are from the past ten years.
In June 2008, J.K.W. collected M. rubra at eight sites
on Prince Edward Island, a Canadian province where M.
rubra has not been previously reported (geo-coordinates
in parentheses): Dalvay, by hotel (46.416, -63.074), Char-
lottetown, Victoria Park (46.229, -63.139), Brookvale Pro-
vincial Park, forest (46.275, -63.421), Darnley, forest patch
(46.546, -63.642), Brundenell River Provincial Park, by
river (46.199, -62.576), Sally’s Beach Provincial Park, near
beach (46.264, -62.380), Seal Cove, campground (46.046,
-62.523), and Portage, by bike trail (46.665, -64.058). J.K.W.
also collected M. rubra at a highway rest area in Med-
way, Maine (45.596, -68.535) and on the roadside by the
ferry terminal on Deer Island, New Brunswick (44.928,
-66.985). The records came from a wide variety of habitats,
including an urban park, the manicured grass of a hotel,
beachfront scrub, and intact forest.
Questionable records: We have mapped ten question-
able records from sites that fall outside the confirmed range
of M. rubra (question marks in Fig. 4). Although we be-
lieve that all these records of M. rubra are probably based
on errors of identification or location, we included them
because they have been largely overlooked and deserved to
Two early voyages to Arctic Greenland and neighbor-
ing parts of North America, both listed M. rubra as the
only ant species recorded. From an 1824 - 1825 voyage,
ROSS (1826) wrote that Formica rubra (= M. rubra) was
"abundant at the Whale-fish Islands; it was also found, on
the preceding voyage, on several parts of the Melville Pen-
insula" (in Greenland and Nunavut, respectively). From an
1829 - 1833 Arctic voyage, CURTIS (1835) listed Myrmica
rubra as "Numerous, under stones," but gave no site infor-
mation. These records, however, predate the descriptions of
many related species (including M. ruginodis), and could
be misidentifications. WEBER (1953) asserted that these rec-
ords were probably misidentifications of Formica fusca
LINNAEUS, but gave no rationale for this conclusion. How-
ever, it seems unlikely that CURTIS (1835) would confuse
ants in two such fundamentally dissimilar genera, the name-
sakes of two subfamilies (Myrmica in Myrmicinae versus
Formica in Formicinae). In fact, CURTIS (1854) described
Myrmica longiscapus (= M. rubra) and wrote keys distin-
guishing Myrmica from Formica. It seems more likely that
the ants were a species of Myrmica (e.g., Myrmica alask-
ensis WHEELER, 1917) or another myrmicine species (e.g.,
Leptothorax acervorum (FABRICIUS, 1793)) known from
Arctic Canada and Alaska. Remarkably, we have found no
other ant records of any kind from Nunavut or Greenland.
In fact, recent authors commonly state that no ants now live
in Greenland, though there are fossil ants from northern
Greenland (e.g., BENNIKE & BÖCHER 1990, HEINZE 1993).
Someone needs to go look for these ants.
ANDRÉ (1883) listed the range of M. laevinodis (= M.
rubra) as including North America, though we do not know
the source of this information. MAYR (1886), in his review
of the ants of North America, wrote of M. laevinodis (= M.
rubra): "This species is known to me only from Labrador,"
then reported M. ruginodis from Colorado and Virginia.
We do not know the sources of these records. ASHMEAD
(1902) reported M. laevinodis (= M. rubra) collected from
Nushagak River in Alaska, noting that this species "occurs
also in Siberia and various parts of the United States." Yet
we have been unable to find any other record of this species
from the United States published before WHEELER (1906).
We also mapped questionable records of M. laevinodis
(= M. rubra) from Kashmir (FOREL 1906), North India (KA-
RAWAJEW 1934), and Turkmenistan (KARAWAJEW 1934).
