ASIAN MYRMECOLOGY Volume 7, 105 – 112, 2015

ISSN 1985-1944 © James K. Wetterer, Benoit Guénard and Douglas B. Booher

Geographic spread of Vollenhovia emeryi(Hymenoptera: Formicidae)
James K. Wetterer1*, Benoit Guénard2 and Douglas B. Booher3
Wilkes Honors College, Florida Atlantic University, 5353 Parkside Drive,
Jupiter, FL 33458, USA
2
School of Biological Sciences & The University of Hong Kong,
Pokfulam Road & Hong Kong, PRC
3
Department of Ecology and Evolutionary Biology, University of California
Los Angeles, 621 Charles E. Young Drive S, Los Angeles,
CA 90095-1606, USA
1*Corresponding author's email: wetterer@fau.edu
ABSTRACT. Vollenhovia emeryi Wheeler is a small and inconspicuous
ant species originally from East Asia that recently has been found in North
America. Here, we examine the geographic spread of V. emeryi and compare
its native range in Asia with its exotic range in North America. We compiled
published and unpublished V. emeryi specimen records from >300 sites. We
documented the earliest known V. emeryi records for 14 geographic areas, ten
in Asia (Japan, North Korea, South Korea, Taiwan, Thailand, and five provincesof China) and four in North America (Maryland, Pennsylvania, Virginia, andWashington DC). Reports of V. emeryi from lower latitude (14.3°N to 25.1°N)
sites in Taiwan, Thailand, and southern China, however, probably represent a
distinct species. Vollenhovia emeryi has a much broader latitudinal spread in
East Asia (at least 13.5° range: 29.7°N to 43.2°N) than it has in North America(1.3° range: 38.7°N to 40.0°N). The North American records of V. emeryi are allat latitudes near the northernmost records in Asia. It seems likely that V. emeryi
will spread further in North America, particularly towards the south and west.
Keywords: biogeography, biological invasion, exotic species, invasive species
INTRODUCTION
Forel (1911) compiled a list of tramp ant species,
spread by human commerce, which had achievedor were in the process of achieving broad
cosmopolitan distributions. The 14 cosmopolitan
species on this list all originated in the tropics and/or subtropics. However, in what appears be a new
trend, several temperate ant species are spreading
outside their native hemisphere (e.g., Myrmica
rubra (L.); Wetterer & Radchenko 2011), including
five ant species native to temperate East Asia thatare now spreading in temperate North America(earliest North American record in parentheses):

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Brachyponera chinensis (Emery) (1932; Guénard
& Dunn 2010), Nylanderia flavipes (Smith) (1939;
Wetterer 2011), Vollenhovia emeryi Wheeler(1986; Kjar & Suman 2007), Strumigenys
hexamera (Brown) (1987; MacGown & Wetterer2012), and Tetramorium tsushimae Emery (1988;
Reuther 2009). Here, we examine the geographic
distribution of V. emeryi and compare its native
range in Asia with its exotic range in North
America.
Wheeler (1906) described V. emeryi fromJapan. Radchenko (2005) designated Vollenhovia
emeryi chosenica Wheeler, described fromSouth Korea, as a junior synonym of V. emeryi.

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106

James K. Wetterer, Benoit Guénard & Douglas B. Booher

Vollenhovia emeryi colonies typically nest in thewood and under the bark of fallen trees (Terayama& Yamauchi 2003; BG, pers. obs.) in old-growthand second-growth deciduous and conifer forests(Japan Ant Database Group 2003, Maeto & Sato
2004, Kwon et al. 2005, Hosoishi et al. 2007).
This species is also occasionally found in urban
parks (Iwata et al. 2005, Harada et al. 2010) andin agricultural fields (Hosoishi et al. 2007).
Much recent research on V. emeryi
concerns their remarkable mode of reproduction.
Vollenhovia emeryi queens produce daughter
queens through parthenogenesis and produce
males that are clones of their mates (Ohkawara
et al. 2006; Kobayashi et al. 2008, 2011, 2012),
a reproductive mode similar to that reported
for some other ant species, including the tramp
ants Wasmannia auropunctata (Roger) andParatrechina longicornis (Latr.) (Fournier et
al. 2005; Pearcy et al. 2011), but with some
additional complexity. Whereas V. emeryi workers
are monomorphic (Fig. 1), V. emeryi queens
have two morphotypes: long-winged (L) andshort-winged (S) (Figs. 2-3). Males also occur
in two genetically discrete (but morphologicallyindistinguishable) types, with L and S males

mating only with L and S queens, respectively(Kobayashi et al. 2011). The queen and malegenomes do not mix in reproductives and there
is no recent gene flow between queens and males(Kobayashi et al. 2011). Phylogenetically, Smorph queens form their own clade, whereas the
L morph queens are paraphyletic, indicating the Lmorph is the primitive condition (Kobayashi et al.
2012). Thus, the S morph V. emeryi would seemin most ways like a separate species from the Lmorph V. emeryi, except for one twist: S males
appear to be more closely related to L queens andL males than they are to S queens (Kobayashi etal. 2008).
METHODS
Using published and unpublished records, wedocumented the worldwide range of V. emeryi. We
obtained unpublished site records from museum
specimens in the collections of the Museum of
Comparative Zoology (MCZ, identified by S.
Cover) and the Smithsonian Institution (SI,identified by M. Smith). In addition, we used
on-line databases with collection information on

