Worldwide spread of the ghost ant, Tapinoma melanocephalum (Hymenoptera: Formicidae)

Abstract

The ghost ant, Tapinoma melanocephalum (FABRICIUS, 1793), is a ubiquitous indoor and outdoor pest throughout much of the tropics and subtropics, and an increasingly common indoor pest in temperate regions. Here, I examine the taxonomy of T. melanocephalum and map records from > 1500 sites to evaluate its worldwide spread and probable geographic origin. I document the earliest known T. melanocephalum records for 154 geographic areas (countries, major islands, island groups, US states, and Canadian provinces), including many areas for which I found no previously published records: and Wisconsin. Tapinoma melanocephalum has one of the widest distributions known for any ant species. It has spread across the Old World and New World in both the northern and southern hemispheres, though at latitudes greater than 30° it is largely restricted to living inside buildings.

Full text

Myrmecological News 12 23-33 Vienna, September 2009
Worldwide spread of the ghost ant, Tapinoma melanocephalum (Hymenoptera: For-
micidae)
James K. WETTERER
Abstract
The ghost ant, Tapinoma melanocephalum (FABRICIUS, 1793), is a ubiquitous indoor and outdoor pest throughout much of
the tropics and subtropics, and an increasingly common indoor pest in temperate regions. Here, I examine the taxonomy of
T. melanocephalum and map records from > 1500 sites to evaluate its worldwide spread and probable geographic origin.
I document the earliest known T. melanocephalum records for 154 geographic areas (countries, major islands, island
groups, US states, and Canadian provinces), including many areas for which I found no previously published records:
Anguilla, Antigua & Barbuda, Barbados, Bonaire, Cambodia, Cape Verde, Cayman Islands, Comoro Islands, Curaçao,
Gabon, Gambia, Grenada, Italy, Kenya, Montserrat, Nevis, Nigeria, Pakistan, Romania, St. Kitts, St. Lucia, St. Martin,
Illinois, Michigan, Minnesota, Missouri, North Carolina, Virginia, and Wisconsin. Tapinoma melanocephalum has one
of the widest distributions known for any ant species. It has spread across the Old World and New World in both the
northern and southern hemispheres, though at latitudes greater than 30° it is largely restricted to living inside buildings.
In most parts of the world, T. melanocephalum can be reliably distinguished from other ant species. In India, Southeast
Asia, and the western Pacific region, however, T. melanocephalum may be confused with closely related taxa, including
Tapinoma minutum MAYR, 1862, Tapinoma indicum FOREL, 1895, and several seemingly intermediate forms, suggest-
ing that T. melanocephalum originated in the Indo-Pacific region. Future research on the phylogeny of Tapinoma and
on the genetic diversity of T. melanocephalum populations in different parts of the world should help elucidate further
the native and exotic ranges of T. melanocephalum.
Key words: Biogeography, biological invasion, exotic species, Formicidae, invasive species.
Myrmecol. News 12: 23-33 (online 24 October 2008)
ISSN 1994-4136 (print), ISSN 1997-3500 (online)
Received 11 June 2008; revision received 12 July 2008; accepted 15 July 2008
Prof. Dr. James K. Wetterer, Wilkes Honors College, Florida Atlantic University, 5353 Parkside Drive, Jupiter, FL 33458,
USA. E-mail: wetterer@fau.edu
Introduction
Tramp ants associate with humans and are spread by hu-
man commerce. Typically, tramp ants thrive only in dis-
turbed environments and do not penetrate intact natural
habitats. But as humans and their disturbance spread, so do
tramp ants. EMERY (1893a) evaluated the geographic ori-
gins of several major tramp ants using different lines of
evidence. EMERY (1893a) proposed the pharaoh ant (Mono-
morium pharaonis (LINNAEUS, 1758)) originated in East
Asia because "in collections from the East Indies, M. pha-
raonis almost always is represented, but in South Ameri-
can and African collections, it usually is not. Indeed, I have
received such specimens from Neotropical and African
regions almost exclusively from coastal areas and islands,
indicating a more recent introduction." EMERY (1893a)
proposed that the big-headed ant (Pheidole megacephala
(FABRICIUS, 1793)) probably came from Africa, "where
its closest relatives live." EMERY (1893a), however, wrote
that for two species, the longhorn crazy ant (Paratrechina
longicornis (LATREILLE, 1802)) and the ghost ant (Tapi-
noma melanocephalum (FABRICIUS, 1793)) "that have al-
ready become cosmopolitan in the tropics, I am not in the
osition to assign an original homeland." WETTERER (2008)
concluded the distributions of three closely related spe-
cies offered good evidence that P. longicornis is native to
Southeast Asia and Melanesia. Here, I evaluate the world-
wide spread and possible geographic origin of T. melano-
cephalum.
p
Tapinoma melanocephalum is a ubiquitous pest through
much of the tropics and subtropics. As LONGINO (2006)
aptly wrote: "regardless of whether you are in Guinea, New
Guinea, or Guyana, if you are sitting at a table with a sugar
dispenser you are likely to see workers of T. melanoce-
phalum running about on the surface." Workers are small
(~ 1.5 mm) and their pale legs and abdomens often blend
into the background, making them difficult to see. Their
dark brown heads and thoraces often look like hovering
specks, unrecognizable as ants. The barely visible "ghostly"
appearance of T. melanocephalum no doubt explains its
common name.
WILSON & TAYLOR (1967) wrote of T. melanocepha-
lum: "the origin of this ubiquitous tramp species is un-
known. Related species are native to various parts of Af-
rica, southeastern Asia, and the New World subtropics
and tropics." FOWLER & al. (1994) listed the assumed geo-

Fig. 1: Worldwide distribution of Tapinoma melanocephalum (some Indo-Pacific records may be misidentifications of
related species).
graphic origin of T. melanocephalum to be tropical Africa.
DLUSSKY (1994) believed T. melanocephalum to be of Asi-
an origin, and in previous papers I have followed DLUSS-
KY (1994) in calling it an Asian species (e.g., WETTERER
2002, WETTERER & VARGO 2003).
When evaluating the native and exotic ranges of a spe-
cies, researchers may consider a spectrum of distributional,
historical, evolutionary, ecological, and genetic information
(see WETTERER 2008). Evidence considered indicative of
a species' native range includes (1) older records largely
confined to a single continuous region, (2) occurrence in
inland native communities, (3) high genetic diversity, (4)
co-occurrence of species-specific symbionts, and (5) proxi-
mity to the ranges of closely related species. In contrast,
evidence indicative of a species' exotic range includes (I)
sudden appearance and spread of the species through an
area discontinuous with other known populations, (II) oc-
currence exclusively in coastal and highly disturbed envi-
ronments, (III) low genetic diversity due to a founder ef-
fect, (IV) absence of species-specific symbionts, and (V)
geographic isolation from closely related species.
Methods
I documented the range of T. melanocephalum using both
published and unpublished records. Unlike many other
major pest ant species, T. melanocephalum is easy to
identify correctly (except in the Indo-Pacific; see below),
and therefore identifications in the literature are fairly re-
liable. I obtained unpublished site records from museum
specimens in the collections of the American Museum of
Natural History (AMNH), the Archbold Biological Sta-
tion (ABS), the British Natural History Museum (BMNH),
the Museum of Comparative Zoology (MCZ), the National
Museums Liverpool (NML), the Oxford University Nat-
ural History Museum (ONHM), the Smithsonian Institute's
National Museum of Natural History (SI), and the Univer-
sity of Minnesota Collection (UMC). In addition, I used
on-line databases with specimen records from the Austra-
lian National Insect Collection (ANIC), California Academy
of Science (CAS), and Essig Museum at UC Berkeley
(UCB). I also received unpublished site records from J.
Cook (Illinois, Texas), G. Dlussky (Samoa), X. Espadaler
(France), A. Francoeur (Brazil, Colombia, Cuba, Mexico,
Quebec), B. Guénard (North Carolina, Quebec), G. Heller
(Canary Islands, Germany), J. LaForest (Michigan), M. Lush
(Australia, Gambia), P. Mattis (Florida), P. Pellitteri (Wis-
consin), F. Rigato (Italy), and B. Taylor (Nigeria). Finally,
I collected T. melanocephalum specimens on numerous Pa-
cific, Atlantic, and Caribbean Islands, and in Arizona and
Florida (e.g., WETTERER & al. 1999, WETTERER 2002,
WETTERER & O'HARA 2002, WETTERER & VARGO 2003).