Tab. 1: Earliest known records for Myrmica rubra from its
native range in Western Europe. ENB = inatura - Erlebnis
Naturschau Dornbirn, ZMUC = University of Copenhagen
≤ Earliest record
Andorra ≤ 2006 (BERNADOU & al. 2006)
Austria ≤ 1853 (J. Möller, ENB): many sites
Belgium ≤ 1897 (GASPAR 1966 as M. laevinodis)
Denmark ≤ 1853 (Meinert, ZMUC): Amager
England ≤ 1852 (ROEBUCK 1877 as M. laevinodis)
Finland ≤ 1846 (NYLANDER 1846 as M. laevinodis)
France ≤ 1855 (MAYR 1855 as M. laevinodis)
Germany ≤ 1852 (SCHENCK 1852 as M. laevinodis)
Ireland ≤ 1898 (CUTHBERT 1898 as M. laevinodis)
Isle of Man ≤ 1976 (BARONI-URBANI & COLLINGWOOD
Italy ≤ 1853 (MAYR 1853 as M. laevinodis)
Liechtenstein ≤ 2004 (RADCHENKO 2004)
Luxembourg ≤ 1953 (STUMPER 1953 as M. laevinodis)
Netherlands ≤ 1887 (BOS 1887 as M. laevinodis)
≤ 1896 (JOHNSON 1896 as M. laevinodis)
Norway ≤ 1898 (STRAND 1898 as M. levinodis)
Portugal ≤ 1979 (COLLINGWOOD 1979)
Scotland ≤ 1825 (CURTIS 1854 as M. longiscapus)
Spain ≤ 1879 (MARTORELL Y PEÑA 1879 as M.
Sweden ≤ 1846 (NYLANDER 1846 as M. laevinodis)
Switzerland ≤ 1855 (MAYR 1855 as M. laevinodis)
Vatican City ≤ 1855 (MAYR 1855 as M. laevinodis)
Wales ≤ 1913 (DONISTHORPE 1913 as M. laevinodis)
It is very likely that the records from Kashmir and North
India are based on misidentifications of one or more close-
ly related Myrmica species found in the Himalayas (see
above). The record from Turkmenistan may be a locality
error based on the record of M. laevinodis (= M. rubra) in
MAYR (1877) from Turkestan, a name formerly used for the
region encompassing much of Kazakhstan, Kyrgyzstan,
Tajikistan, Turkmenistan, and Uzbekistan. MENOZZI (1939)
listed M. laevinodis (= M. rubra) among the 22 species of
Myrmica known from the Himalayas and Tibet.
GLEASON (1909) listed M. rubra on Isle Royale, Mich-
igan, but it seems much more likely that this record was ac-
Tab. 2: Earliest known records for Myrmica rubra from its
native range in Central and Southeastern Europe.
≤ Earliest record
≤ 1898 (WASMANN 1898 as M. laevinodis)
Bulgaria ≤ 1891 (FOREL 1892 as M. laevinodis)
Croatia ≤ 1890 (KORLEVIC 1890 as M. laevinodis)
≤ 1855 (MAYR 1855 as M. laevinodis)
Greece ≤ 1985 (AGOSTI & COLLINGWOOD 1987)
Hungary ≤ 1869 (FRIVALDSZKY 1869 as M. laevinodis)
Macedonia ≤ 1992 (PETROV & COLLINGWOOD 1992)
Montenegro ≤ 1922 (KULMATYCKI 1922 as M. laevinodis)
Poland ≤ 1855 (MAYR 1855 as M. laevinodis)
Romania ≤ 1853 (FUSS 1853 as M. laevinodis)
Serbia ≤ 1950 (ZIVOJINOVIC 1950 in PETROV & al. 2005)
Slovakia ≤ 1907 (CSIKI 1909 as M. laevinodis)
Slovenia ≤ 1855 (MAYR 1855 as M. laevinodis)
≤ 1985 (AGOSTI & COLLINGWOOD 1987)
tually some North American Myrmica that was considered
a subspecies of M. rubra at the time, e.g., M. rubra cham-
plaini (= M. brevispinosa). In 1909, virtually all records
of true M. rubra were reported as M. laevinodis. CAGNIANT
(1962) recorded M. rubra from the Atlas Mountains of
Morocco. However, CAGNIANT (2006) compiled a list of ant
species known from Morocco that did not include M. rubra,
but had four other Myrmica. A.G.R. examined an old spe-
cimen (probably pre-1900) of M. rubra labeled simply "Al-
geria" in the ZISP collection. This record appears to fall
outside the range of M. rubra and could be from an intro-
duced population, intercepted in quarantine, or simply mis-
For five of the questionable records, we had no specific
site within the region; we mapped these as follows: Algiers
in Algeria, Srinagar in Kashmir, Goose Bay in Labrador,
Leh in North India, and Daşoguz in Turkmenistan.