I obtained geographic coordinates for sites from pub-
lished references, from specimen labels, or I looked them
up. For older references and specimens, some site names
are no longer in use and I searched, not always success-
fully, to determine current names. If a site record listed a
geographic region rather than a "point locale," and I had
no other record for this region, I used the coordinates of
the largest town within the region or, in the case of small
islands and natural areas, the center of the region. If a
published source had many sites less than 10 - 20 km
apart (e.g., HUDDLESTON & FLUKER 1968), I did not al-
ways plot every site. I did not map records of T. melano-
cephalum on boats or intercepted in transit by quarantine
inspectors. An undated specimen collected by Theodore
Pergande in Antigua must have been collected before Per-
gande's death in 1916.
Results
Published and unpublished specimen records of T. melano-
cephalum came from > 1500 sites in 154 different geo-
graphic areas (i.e., countries, island groups, major islands,
US states, and Canadian provinces), including many areas
for which I found no previously published records, e.g.,
Anguilla, Antigua & Barbuda, Barbados, Bonaire, Cam-
bodia, Cape Verde, Cayman Islands, Comoro Islands, Cura-
24

Tab. 1: Earliest known records for Tapinoma melanocepha-
lum from Asia, the Middle East, and neighboring islands.
Unpublished records include collector, museum source,
and site. SI = Smithsonian Institute, MCZ = Museum of
Comparative Zoology, + = no known published record.
Some records from India and Southeast Asia may be mis-
identifications of related species.
≤ Earliest record
+ India ≤ 1851 (JERDON 1851)
+ Singapore ≤ 1857 (SMITH 1857)
+ Indonesia ≤ 1860 (SMITH 1860)
+ Philippines ≤ 1890 (EMERY 1893d)
+ Malaysia ≤ 1892 (EMERY 1893e)
+ Sri Lanka ≤ 1892 (EMERY 1893c)
+ Papua New Guinea ≤ 1897 (EMERY 1897)
+ Burma/Myanmar ≤ 1913 (WHEELER 1913a)
+ Iraq ≤ 1918 (DONISTHORPE 1918)
+ Vietnam ≤ 1920 (SANTSCHI 1920)
+ China ≤ 1921 (WHEELER 1921)
+ Asian Russia ≤ 1926 (KARAWAJEW 1926)
+ Japan ≤ 1927 (TERANISHI 1927
≤ in ONOYAMA 1980)
+ Taiwan ≤ 1929 (WHEELER 1929)
+ Christmas Island ≤ 1934 (DONISTHORPE 1935)
+ Yemen ≤ 1955 (E.O. Wilson, MCZ): Aden
+ Cambodia ≤ 1962 (J.L. Nickel, SI): Siem Reap,
≤ Kok Patry Station
+ Saudi Arabia ≤ 1978 (COLLINGWOOD 1985)
+ Oman ≤ 1984 (COLLINGWOOD 1985)
+ Kuwait ≤ 1996 (COLLINGWOOD & AGOSTI 1996)
+ UAE ≤ 1997 (COLLINGWOOD & al. 1997)
+ Thailand ≤ 2001 (MEKLOY 2001)
+ South Korea ≤ 2005 (ANONYMOUS 2005)
+ Bangladesh ≤ 2005 (HANNAN 2007)
+ Pakistan ≤ 2007 (S. & Z. Valliani, MCZ):
≤ Karachi
çao, Gabon, Gambia, Grenada, Italy, Kenya, Montserrat,
Nevis, Nigeria, Pakistan, Romania, St. Kitts, St. Lucia, St.
Martin, Illinois, Michigan, Minnesota, Missouri, North
Carolina, Virginia, and Wisconsin (Fig. 1, Tabs. 1 - 7).
Tab. 2: Earliest known records for Tapinoma melanocepha-
lum from Australia and Oceania. Some records from the
western Pacific may be misidentifications of related spe-
cies. Abbreviations as in Table 1.
≤ Earliest record
Samoa ≤ 1876 (MAYR 1876)
Tonga ≤ 1876 (MAYR 1876)
Hawaii ≤ 1887 (BLACKBURN & CAMERON
≤ 1887)
Society Islands ≤ 1906-07 (WILSON & TAYLOR 1967)
New Caledonia ≤ 1914 (EMERY 1914)
Fiji ≤ 1915 (MANN 1921)
Banaba ≤ 1923 (TIMBERLAKE 1926)
Tokelau Islands ≤ 1924 (WILSON & TAYLOR 1967)
Marquesas Islands ≤ 1925 (CHEESMAN & CRAWLEY 1928)
Vanuatu ≤ 1925 (SANTSCHI 1928b)
Australia ≤ 1932 (Darlington, MCZ): Lankelly
≤ Creek
Pitcairn Islands ≤ 1934 (WHEELER 1936)
Tuamotu Islands ≤ 1934 (WHEELER 1936)
Austral Islands ≤ 1934 (WHEELER 1936)
Line Islands ≤ 1934 (WHEELER 1936)
Cook Islands ≤ 1935 (WHEELER 1935)
Solomon Islands ≤ 1935 (WHEELER 1935)
FS Micronesia ≤ 1937 (T. Esaki, MCZ): Malem &
≤ Kolonia
Phoenix Islands ≤ 1941 (VAN ZWALUWENBURG 1943)
Mariana Islands ≤ 1945 (D.J. Borror, SI): Agrihan
Marshall Islands ≤ 1947 (COLE 1949)
Palau ≤ 1952 (Gressitt, MCZ): Babeldaup,
≤ Ngaremeskang
Gilbert Islands ≤ 1957 (CLOUSE 2007)
New Zealand ≤ 1959 (TAYLOR 1961)
Wake Island ≤ 1959 (Y. Oshiro, MCZ): site un-
≤ known
Niue ≤ 1964 (TAYLOR 1967)
Wallis & Futuna ≤ 1965 (WILSON & HUNT 1967)
US Pacific Territories ≤ 2005 (HANDLER & al. 2007)
25

Tab. 3: Earliest known records for Tapinoma melanocepha-
lum from Africa and neighboring islands. Abbreviations as
in Table 1, and MP = Museum Paris.
≤ Earliest record
+ Madagascar ≤ 1893 (Grandidier, MP): Toamasina
+ Réunion ≤ 1895 (FOREL 1895)
+ Seychelles ≤ 1897 (FOREL 1897)
+ Guinea ≤ 1914 (SANTSCHI 1914)
+ Somalia ≤ 1926 (MENOZZI 1930)
+ Equatorial Guinea ≤ 1940 (MENOZZI 1942)
+ Mauritius ≤ 1949 (MAMET 1954)
+ Tanzania ≤ 1953 (WAY 1953)
+ Saint Helena ≤ 1967 (TAYLOR 1976)
+ Ghana ≤ 1970 (ROOM 1971)
+ Kenya ≤ 1973-74 (Allen & Brooks, MCZ):
≤ Mombasa
+ Gabon ≤ 1974 (W. Gotwald, MCZ): Makokou
+ Nigeria ≤ 1976 (B. Taylor, pers. comm.):
≤ Gambari
+ Sierra Leone ≤ 1976 (GRIFFITHS 1980)
+ Canary Islands ≤ 1989 (ESPADALER 2007)
+ Cameroon ≤ 1991 (DEJEAN & al. 1994)
+ Comoro Islands ≤ 1994 (Roger, MCZ): Dzaoudi &
≤ Moroni
+ Cape Verde ≤ 2003 (J.K. Wetterer, unpubl.):
≤ Santa Maria
+ Gambia ≤ 2007 (M. Lush, pers. comm.): Kololi
In both the Old and New World tropics, T. melano-
cephalum has been found in most maritime countries (Fig.
1), including all tropical countries of Oceania except Nauru
and Tuvalu, and all island groups of the West Indies ex-
cept the Turks and Caicos Islands. In tropical Asia, T.
melanocephalum has been recorded from all countries ex-
cept Brunei and Laos. In continental Africa, however,
there have been relatively few records of this ant (Fig. 1).
In the continental US, most records of T. melanocepha-
lum were from Florida. Although the majority of records
came from disturbed habitats, some were from relatively
intact habitats. For example, I collected T. melanocephalum
nesting in a cypress tree in a relatively natural marsh area
of Loxahatchee National Wildlife Refuge in Palm Beach
County, Florida.