Erroneous records: We excluded from the map the
many published records of M. rubra from East Asia, in-
cluding the Far East of Russia (ONOYAMA 1989, KUPY-
ANSKAYA 1990, KUPYANSKAYA & LELEJ 2000, KUPYANS-
KAYA & al. 2000), Japan (FOREL 1901, 1907, TERANISHI
1929, EMERY 1908, 1921, ONOYAMA 1989, IMAI & al.
2003), and China (e.g., EMERY 1908, COLLINGWOOD 1962,
CHANG & HE 2001, AZUMA & al. 2005), because all are al-
most certainly misidentifications. For example, RADCHENKO
(2005) determined the East Asian M. rubra records in FO-
REL (1907), EMERY (1908, 1921), WEBER (1948), and ONO-
YAMA (1989) all to be M. kotokui.
HOLGERSEN (1943) recorded M. rubra (as M. laevino-
dis) from "Sajan, Sistikem, Mongolia," but Systyg Khem in
Tab. 3: Earliest known records for Myrmica rubra from its
native range in Eastern Europe, Russia, the Caucasus, and
Asia. ZMMU = Zoological Museum of the Moscow State
University. + = no previously published records.
≤ Earliest record
+ Armenia ≤ 1900 (RUZSKY 1905 as M. laevinodis)
+ Azerbaijan ≤ 1898 (RUZSKY 1905 as M. laevinodis)
+ Belarus ≤ 1904 (RUZSKY 1905 as M. laevinodis)
+ Estonia ≤ 1887 (MÜHLEN 1887 as M. laevinodis)
+ Georgia ≤ 1899 (RUZSKY 1905 as M. laevinodis)
+ Kaliningrad ≤ 1939 (JACOBSON 1939 as M. laevinodis)
+ Kazakhstan ≤ 1896 (RUZSKY 1905 as M. laevinodis)
+ Kyrgyzstan ≤ 1976 (TARBINSKY 1976)
+ Latvia ≤ 1889 (NASSONOV 1889 as M. laevinodis)
+ Lithuania ≤ 1889 (NASSONOV 1889 as M. laevinodis)
+ Moldova ≤ 1965 (CÎRDEI & BULIMAR 1965 as M.
≤ laevinodis in WOJCIK & PORTER 2008)
+ Russia: Central ≤ 1889 (NASSONOV 1889 as M. laevinodis)
+ Russia: Far
≤ date unknown (collector unknown,
≤ ZMMU): Sakha Republic, Saldykel'
+ Russia, N.
≤ 1899 (Ruzsky 1905 as M. laevinodis)
+ Russia: North-
≤ 1889 (NASSONOV 1889 as M. laevinodis)
+ Russia: Siberian ≤ 1856 (NYLANDER 1856 as M. laevinodis)
+ Russia: Southern ≤ 1849 (NYLANDER 1849 as M. laevinodis)
+ Russia: Urals ≤ 1894 (RUZSKY 1905 as M. laevinodis)
+ Russia: Volga ≤ 1894 (RUZSKY 1905 as M. laevinodis)
+ Tajikistan ≤ 1870 (MAYR 1877 as M. laevinodis)
+ Turkey: Kars ≤ 1902 (RUZSKY 1905 as M. laevinodis)
+ Ukraine ≤ 1892 (NASSONOV 1892 as M. laevinodis)
the Sayan Mountains is actually located in Tuva, Siberia.
Subsequent lists of Mongolian ant species that include M.
rubra appear to be based on this error.
Myrmica rubra has a very broad native distribution in Eur-
asia, spread over 30° of latitude and extending 8000 km
across Europe and Siberia (Fig. 4). SEIFERT (1988) wrote
that M. rubra ranged from Portugal to eastern Siberia and
from Italy to northern Scandinavia, but in the Mediterrane-
an region it was only found in moist places. RADCHENKO
(2005) wrote: "M. rubra is a Euro-Siberian species, whose
eastern limit of distribution is Lake Baikal." The present an-
alysis largely agrees with these earlier assessments, though
Tab. 4: Earliest known records for Myrmica rubra from its
exotic range in North America. ABS = Archbold Biological
Station, MCZ = Museum of Comparative Zoology. + = no
previously published records.