At latitudes greater than 30°, T. melanocephalum is
largely restricted to living inside buildings. The highest lati-
tude records came from indoor sites in Scandinavia (up to
65.0° N; SORVARI 2003), Scotland (up to 55.9° N; GOD-
Tab. 4: Earliest known records for Tapinoma melanocepha-
lum from South and Central America and neighboring is-
lands. Abbreviations as in Table 1, and BMNH = Natural
History Museum in London.
≤ Earliest record
French Guiana ≤ 1793 (FABRICIUS 1793)
Brazil ≤ 1881 (FOREL 1881)
Galapagos ≤ 1892 (EMERY 1893b)
Mexico ≤ 1894 (PERGANDE 1896)
Belize ≤ 1905-06 (WHEELER 1907)
Costa Rica ≤ 1908 (FOREL 1908)
Guyana ≤ 1914 (G.E. Bokin, BMNH): Georgetown
Panama ≤ 1918 (no collector data, SI): Ancon
Venezuela ≤ 1936 (WEBER 1948)
Colombia ≤ 1939 (Gallego, SI): Medellin Valley
Nicaragua ≤ 1943 (Woke, SI): Chinandega
Surinam ≤ 1959 (KEMPF 1961)
Peru ≤ 1964 (PARDO 1964)
Honduras ≤ 1972 (KEMPF 1972)
Paraguay ≤ 1997 (A. Wild, pers. comm.): Asunción
FREY 1907), and Russia (up to 55.8° N; KUNASHEV & NI-
YAZOVA 1998). In North America, the highest latitude re-
cords come from indoor sites in Manitoba (49.9° N; AYRE
1977), Quebec (45.5° N; B. Guénard, pers. comm.), Oregon
(45.5° N; NUGENT & al. 2005), Minnesota (45.0° N; P.
Clausen, pers. comm.), Wisconsin (up to 43.6° N; P. Pel-
litteri, pers. comm.), Michigan (43.6° N; J. LaForest, pers.
comm.), and Maine (43.4° N; DEARBORN & GRANGER
2000). In the southern hemisphere, the highest latitude
records came from sites in New Zealand (up to 37.6° S;
TAYLOR 1961).
Nest symbionts
Although T. melanocephalum workers readily tend a wide
range of Hemiptera, I found only one record of a pur-
ported species-specific symbiont of T. melanocephalum.
SHEPARD & GIBSON (1972) reported salticid spiders of
the genus Cotinusa present in 61% of T. melanocephalum
nests on the Osa Peninsula, Costa Rica "where these ants
build their nests on the lower surface of leaves in the lower
forest canopy." I believe, however, that the ants were mis-
identified (see Discussion).
Related Tapinoma
In the Indo-Pacific, T. melanocephalum may be confused
with a number of closely related taxa, including Tapinoma
minutum MAYR, 1862 and Tapinoma indicum FOREL, 1895.
Tapinoma minutum is brown with a somewhat darker ab-
domen; Tapinoma indicum is very similar to T. minutum,
26

Tab. 5: Earliest known records for Tapinoma melanocepha-
lum from the West Indies. Abbreviations as in Table 1.
≤ Earliest record
+ Virgin Islands ≤ 1881 (FOREL 1881)
+ Jamaica ≤ 1891 (Pergande collection, SI): Kingston
+ St Vincent ≤ 1893 (FOREL 1893)
+ Puerto Rico ≤ 1899 (Busck, SI): San Juan
+ Bahamas ≤ 1904 (WHEELER 1905)
+ Trinidad &
+ Tobago
≤ 1910 (WHEELER 1916)
+ Haiti ≤ 1912-13 (WHEELER & MANN 1914)
+ Dominica ≤ 1913 (WHEELER 1913b)
+ Cuba ≤ 1913 (WHEELER 1913b)
+ Antigua &
+ Barbuda
≤ 1916 (Pergande collection, SI): Antigua
+ Dominican
+ Republic
≤ 1930 (MENOZZI & RUSSO 1930)
+ Nevis ≤ 1934 (N.A. Weber, MCZ): site unknown
+ St Lucia ≤ no date (N.A. Weber, MCZ): site un-
≤ known
+ Montserrat ≤ 1991 (J.P.E.C. Darlington, MCZ): Salem
+ Aruba ≤ 1994 (JAFFE & LATTKE 1994)
+ Martinique ≤ 1994 (JAFFE & LATTKE 1994)
+ Guadeloupe ≤ 1994 (JAFFE & LATTKE 1994)
+ Barbados ≤ 1998 (S. Cover & E.O. Wilson, MCZ):
≤ Graeme Hall Swamp
+ Grenada ≤ 2003 (J.K. Wetterer, MCZ): St George's
+ Curaçao ≤ 2004 (J.K. Wetterer, MCZ): Piscadera
+ Anguilla ≤ 2006 (J.K. Wetterer, MCZ): Meads Bay
+ St Martin ≤ 2006 (J.K. Wetterer, MCZ): Baie Nettle
+ St. Kitts ≤ 2007 (J.K. Wetterer, MCZ): Bird Rock
+ Cayman
+ Islands
≤ 2008 (J.K. Wetterer, MCZ): Grand
≤ Cayman; Websters
+ Bonaire ≤ 2008 (J.K. Wetterer, MCZ): Barkadera
but with lighter color and longer antennae (though with
shorter antennae than those of T. melanocephalum).
FOREL (1913) described Tapinoma melanocephalum
var. malesianum FOREL, 1913 from sites in Sumatra and
Java, as a variety with more uniform color, thinner palps,
and shorter antennae than the type. Although this remains
a valid subspecies of T. melanocephalum, its description is
very similar to that of T. indicum. SANTSCHI (1928a) de-
Tab. 6: Earliest known records for Tapinoma melanocepha-
lum from Europe. Abbreviations as in Table 1, and BMNH
= Natural History Museum in London.
≤ Earliest record
+ England ≤ 1886 (BILLUPS 1887)
+ Scotland ≤ 1904 (GODFREY 1907)
+ Germany ≤ 1926 (Dorn, BMNH): Leipzig Zoo
+ Scandinavia ≤ 1979 (COLLINGWOOD 1979)
+ Finland ≤ 1991 (SORVARI 2003)
+ Sweden ≤ 1995 (DOUWES 1995)
+ France ≤ 1995 (HUGEL & al. 2003)
+ Switzerland ≤ 1997 (DORN & al. 1997)
+ European Russia ≤ 1997 (KUNASHEV & NIYAZOVA 1998)
+ Spain ≤ 1999 (ESPADALER & ESPEJO 2002)
+ Austria ≤ 2001 (STEINER & al. 2003)
+ Denmark ≤ 2002 (JESPERSEN & CHRISTENSEN
≤ 2003)
+ Netherlands ≤ 2002 (VIERBERGEN 2003)
+ Romania ≤ 2004 (RADCHENKO 2004):
≤ site unknown
+ Norway ≤ 2006 (BIRKEMOE 2006)
+ Belgium ≤ 2006 (DEKONINCK & al. 2006)
+ Italy ≤ 2006 (F. Rigato, pers. comm.): Milan
scribed Tapinoma melanocephalum var. australis SANT-
SCHI, 1928 from several sites in Samoa, and mentioned that
Buxton also collected this variety in Vanuatu. SANTSCHI
(1928b) described Tapinoma melanocephalum var. aus-
trale SANTSCHI, 1928 based on specimens collected by
Buxton in "Noq Narbow," Vanuatu (= misreading of "Hog
Harbour," hand-written on specimen labels in BMNH and
SI), mentioning that this variety also occurred in Samoa.
Both australis and australe have a dark head and thorax
and light abdomen, like the melanocephalum type, but
have narrower palps, like those of T. melanocephalum ma-
lesianum. From the matching descriptions, it is clear that
SANTSCHI (1928a, b) described the same taxon twice, using
slightly different names: australis and australe, however,
as BOLTON (1995) pointed out "the type-localities of the
two are different and the names must be treated as sepa-
rate." Although both names were later synonymized with
T. melanocephalum (by WILSON & TAYLOR 1967 and BOL-
TON 1995, respectively), taxonomic questions remain. In
many surveys of Indo-Pacific ants, researchers were un-
able to identify satisfactorily Tapinoma specimens. For ex-
ample, ONOYAMA (1976) recorded Tapinoma sp. A in Oki-
nawa that "seems to be recorded as melanocephalum by
several authors in Japan, but does not agree with the fig-
ure and description of melanocephalum by WILSON & TAY-
27

Tab. 7: Earliest known records for Tapinoma melanocepha-
lum from the United States and Canada. Abbreviations as in
Table 1, and UMC = University of Minnesota Collection.
≤ Earliest record
+ Washington DC ≤ 1894 (PERGANDE 1896)
+ New York ≤ 1907 (WHEELER 1907)
+ Florida ≤ 1930 (SMITH 1930)
+ Virginia ≤ 1932 (H.T. Vanderford, SI): Norfolk
+ Minnesota ≤ 1935 (D.G. Denning, UMC): Min-
≤ neapolis
+ Quebec ≤ 1939 (E.R. Bellemare, A Francoeur
≤ pers. comm.): Montréal
+ Iowa ≤ 1947 (KING 1948)
+ Georgia ≤ 1951 (SMITH 1951)
+ New Mexico ≤ 1967 (Elson, SI): Cloudcroft
+ Manitoba ≤ 1977 (AYRE 1977)
+ California ≤ 1979 (SMITH 1979)
+ Ontario ≤ 1979 (SMITH 1979)
+ Connecticut ≤ 1979 (FROST 1979)
+ Kansas ≤ 1980 (DUBOIS & DANOFF-BURG 1994)
+ Texas ≤ 1994 (COOK & al. 1994)
+ Missouri ≤ 1995 (J. Trager, pers. comm.): Chest-
≤ erfield
+ Arizona ≤ 1996 (WETTERER & al. 1999)
+ Illinois ≤ 1997 (J.L. Cook, pers. comm.): Chicago
+ Louisiana ≤ 1999 (DASH 2004)
+ Maine ≤ 1999 (DEARBORN & GRANGER 2000)
+ Washington ≤ 2000 (HEDGES 2000)
+ Wisconsin ≤ 2000 (P. Pellitteri, pers. comm.): Wis-
≤ consin Dells
+ Michigan ≤ 2004 (J. LaForest, pers. comm.): Mid-
≤ land; Dow Gardens
+ Oregon ≤ 2005 (NUGENT & al. 2005)
+ North Carolina ≤ 2007 (B. Guénard; pers. comm.): Ra-
≤ leigh
LOR (1967)." ONOYAMA (1976) wrote that the decisive dif-
ference was that Tapinoma sp. A has shorter antennae, with
the scape extending beyond the occiput by about half the
length of the first funicular segment, rather than by more
than the length of the first two funicular segments as in
T. melanocephalum. CLOUSE (2007) found that in Micro-
nesia, most Tapinoma specimens could be unambiguously
assigned either to T. melanocephalum or to T. minutum.
However, some specimens were intermediate, with anten-
nae like T. melanocephalum but color like T. minutum or
the reverse, which CLOUSE (2007) referred to as Tapinoma
A and Tapinoma B, respectively. Tapinoma B may or may
not correspond to T. melanocephalum australis/australe
and ONOYAMA's (1976) Tapinoma sp. A.
FOREL (1908) described the only other valid subspecies,
T. melanocephalum coronatum FOREL, 1908 from Costa
Rica, reporting that this subspecies also occurred in Ja-
maica and "etc." This taxon has the same coloration and
antenna length as the type, but with shorter and narrower
palps, no indentation in the thorax, and less shiny head
and thorax.
Discussion
Tapinoma melanocephalum has one of the widest distri-
butions known for any ant species. It has spread across
the Old World and New World in both the northern and
southern hemispheres. It is tropical and subtropical; at lati-
tudes greater than 30°, it is largely restricted to living in-
side buildings. The only ants with broader known distri-
butions than T. melanocephalum are the longhorn crazy
ant, Paratrechina longicornis (WETTERER 2008), and the
pharaoh ant, Monomorium pharaonis (L.) (J.K. Wetterer,
unpubl.). I expect that T. melanocephalum will eventually
spread to the disturbed lowlands of virtually every trop-
ical country. This species also appears to be an increas-
ingly common indoor pest in temperate areas, and some-
day may be found in every major city of the world. Be-
cause T. melanocephalum can live anywhere that humans
live, there is no limit to the latitude where indoor popula-
tions of this species may be found.
Tapinoma melanocephalum is particularly noted as a
pest in homes, restaurants, hospitals, and greenhouses. For
example, T. melanocephalum was the most common house
ant in Brazil (e.g., KAMURA & al. 2007) and on Panang
Island, Malaysia (LEE & al. 2002), and it was the most fre-
quently reported pest ant in greater Miami (KLOTZ & al.
1995). WHEELER (1907) noted that T. melanocephalum
had been "introduced into well-heated buildings in New
York," and observed workers in the winter "visiting the
sugar bowls on the tables of the Columbia University com-
mons." KARAWAJEW (1926) reported T. melanocephalum
from hotel tables both on Ambon Island, Indonesia and
in Vladivostok, Russia. James Trager (pers. comm.) ob-
served T. melanocephalum inside the Sachs Butterfly House
in Chesterfield, Missouri, beginning shortly after it opened
in 1995. Tapinoma melanocephalum also has been a long-
term problem in the greenhouses of the Dow Estate in Mid-
land, Michigan, where fumigation is incompatible with a
butterfly exhibit (J. LaForest, pers. comm.).
Although T. melanocephalum does not sting and its bite
is not very painful (pers. obs.), it can be a serious pest. For
example, FOWLER & al. (1993) and MOREIRA & al. (2005)
found that T. melanocephalum was the most prevalent ant
in hospitals in Brazil, and had the highest potential as a
vector for pathogens. In fact, MOREIRA & al. (2005) found
at least 14 different types of bacteria on T. melanocephalum
workers collected in the hospitals, including antibiotic re-
sistant strains. Tapinoma melanocephalum also can be an
agricultural pest because it tends populations of phloem-
feeding Hemiptera, such as mealybugs, scale insects, and
28

aphids, for their honeydew (VENKATARAMAIAH & REHMAN
1989). Hemiptera cause damage by sapping plants of nu-
trients and increasing the occurrence of diseases, including
viral and fungal infections. FOWLER & al. (1994) concluded
that T. melanocephalum had a great impact on other ant
species on banana plantations in Brazil. Under some con-
ditions, T. melanocephalum may be beneficial as a bio-
control agent. For example, OSBORNE & al. (1995) found
that in greenhouses of central Florida, naturally occurring
populations of T. melanocephalum were important in con-
trolling a serious plant pest, the two-spotted spider mite,
Tetranychus urticae KOCH, 1836.
Whereas T. melanocephalum is most prevalent in dis-
turbed habitats, it has also been reported from natural re-
serves around the world, including the Baiyer River Wild-
life Sanctuary in Papua-New Guinea (WHALEN & MAC-
KAY 1988), Reserva Biológica de Una in Brazil (DELABIE
& al. 1997), primary rainforest in Chocó, Colombia (ARM-
BRECHT & al. 2001), and on many Galapagos islands (CAU-
STEN & al. 2006). In the Wapoga River Area of Irian Jaya,
Indonesia, SNELLING (2000) found that T. melanocephalum
"had successfully invaded forest areas where they appa-
rently became quickly assimilated into the native mix; their
presence did not seem to adversely affect other ant spe-
cies there. This apparently benign situation probably would
be reversed in the presence of continued disturbance of
the habitat."
Tapinoma melanocephalum colonies are polygynous
and frequently relocate their nests. In a matter of hours,
colonies can move into cargo, which is then shipped world-
wide. In fact, LESTER (2005) reported that T. melanocepha-
lum was the second most common ant species (after Phei-
dole megacephala) intercepted in freight entering New Zea-
land between 1997 and 2002. WEBER (1939) reported col-
lecting T. melanocephalum from riverboats in Venezuela
and Guyana and on a ship between Haiti and New York.
Tapinoma melanocephalum was found on a train in France,
traveling from Paris to Rennes (X. Espadaler, pers. comm.).
In Le Carbet, Martinique, a T. melanocephalum colony
moved overnight into luggage packed for departure, requir-
ing me to shake out everything before repacking.