≤ Earliest record
+ Massachusetts ≤ 1900 (WHEELER 1906 as M. laevi-
≤ nodis bruesi)
+ Quebec ≤ 1915 (GRODEN & al. 2005)
+ New York ≤ 1936 (LINDSEY 1939)
+ Maine ≤ 1952 (GRODEN & al. 2005)
+ Rhode Island ≤ 1966 (GRODEN & al. 2005)
+ Ontario ≤ 1975 (GRODEN & al. 2005)
+ New Hampshire ≤ 1977 (GRODEN & al. 2005)
+ Washington ≤ 1988 (M. Deyrup, ABS): Seattle
+ Nova Scotia ≤ 1998 (GRODEN & al. 2005)
+ New Brunswick ≤ 2005 (GRODEN & al. 2005)
+ Vermont ≤ 2005 (GRODEN & al. 2005)
+ Prince Edward Is. ≤ 2008 (J.K. Wetterer, MCZ): Dalvay
with a distribution reaching somewhat east of Lake Bai-
kal, with one record from the Sakha Republic of the Russi-
an Far Eastern Federal District (Tab. 3, Fig. 4). We found
no M. rubra records from Uzbekistan and most of Kaz-
akhstan. We believe that this gap in the distribution of M.
rubra in Central Asia is real and not due solely to poor
sampling. Almost all site records from Kazakhstan came
from the mountainous regions along its northern and south-
eastern borders. The arid conditions over much of Uzbeki-
stan and Kazakhstan may exclude M. rubra.
DONISTHORPE (1915) noted that M. rubra (as M. laevi-
nodis) occurs "further south in the mountains." Indeed, most
of the lowest latitude records of M. rubra across Eurasia
(39 - 42° N) were from high elevation regions in the Py-
renees, Alps, Rhodope, Caucasus, Pamir, and Tian Shan
Mountains. During the last glacial maximum ~ 16,000 years
ago, populations of M. rubra no doubt extended into much
lower latitudes. But with the latest interglacial warming, pop-
ulations retreated to higher latitudes or higher elevations.
In contrast to its very broad native range, M. rubra has
a very limited distribution in North America. The few pub-
lished records of M. rubra from Greenland, Nunavut, La-
brador, and Alaska are probably based on misidentifica-
tions. The remaining M. rubra records in the US and Cana-
da come from a fairly narrow band of latitude (Fig. 4), from
Woods Hole, Massachusetts (41.5° N) and Newport, Rhode
Island (41.5° N) in the south to Quebec City, Quebec
(46.8° N) and Portage, Prince Edward Island (46.7° N) in
the north. Given the enormous range of this species in the
Palaearctic and how often it has been found in cargo from
Europe, perhaps the most striking aspect about M. rubra
in North America is how little it has spread over the past
One possible explanation may be that the North Ame-
rican populations of M. rubra have a fairly narrow range
of climatic tolerances. In Europe, different M. rubra popu-
lations show physiological adaptations to the local climate
(ELMES & al. 1999). Thus, although the species as a whole
is able to live under a great diversity of climatic conditions
in Europe and Asia, any one population has a much nar-
rower tolerance range. It is possible that the populations
of M. rubra now in North America are descended from
colonists from a narrow geographic area or even a single
locale and are not adapted to spread much beyond their cur-
rent latitudinal range in the New World. Most M. rubra
records in North America are from the past ten years, sug-
gesting that the North American populations are expand-
ing. While it is not clear what the latitudinal limits are, there
appears to be no geographic barriers that would prevent M.
rubra from spreading across the US and Canada from the
Atlantic to Pacific coasts.
Impact: In addition to its sting, M. rubra may become
a crop pest through tending plant-feeding aphids. SEIFERT
(1988) wrote that M. rubra "tends aphids more frequently
than other members of the genus." In some areas (e.g.,
along rivers and streams in Carpathian Mountains), M. ru-
bra forms huge supercolonies consisting of hundreds of in-
terconnected nests with hundred of thousands (or even mil-
lions) of workers. In these areas, population densities of M.
rubra may become very high (A.G. Radchenko, unpubl.).