Many of the most recent first records reported here
come from the West Indies, but this is likely an artifact of
poor past collection records. For example, PERGANDE (1896)
wrote that T. melanocephalum was "common on all of
the West Indian islands," though at the time the only pub-
lished West Indian records were from St Thomas and St
Vincent (Tab. 5). In contrast, the many first records of T.
melanocephalum from well-studied parts of Europe and
North America dating from the past 15 years (Tabs. 6, 7)
suggests a recent spread in these areas, perhaps as a re-
sult of the increasing prevalence of air commerce.
Native and exotic range
The general consensus holds that T. melanocephalum is
from the Old World tropics. The earliest T. melanoce-
phalum record from continental Africa (SANTSCHI 1914;
Tab. 3) was reported long after this species was already
known from many sites in Asia and the Pacific (Tabs. 1, 2),
suggesting that T. melanocephalum may be a relatively
recent arrival in Africa. In addition, site records from con-
tinental Africa remain scarce (Fig. 1). In contrast, T. mela-
nocephalum has widespread records from India, South-
east Asia, and the western Pacific. The close resemblance
between T. melanocephalum and several species in this re-
gion further indicates that T. melanocephalum most pro-
bably originated in the Indo-Pacific.
A few lines of evidence appear to support the hypo-
thesis that T. melanocephalum originated in the Neotrop-
ics: (a) T. melanocephalum was first described from French
Guiana (FABRICIUS 1793), (b) the salticid spider Continusa
sp., which is the sole proposed symbiont of T. melano-
cephalum, has only been reported from Costa Rica (SHEP-
ARD & GIBSON 1972), and (c) the subspecies T. melano-
cephalum coronatum is known only from Costa Rica and
Jamaica (FOREL 1908). Closer inspection, however, calls
into question all three of these lines of evidence. First, sub-
sequent to FABRICIUS (1793), I found no other published
report of T. melanocephalum from French Guiana. Sec-
ond, SHEPARD & GIBSON (1972) found that Continusa spi-
ders were extremely common in T. melanocephalum nests
on the Osa Peninsula, but spiders have never been reported
in T. melanocephalum nests elsewhere. It is also very un-
usual that SHEPARD & GIBSON (1972) found T. melanoce-
phalum was common in a forest canopy and was nesting
on the undersides of leaves. I am confident that the ants in
this study were misidentified Tapinoma ramulorum inrec-
tum FOREL, 1908. According to LONGINO (2006), T. ramu-
lorum inrectum "has very distinctive nesting habits and
thus is more easily identified in the field than from mu-
seum specimens. It makes small carton nests on the un-
dersides of leaves." Finally, after FOREL (1908) described
T. melanocephalum coronatum, no subsequent researcher
has ever reported this taxon, despite extensive ant samp-
ling particularly in Costa Rica, calling its validity into ques-
tion.
A thorough taxonomic revision of Tapinoma would be
valuable. Future research on the phylogeny of Tapinoma
and on the genetic diversity of T. melanocephalum popu-
lations in different parts of the world should help eluci-
date further the native and exotic ranges of T. melanoce-
phalum.
Acknowledgements
I thank A. Wetterer, M. Wetterer, C.-Y. Lee, and R. Clouse
for comments on this manuscript; S. Cover for help, en-
couragement, and ant identification; B. Taylor, F. Rigato,
P. Pellitteri, P. Mattis, M. Lush, J. LaForest, B. Guénard, A.
Francoeur, X. Espadaler, G. Dlussky, and J. Cook for pro-
viding unpublished records; X. Espadaler for informing me
of published records I overlooked; B. Bolton (BMNH), J.
Carpenter (AMNH), P. Clausen (UMC), S. Cover (MCZ),
M. Deyrup (ABS), J. Hogan (ONHM), G. Knight (NML),
and T. Schultz (SI) for help with their respective ant col-
lections; W. O'Brien for GIS help; D.P. Wojcik and S.D.
Porter for compiling their valuable FORMIS bibliography;
C. Scheid and R. Pasos of the FAU library for processing so
many interlibrary loans; FAU, the National Science Foun-
dation, and the National Geographic Society for financial
support.
Zusammenfassung
Die Ameisenart Tapinoma melanocephalum (FABRICIUS,
1793), auch "ghost ant" genannt, ist ein ubiquitärer Lästling
innerhalb und außerhalb von Gebäuden in großen Teilen
der Tropen und Subtropen und zunehmend auch ein Läst-
29

ling in Gebäuden temperater Gebiete. Ich erörtere hier die
Taxonomie von T. melanocephalum und zeige Nachweise
von > 1500 Fundorten, um die weltweite Ausbreitung und
den wahrscheinlichen geographischen Ursprung der Art zu
evaluieren.
Ich habe die frühesten bekannten Nachweise von T.
melanocephalum für 154 geographische Gebiete (Länder,
große Inseln, Inselgruppen, US-Bundesstaaten und Kanadi-
sche Provinzen) zusammengetragen, einschließlich vieler,
für die ich keine bisher veröffentlichten Nachweise gefun-
den habe: Anguilla, Antigua & Barbuda, Barbados, Bonaire,
Curaçao, Gabon, Gambia, Grenada, Illinois, Italien, Kaiman-
Inseln, Kambodja, Kap Verde, Kenya, Komoren, Montser-
rat, Nevis, Nigeria, Pakistan, Rumänien, St. Kitts, St. Lucia,
St. Martin, Michigan, Minnesota, Missouri, North Caroli-
na, Virginia und Wisconsin. Tapinoma melanocephalum
hat somit eine der weitesten Verbreitungen, die man von
Ameisen kennt. Die Art hat sich über die Alte Welt und die
Neue Welt, sowohl auf der nördlichen als auch auf der süd-
lichen Hemisphäre, ausgebreitet. In Breitengraden höher
als 30° ist die Art aber im Wesentlichen auf Vorkommen
innerhalb von Gebäuden beschränkt.
Für die meisten Teile der Welt kann T. melanocephalum
verlässlich von anderen Ameisenarten unterschieden wer-
den. In Indien, Südostasien und der westlichen Pazifik-Re-
gion allerdings leben nahe verwandte Taxa, mit denen T.
melanocephalum verwechselt werden kann. Dies sind Tapi-
noma minutum MAYR, 1862, Tapinoma indicum FOREL,
1895 sowie einige scheinbar intermediäre Formen; daraus
kann geschlussfolgert werden, dass T. melanocephalum ur-
sprünglich aus der Indo-Pazifischen Region stammt. Zukünf-
tige Untersuchungen der Phylogenie von Tapinoma und der
genetischen Vielfalt von T. melanocephalum-Populationen
in verschiedenen Teilen der Welt sollten die Einteilung der
Vorkommen in native und exotische im Detail klären kön-
nen.
References
ANONYMOUS 2005: Ghost ants. – Korea Times 29 November 2005.
ARMBRECHT, I., JIMÉNEZ, E., ALVAREZ, G., ULLOA-CHACON, P. &
ARMBRECHT, H. 2001: An ant mosaic in the Colombian rain
forest of Chocó (Hymenoptera: Formicidae). – Sociobiology
37: 491-509.
AYRE, G.L. 1977: Exotic ants in Winnipeg. – Manitoba Ento-
mologist 11: 4111-4144.
BILLUPS, T.R. 1887: Tapinoma melanocephalum, FOR. – The
Entomologist 20: 184-185.
BIRKEMOE, T. 2006: "Spøkelsesmaur" (Tapinoma melanocepha-
lum), Nasjonalt folkehelseinstitutt. – Norwegian Biodiversity
Information Centre, Trondheim 1 p.
BLACKBURN, T. & CAMERON, P. 1887: On the Hymenoptera of
the Hawaiian Islands. – Memoirs of the Manchester Literary
& Philosophical Society 10: 194-244.
BOLTON, B. 1995: A new general catalogue of the ants of the
world. – Harvard University Press, Cambridge, MA, 504 pp.
CAUSTEN, C.E., PECK, S.B., SINCLAIR, B.J., ROQUE-ALBELO, L.,
HODGSON, C.J. & LANDRY, B. 2006: Alien insects: threats and
implications for conservation of Galápagos Islands. – Annals
of the Entomological Society of America 99: 121-143.
CHEESMAN, L.E. & CRAWLEY, W.C. 1928: A contribution to-
wards the insect fauna of French Oceania – Part III. Formicidae.
– Annals & Magazine of Natural History (10)2: 514-525.
CLOUSE, R.M. 2007: The ants of Micronesia (Hymenoptera: For-
micidae). – Micronesica 39: 171-295.
COLE, A.C. 1949: The ants of Bikini Atoll, Marshall Islands. –
PanPacific Entomologist 25: 172-174.
COLLINGWOOD, C.A. 1979: The Formicidae (Hymenoptera) of
Fennoscandia and Denmark. – Fauna Entomologica Scandi-
navica 8: 1-174.
COLLINGWOOD, C.A. 1985: Hymenoptera: Fam. Formicidae of
Saudi Arabia. – Fauna of Saudi Arabia 7: 230-302.
COLLINGWOOD, C.A. & AGOSTI, D. 1996: Formicidae (Insecta:
Hymenoptera) of Saudi Arabia (Part 2). – Fauna of Saudi Ara-
bia 15: 300-385.
COLLINGWOOD, C.A., TIGAR, B.J. & AGOSTI, D. 1997: Introduced
ants in the United Arab Emirates. – Journal of Arid Environ-
ments 37: 505-512.
COOK, J.L., MARTIN, J.B. & GOLD, R.E. 1994: First record of
Tapinoma melanocephalum (Hymenoptera: Formicidae) in
Texas. – Southwestern Entomologist 19: 409-410.
DASH, S.T. 2004: Species diversity and biogeography of ants
(Hymenoptera: Formicidae) in Louisiana, with notes on their ec-
ology. – MSc Thesis, Louisiana State University, Baton Rouge,
290 pp.
DEARBORN, R.G. & GRANGER, C.A. 2000: Forest and shade tree
insect and disease conditions for Maine. A summary of the
1999 situation. – Forest Health and Monitoring Division Sum-
mary Report 14: 1-68.
DEJEAN, A., AKOA, A., DJIETO-LORDON, C. & LENOIR, A. 1994:
Mosaic ant territories in an African secondary rain forest (Hy-
menoptera: Formicidae). – Sociobiology 23: 275-292.
DEKONINCK, W., WEGNEZ, P., ESPADALER, X. & GROOTAERT, P.
2006: First record of infestation of the ghost ant Tapinoma
melanocephalum (FABRICIUS, 1793) in Belgium (Hymenoptera,
Formicidae) – a new indoor pest problem for the country? –
Bulletin van de Koninklijke Belgische Vereniging voor Ento-
mologie 142: 25-28.
DELABIE, J.H.C., MARIANO, C.S.F. & DO NASCIMENTO, I.C. 1997:
Inventário preliminar das formigas (Insecta; Hymenoptera; For-
micidae) da reserva biológica e áreas conexas no município
de Una, Bahia. In: SARACURA, V.F. (Ed.): Plano de manejo da
Reserva Biológica de Una - Ba. (versão preliminar). – Pro-
grama Nacional de Meio Ambiente, Ministério do Meio Ambi-
ente, Brasília, pp. 105-111.
DLUSSKY, G.M. 1994: Zoogeography of southwestern Oceania.
In: PUZATCHENKO, Y.G., GOLOVATCH, S.I., DLUSSKY, G.M.,
DIAKONOV, K.N., ZAKHAROV, A.A. & KORGANOVA, G.A. (Eds.):
Animal population of the islands of Southwestern Oceania
(ecogeographic studies). – Nauka Publishers, Moscow, pp. 48-
93. (in Russian)
DONISTHORPE, H. 1918: A list of ants from Mesopotamia; with
a description of a new species and a new variety. – Entomolo-
gist's Record & Journal of Variation 30: 165-168.
DONISTHORPE, H. 1935: The ants of Christmas Island. – Annals
& Magazine of Natural History (10)15: 629-635.
DORN, K., LANDAU, I. & CHERIX, D. 1997: Einschleppung von
Tapinoma melanocephalum (Formicidae) in der Schweiz. –
Mitteilungen der Schweizerischen Entomologischen Gesell-
schaft 70: 242-243.
DOUWES, P. 1995: Sveriges myror. – Entomologisk Tidskrift 116:
83-99.
DUBOIS, M.B. & DANOFF-BURG, J. 1994: Distribution of ants in
Kansas: subfamilies Dolichoderinae and Formicinae (Hymeno-
ptera: Formicidae). – Sociobiology 24: 147-178.
EMERY, C. 1893a: Über die Herkunft der Pharao-Ameise. – Bio-
logisches Centralblatt 13: 435-436.
30

EMERY, C. 1893b: Notice sur quelques fourmis des îles Galapa-
gos. – Annales de la Société Entomologique de France 62:
89-92.
EMERY, C. 1893c: Voyage de M. E. Simon à l'île de Ceylan (jan-
vier-février 1892). Formicides. – Annales de la Société Ento-
mologique de France 62: 239-258.
EMERY, C. 1893d: Voyage de M. E. Simon aux îles Philippines
(mars & avril 1890). Formicides. – Annales de la Société Ento-
mologique de France 62: 259-270.
EMERY, C. 1893e: Formicides de l'Archipel Malais. – Revue
Suisse de Zoologie 1: 187-229.
EMERY, C. 1897: Viaggio di Lamberto Loria nella Papuasia orien-
tale. XVIII. Formiche raccolte nella Nuova Guinea dal Dott.
Lamberto Loria. – Annali del Museo Civico di Storia Natu-
rale 38[=(2)18]: 546-576.
EMERY, C. 1914: Les fourmis de la Nouveller-Calédonie and des
îles Loyalty In: SARASIN, F. & ROUX, J. (Eds.): Nova Caledonia.
A. zoologie. – C.W. Kreidels Verlag, Wiesbaden, pp. 393-437.
ESPADALER, X. 2007: The ants of El Hierro (Canary Islands). –
Memoirs of the American Entomological Institute 80: 113-127.
ESPADALER, X. & ESPEJO, F. 2002: Tapinoma melanocephalum
(FABRICIUS, 1793), a new exotic ant in Spain (Hymenoptera,
Formicidae). – Orsis 17: 101-104.
FABRICIUS, J.C. 1793: Entomologia systematica emendata et aucta.
Secundum classes, ordines, genera, species, adjectis synonimis,
locis observationibus, descriptionibus. Tome 2. – C.G. Proft,
Hafniae, 519 pp.
FOREL, A. 1881: Die Ameisen der Antille St. Thomas. – Mittei-
lungen des Münchner Entomologischen Vereins 5: 1-16.
FOREL, A. 1893: Formicides de l'Antille St. Vincent, récoltées par
Mons. H.H. Smith. – Transactions of the Entomological So-
ciety of London 1893: 333-418.
FOREL, A. 1895: Nouvelles fourmis de diverses provenances,
surtout d'Australie. – Annales de la Société Entomologique
de Belgique 39: 41-49.
FOREL, A. 1897: Ameisen aus Nossi-Bé, Majunga, Juan de Nova
(Madagaskar), den Aldabra-Inseln und Sansibar, gesammelt
von Herrn Dr. A. Voeltzkow aus Berlin. Mit einem Anhang
über die von Herrn Privatdocenten Dr. A. Brauer in Marburg
auf den Seychellen und von Herrn Perrot auf Ste. Marie (Ma-
dagaskar) gesammelten Ameisen. – Abhandlungen der Sen-
ckenbergischen Naturforschenden Gesellschaft 21: 185-208.
FOREL, A. 1908: Fourmis de Costa-Rica récoltées par M. Paul
Biolley. – Bulletin de la Société Vaudoise des Sciences Na-
turelles 44: 35-72.
FOREL, A. 1913: Wissenschaftliche Ergebnisse einer Forschungs-
reise nach Ostindien ausgeführt im Auftrage der Kgl. Preuss.
Akademie der Wissenschaften zu Berlin von H. v. Buttel-Ree-
pen. II. Ameisen aus Sumatra, Java, Malacca und Ceylon. Ge-
sammelt von Herrn Prof. Dr. v. Buttel-Reepen in den Jahren
1911-1912. – Zoologische Jahrbücher, Abteilung für Syste-
matik, Geographie und Biologie der Tiere 36: 1-148.
FOWLER, H.G., BUENO, O.C., SADATSUNE, T. & MONTELLI, A.C.