STURTEVANT (1931) wrote of M. laevinodis bruesi (= M.
rubra) in Woods Hole, Massachusetts, where M. rubra was
first found in 1900: "This form was described from speci-
mens taken in the woods adjoining the Fay Rose Gardens.
It is now the dominant ant in these woods, but I have been
unable to find a single specimen in any other place. The
numerous nests seem to represent branches of a single fam-
ily, since transfers of workers never lead to fighting in spite
of the fact that the species is very pugnacious and has the
most painful sting of any ant I have encountered in the
northeastern states." GARNAS & al. (2007), however, did
not find large-scale supercolonies in the M. rubra popula-
tions of Maine.
In the native range of M. rubra, a complex of co-
evolved competitors and natural enemies may keep most
M. rubra populations in check. For example, SEIFERT (1988)
wrote that M. rubra and M. ruginodis "have a high ecologi-
cal similarity or overlap of their fundamental niches (63%)
but their real habitat overlap is 12.9% only ... which indi-
cates a strong competitive displacement."
In North America, the absence of co-evolved competi-
tors and natural enemies may allow M. rubra populations
to expand to levels where they become significant pests.
Reports of M. rubra as a serious pest in North America are
all recent. SPIERING (2009) wrote that at the Tifft Nature
Preserve in Buffalo, New York, M. rubra "were discov-
ered on the preserve in the mid-1980’s and soon became
such a serious pest that sections of the preserve needed to
be closed to the public at times." The earliest record of M.
rubra from New Brunswick dates to 1998, but now re-
ports of this species as a pest come from all over the pro-
vince (TOAL 2008). Recently there have been enormous
outbreaks of M. rubra in coastal Maine, particularly on
Mount Desert Island (GRODEN & al. 2005).
Why is M. rubra suddenly emerging as a pest after more
than a century of residence? One possibility is that a new
strain of M. rubra has been recently introduced to North
America. If the M. rubra population found in Woods Hole,
Massachusetts in 1900 still persists, it would be interesting
to examine whether or not it genetically matches popula-
tions in other parts of North America. In any event, it seems
likely that if M. rubra populations in North America con-
tinue expanding, this species will become a familiar sting-
ing pest ant in temperate North America.
We thank M. Wetterer and A. Wetterer for comments on
this manuscript; S. Cover (MCZ) and M. Deyrup (ABS)
for help with their respective ant collections; H. Zettel, S.
Tschesnokova, R. Schultz, J. Huber, A. Francoeur, G. El-
mes, and M. Bustos for supplying unpublished records; W.
O'Brien for GIS help; D.P. Wojcik and S.D. Porter for com-
piling their valuable FORMIS bibliography; R. Pasos and
W. Howerton of the FAU library for processing so many
interlibrary loans; and Florida Atlantic University for finan-
Die Ameise Myrmica rubra (LINNAEUS, 1758) (früher Myr-
mica laevinodis NYLANDER, 1846) ist eine aggressive eura-
sische Art mit mächtigem Stich, die sich derzeit im tem-
peraten Nordamerika ausbreitet. Hier wird der Trivialname
"ruby ant" vorgeschlagen. Um die weltweite Verbreitung
von M. rubra zu dokumentieren und ihr Potenzial zur wei-
teren Ausbreitung abzuschätzen, haben wir veröffentlichte
und unveröffentlichte Nachweise von > 2000 Fundorten
zusammengetragen. Wir berichten von den frühesten be-
kannten Nachweisen der Art für 71 geographische Gebiete
(Länder, große Inseln, US-Bundesstaaten, kanadische Pro-
vinzen und russische Föderationskreise), einschließlich drei-
er Gebiete ohne bisher veröffentlichte Nachweise: Prince
Edward Island, der Staat Washington und der russische
Föderationskreis Ferner Osten. Bei allen veröffentlichten
Nachweisen von M. rubra aus Ostasien, einschließlich Ja-
pans, Chinas und des östlichsten Teils Russlands, scheint
es sich um Fehlbestimmungen von Myrmica kotokui FO-
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