1993: Ants as potential vectors of pathogens in hospitals in
the state of São Paulo, Brazil. – Insect Science and Its Appli-
cation 14: 367-370.
FOWLER, H.G., SCHLINDWEIN, M.N. & MEDEIROS, M.A. 1994:
Exotic ants and community simplification in Brazil: a review
of the impact of exotic ants on native ant assemblages. In:
WILLIAMS D.F. (Ed.): Exotic ants: biology, impact, and control
of introduced species. – Westview Press, Boulder, pp. 151-162.
FROST, S.W. 1979: A preliminary study on North American in-
sects associated with elderberry flowers. – Florida Entomolo-
gist 62: 341-355.
GODFREY, R. 1907: Notes on the animal life of the hothouses of
the Royal Botanic Garden, Edinburgh. – Notes from the Roy-
al Botanic Garden, Edinburgh 4: 99-103.
GRIFFITHS, D. 1980: The feeding biology of ant-lion larvae: prey
capture, handling and utilization. – Journal of Animal Ecology
49: 99-125.
HANDLER, A.T., GRUNER, D.S., HAINES, W.P., LANGE, M.W. &
KANESHIRO, K.Y. 2007: Arthropod surveys on Palmyra Atoll,
Line Islands, and insights into the decline of the native tree
Pisonia grandis (Nyctaginaceae). – Pacific Science 61: 485-502.
HANNAN, M.A. 2007: Ant records from Savar, Dhaka, Bangla-
desh. – Asian Myrmecology 1: 101-104.
HEDGES, S.A. 2000: Coast to coast. – PCT Online 77: 12 Octo-
ber 2000, <http://pctonline.com/articles/article.asp?ID=1273
&IssueID=77>, retrieved on 28 November 2006.
HUDDLESTON, E.W. & FLUKER, S.S. 1968: Distribution of ant
species of Hawaii. – Proceedings of the Hawaiian Entomo-
logical Society 20: 45-69.
HUGEL, S., CALLOT, H. & DELECOLLE, J.C. 2003: Insectes exo-
tiques et/ou nouveaux pour la France dans les serres du Jar-
din Botanique de Strasbourg. – Bulletin de la Société entomo-
logique de Mulhouse 59: 69-73.
JAFFE, K. & LATTKE, J.E. 1994: Ant fauna of the French and
Venezuelan islands in the Caribbean. In: WILLIAMS, D.F. (Ed.):
Exotic ants: biology, impact, and control of introduced spe-
cies. – Westview Press, Boulder, pp. 181-190.
JERDON, T.C. 1851: A catalogue of the species of ants found in
Southern India. – Madras Journal of Literature & Science 17:
103-127.
JESPERSEN, J.B. & CHRISTENSEN, M. 2003: Annual report 2002.
– Danish Pest Infestation Laboratory, Kongens Lyngby, 60 pp.
KAMURA, C.M., MORINI, M.S.C., FIGUEIREDO, C.J., BUENO, O.C.
& CAMPOS-FARINHA, A.E.C. 2007: Ant communities (Hyme-
noptera: Formicidae) in an urban ecosystem near the Atlantic
Rainforest. – Brazilian Journal of Biology 67: 631-637.
KARAWAJEW, V. 1926: Ameisen aus dem Indo-Australischen Ge-
biet. – Treubia 8: 413-445.
KEMPF, W.W. 1961: A survey of the ants of the soil fauna in
Surinam (Hymenoptera: Formicidae). – Studia Entomologica
4: 481-524.
KEMPF, W.W. 1972: Catálogo abreviado das formigas da região
neotropical (Hymenoptera: Formicidae). – Studia Entomolo-
gica 15: 3-344.
KING, R.L. 1948: A tropical ant temporarily established in Iowa.
– Proceedings of the Iowa Academy of Science 55: 395.
KLOTZ, J.H., MANGOLD, J.R., VAIL, K.M., DAVIS, L.R. JR. &
PATTERSON, R.S. 1995: A survey of the urban pest ants (Hy-
menoptera: Formicidae) of peninsular Florida. – Florida Ento-
mologist 78: 109-118.
KUNASHEV, M.V. & NIYAZOVA, M.V. 1998: On the discovery of
Tapinoma melanocephalum (Dolichoderinae) in Moscow. In:
Ants and forest protection. Materials of the 10th All-Russian
Myrmecological Symposium, Peshki, 24 - 28 August 1988,
p. 153.
LEE, C.Y, LIM, C.Y. & DARAH, I. 2002: Survey on structure-in-
festing ants (Hymenoptera; Formicidae) in food preparative
outlets. – Tropical Biomedicine 19: 21-26.
LESTER, P.J. 2005: Determinants for the successful establishment
of exotic ants in New Zealand. – Diversity and Distributions
11: 279-288.
LONGINO, J.T. 2006: Ants of Costa Rica. – <http://academic.
evergreen.edu/projects/ants/AntsofCostaRica.html>, retrieved
on 1 December 2006.
31

MAMET, R. 1954: The ants (Hymenoptera Formicidae) of the
Mascarene Islands. – Mauritius Institute Bulletin 3: 249-259.
MANN, W.M. 1921: The ants of the Fiji Islands. – Bulletin of the
Museum of Comparative Zoology 64: 401-499.
MAYR, G. 1876: Die australischen Ameisen. – Journal des Mu-
seums Godeffroy Hamburg 12: 56-115.
MEKLOY, P. 2001: All about ants. – Bangkok Post 15 January
2001.
MENOZZI, C. 1930: Formiche della Somalia italiana meridionale.
– Memorie della Società Entomologica Italiana 9: 76-130.
MENOZZI, C. 1942: Formiche dell'isola Fernando Poo e del terri-
torio del Rio Muni (Guinea Spagnol). 24. Beitrag zu den Wis-
senschaftlichen Ergebnissen der Forschungsreise H. Eidmann
nach Spanisch-Guinea 1939 bis 1940. – Zoologischer Anzei-
ger 140: 164-182.
MENOZZI, C. & RUSSO, G. 1930: Contributo alla conoscenza del-
la mirmecofauna della Repubblica Dominicana (Antille). – Bol-
lettino del Laboratorio di Zoologia Generale e Agraria della
Reale Scuola Superiore d'Agricoltura, Portici 4: 148-173.
MOREIRA, D.D.O., DE MORAIS, V., VIEIRA-DA-MOTTA, O., CAM-
POS-FARINHA, A.E. DE C. & TONHASCA, A. JR . 2005: Ants as
carriers of antibiotic-resistant bacteria in hospitals. – Neotrop-
ical Entomology 34: 999-1006.
NUGENT, M., SYTSMA, M., CHAN, S., HILBURN, D., CUDD, S.,
DEMASI, R., ALEXANIAN, K., TU, M., SCHWAMBERGER, E., GUN-
TERMANN, C., HENRY, R. & REYNOLDS, B. 2005: Invasive spe-
cies in Oregon. Report card 2005. – Oregon Invasive Species
Council, Salem, 10 pp.
ONOYAMA, K. 1976: A preliminary study on the ant fauna of
Okinawa-Ken, with taxonomic notes (Japan; Hymenoptera:
Formicidae). In: IKEHARA, S. (Ed.): Ecological studies of na-
ture conservation of the Ryukyu Islands. II. – University of
Ryukyus, Naha, Okinawa, pp. 121-141.
ONOYAMA, K. 1980: An introduction to the ant fauna of Japan,
with a check list (Hymenoptera, Formicidae). – Kontyû 48:
193-212.
OSBORNE, L.S., PEÑA, J.E. & OI, D.H. 1995: Predation by Tapi-
noma melanocephalum (Hymenoptera: Formicidae) on two-
spotted spider mites (Acari: Tetranychidae) in Florida green-
houses. – Florida Entomologist 78: 565-570.
PARDO, V.R. 1964: Clave para identificar los Formicidae de la
provincia de Chiclayo. – Revista de Peruana Entomologia 7:
98-102.
PERGANDE, T. 1896: Mexican Formicidae. – Proceedings of the
California Academy of Sciences (2)5: 858-896.
RADCHENKO, A. 2004: Formicidae. – Fauna Europaea version 1.1.
– <http://www. faunaeur.org/>, retrieved on 11 December 2006.
ROOM, P.M. 1971: The relative distributions of ant species in
Ghana's cocoa farms. – Journal of Animal Ecology 40: 735-751.
SANTSCHI, F. 1914: Formicides de l'Afrique occidentale & aus-
trale du voyage de Mr. le Professeur F. Silvestri. – Bollettino del
Laboratorio di Zoologia Generale ed Agraria, Portici 8: 309-385.
SANTSCHI, F. 1920: Fourmis d'Indo-Chine. – Annales de la So-
ciété Entomologique de Belgique 60: 158-176.
SANTSCHI, F. 1928a: Formicidae (Fourmis). – Insects of Samoa
5: 41-58.
SANTSCHI, F. 1928b: Nouvelles fourmis d'Australie. – Bulletin de
la Société Vaudoise des Sciences Naturelles 56: 465-483.
SHEPARD, M. & GIBSON, F. 1972: Spider-ant symbiosis: Cotinusa
spp. (Araneida: Salticidae) and Tapinoma melanocephalum
(Hymenoptera: Formicidae). – Canadian Entomologist 104:
1951-1954.
SMITH, F. 1857: Catalogue of the hymenopterous insects collect-
ed at Sarawak, Borneo; Mount Ophir, Malacca; and at Singa-
pore, by A. R. Wallace. – Journal and Proceedings of the Lin-
nean Society of London Zoology 2: 42-88.
SMITH, F. 1860: Catalogue of hymenopterous insects collected by
Mr. A. R. Wallace in the islands of Bachian, Kaisaa, Amboy-
na, Gilolo, and at Dory in New Guinea. – Journal and Pro-
ceedings of the Linnean Society of London Zoology 5(17b):
93-143.
SMITH, M.R. 1930: A list of Florida ants. – Florida Entomolo-
gist 14: 1-6.
SMITH, M.R. 1951: Family Formicidae. In: MUESEBECK, C.F.,
KROMBEIN, K.V. & TOWNES, H.K. (Eds.): Hymenoptera in
America north of Mexico. Synoptic catalog. – USDA Agri-
cultural Monograph 2: 778-875.
SMITH, M.R. 1979: Superfamily Formicoidea. In: KROMBEIN,
K.V., HURD, P.D. JR., SMITH, D.R. & BURKS, B.D. (Eds.): Cata-
log of Hymenoptera in America North of Mexico. Volume 2.
Apocrita (Aculeata). – Smithsonian Institution Press, Wash-
ington, D.C., pp. 1323-1467.
SNELLING, R. 2000: Ants recorded at two camps in the Wapoga
area, Irian Jaya, Indonesia. – RAP Bulletin of Biological As-
sessment 14: 95-100.
SORVARI, J. 2003: Tapinoma melanocephalum (FABRICIUS 1793),
asunnoissa pesivä muurahainen – uusia havaintoja Suomessa
(Hymenoptera, Formicidae, Dolichoderinae). – Sahlbergia 8: 20.
STEINER, F.M., SCHLICK-STEINER, B.C., SCHÖDL, S. & ZETTEL,
H. 2003: Neues zur Kenntnis der Ameisen Wiens (Hymeno-
ptera: Formicidae). – Myrmecologische Nachrichten 5: 31-35.
TAYLOR, R.W. 1961: Notes and new records of exotic ants in-
troduced into New Zealand. – New Zealand Entomologist 6:
28-37.
TAYLOR, R.W. 1967: Entomological survey of the Cook Islands
and Niue. 1 - Hymenoptera-Formicidae. – New Zealand Jour-
nal of Science 10: 1092-1095.
TAYLOR, R.W. 1976: La faune terrestre de l'Ile de Sainte-Hé-
lène. 7. Superfam. Formicoidea. – Annales du Musée Royal
de l'Afrique Centrale, Sciences Zoologiques 215: 192-199.
TIMBERLAKE, P.H. 1926: Hymenoptera. – Bulletin of the B. P.
Bishop Museum 31: 17-43.
VAN ZWALUWENBERG, R.H. 1943: The insects of Canton Island. –
Proceedings of the Hawaiian Entomological Society 11: 300-
312.
VENKATARAMAIAH, G.H. & REHMAN, P.A. 1989: Ants associat-
ed with the mealybugs of coffee. – Indian Coffee 43: 13-14.
VIERBERGEN, B. 2003: Technomyrmex albipes en andere extoten
in Nederland. – Forum Formicedarum 4: 4-7.
WAY, M.J. 1953: The relationship between certain ant species
with particular reference to biological control of the coreid, The-
raptus sp. – Bulletin of Entomological Research 44: 669-691.
WEBER, N.A. 1939: Tourist ants. – Ecology 20: 442-446.
WEBER, N.A. 1948: Studies on the fauna of Curaçao, Aruba,
Bonaire and the Venezuelan islands: No. 14. Ants from the Lee-
ward Group and some other Caribbean localities. – Natuur-
wetenschappelijke Studiekring voor Suriname en de Neder-
landse Antillen 5: 78-86.
WETTERER, J.K. 2002: Ants of Tonga. – Pacific Science 56:
125-135.
WETTERER, J.K. 2008: Worldwide spread of the longhorn crazy
ant, Paratrechina longicornis. – Myrmecological News 11:
137-149.
WETTERER, J.K. & O'HARA, B.C. 2002: Ants of the Dry Tortu-
gas, the outermost Florida Keys. – Florida Entomologist 85:
303-307.
WETTERER, J.K., MILLER, S.E., WHEELER, D.E., OLSON, C.A.,
POLHEMUS, D.A., PITTS, M., ASHTON, I.W., HIMLER, A.G.,
32

33
YOSPIN, M., HELMS, K.R., HARKEN, E.L., GALLAHER, J., DUN-
NING, C.E., NELSON, M., LITSINGER, J., SOUTHERN, A. & BUR-
GESS, T.L. 1999: Ecological dominance by Paratrechina longi-
cornis (Hymenoptera: Formicidae), an invasive tramp ant, in
Biosphere 2. – Florida Entomologist 82: 381-388.
WETTERER, J.K. & VARGO, D.L. 2003: Ants (Hymenoptera: For-
micidae) of Samoa. – Pacific Science 57: 409-419.
WHALEN, M.A. & MACKAY, D.A. 1988: Patterns of ant and her-
bivore activity on five understory euphorbiaceous saplings in
submontane Papua New Guinea. – Biotropica 20: 294-300.
WHEELER, W.M. 1905: The ants of the Bahamas, with a list of
the known West Indian species. – Bulletin of the American
Museum of Natural History 21: 79-135.
WHEELER, W.M. 1907: A collection of ants from British Hon-
duras. – Bulletin of the American Museum of Natural History
23: 271-277.
WHEELER, W.M. 1913a: Zoological results of the Abor Expedi-
tion, 1911-1912, XVII. Hymenoptera, II. Ants (Formicidae).
– Records of the Indian Museum 8: 233-237.
WHEELER, W.M. 1913b: Ants collected in the West Indies. –
Bulletin of the American Museum of Natural History 32:
239-244.
WHEELER, W.M. 1916: Ants collected in Trinidad by Professor
Roland Thaxter, Mr. F. W. Urich, and others. – Bulletin of the
Museum of Comparative Zoology 60: 323-330.
WHEELER, W.M. 1921: Chinese ants collected by Prof. C. W.
Howard. – Psyche 28: 110-115.
WHEELER, W.M. 1929: Ants collected by Professor F. Silvestri
in Formosa, the Malay Peninsula and the Philippines. – Bol-
lettino del Laboratorio di Zoologia Generale e Agraria della
Reale Scuola Superiore d'Agricoltura, Portici 24: 27-64.
WHEELER, W.M. 1935: Check list of the ants of Oceania. – Oc-
casional Papers of the Bernice Pauhahi Bishop Museum 11
(11): 1-56.
WHEELER, W.M. 1936: Ants from the Society, Austral, Tuamotu
and Mangareva Islands. – Occasional Papers of the Bernice
Pauhahi Bishop Museum 12(18): 3-17.
WHEELER, W.M. & MANN, W.M. 1914: The ants of Haiti. –
Bulletin of the American Museum of Natural History 33: 1-61.
WILSON, E.O. & HUNT, G.L. 1967: Ant fauna of Futuna and Wal-
lis Islands, stepping stones to Polynesia. – Pacific Insects 9:
563-584.
WILSON, E.O. & TAYLOR, R.W. 1967: Ants of Polynesia. – Pa-
cific Insects Monographs 14: 1-109.