Table 1. Earliest known records for Vollenhovia emeryi. * = may be based on misidentifications (see text)
Asia

Earliest recordJapan

1905 (Wheeler 1906)

Taiwan1908 (Forel 1912)

South Korea≤1928 (Wheeler 1928 as V. emeryi chosenica)

North Korea

1970 (Radchenko 2005)

Thailand*

1981 (Brown 1988)

Yunnan, China*≤1999 (Xu et al. 1999)

Zhejiang, China*≤2001 (Zhou 2001)

Guangxi, China*≤2001 (Zhou 2001)

Hunan, China*≤2005 (Huang & al. 2005)

Hubei, China*≤2009 (Wang & Zhao 2009)

North America

Earliest record

Washington DC

1986 (Kjar & Suman 2007)

Pennsylvania

1993 (King & Green 1993)

Virginia

2002 (Kjar & Suman 2007)

Maryland2003 (Kjar & Suman 2007)

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Geographic spread of Vollenhovia emeryi

107

Fig. 1. Vollenhovia emeryi worker from Cabin John Regional Park, Maryland (photo by Joe MacGown fromwww.antweb.org).

Fig. 2. Vollenhovia emeryi long-winged queen from Rock Creek Regional Park, Maryland (photo by Joe MacGown
from www.antweb.org).

specimens by Antweb (www.antweb.org), theGlobal Biodiversity Information Facility (www.
gbif.org), and the Japanese ant image database(Terayama & Yamauchi 2003).
We obtained geo-coordinates forcollection sites from published references,
specimen labels, maps, or geography web sites

10 - 18 - AM7 - Geographic spread of Vollenhovia emeryi.indd 107(e.g., earth.google.com and www.tageo.com). If asite record listed a geographic region rather than a“point locale,” and we had no other record for thisregion, we used the coordinates of the capital or
largest town of the region or, in the case of small
islands and natural areas, the center of the region.

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108James K. Wetterer, Benoit Guénard & Douglas B. Booher

We did not map records of V. emeryifound in newly imported goods or intercepted intransit by quarantine inspectors at a port of entry
because the original source of such material isnever known. For example, in 1961, V. emeryi
was intercepted in Hawaii in a shipment of pinefrom Japan (antweb.org).
Radchenko (2005) determined thatspecimens from North Korea that Collingwood(1976) identified as Leptothorax congruus Smith
were actually V. emeryi. We assumed Forel’s (1912)
record of V. emeryi from Pilam (now Beinan),Taiwan was based on the specimen that T. Shirakicollected in 1908 at Pilam (Taiwan Agricultural
Research Institute; http://digiins.tari.gov.tw/tarie/
search_Result2E.php?id=form08159001).
RESULTS AND DISCUSSION
We compiled published and unpublished V. emeryi
specimen records from >300 sites (Fig. 4). We
documented the earliest known V. emeryi records
for 14 geographic areas: ten in Asia (Japan,
North Korea, South Korea, Taiwan, Thailand,
and five provinces of China) and four in North
America (Maryland, Pennsylvania, Virginia, andWashington DC) (Table 1). Records ranged fromsea level (Ogata et al. 1994) to 1450 m elevation(Kwon et al 2014).
At the northern end of the Asian range,
we found only seven records of V. emeryi atlatitudes higher than 40.0°N: five from Japan (upto 43.2°N in Hokkaido; Morisita 1945, Hayashida
1957, 1961, Terayama 1994) and two from North
Korea (up to 41.8°N; Radchenko 2005). Terayama& Kinomura (1997) wrote that V. emeryi “iscommonly and widely distributed from Hokkaido
to Yaku-shima.” We found only two records of V.
emeryi in Japan south of Yaku-shima, i.e., from
Nakano-shima (29.9°N; Harada et al. 2014) andTaira-jima (29.7°N; Ogata 1981). However, wefound records from much lower latitudes in otherparts of Asia, i.e., in Thailand (14.3°N; Brown
1988), Taiwan (22.8°N & 25.1°N; see Results),
and southern China (Xishuangbanna, Yunnan
Province: 22.0°N, Xu et al. 1999; Guangxi
Province: ~22.8°N, Zhou 2001).
The absence of V. emeryi from themany Japanese islands south of Taira-jima

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combined with the great latitudinal difference
between Taira-jima and the records from much
lower latitudes in Thailand, Taiwan, and Chinasuggests that the latter records may be basedon misidentifications. In fact, Terayama (2009)
explicitly excluded V. emeryi from the list ofants of Taiwan, writing: “although Forel (1912)
recorded V. emeryi from Pilam (= Taitung Pref.),no reliable additional record is known.” Forel’s(1912) ‘V. emeryi’ specimens from Taiwan may
be the same species as Vollenhovia cf. emeryi
reported by Fellowes et al. (2002, 2003) fromsouthern China (Hainan Island: 18.8°N; Guangxi
Province: 23.7°N). In fact, it is possible that themore temperate records of V. emeryi from China(Hubei Province: ~30.6°N, Wang & Zhao 2009;
Zhejiang Province: 29.0°N, Zhou 2001; Hunan
Province: 25.3-29.7°N, Huang et al. 2005) arealso based on misidentifications. Morphologicaland genetic evaluation of putative V. emeryi
specimens from China, Taiwan, and Thailand
would be valuable in establishing the geographic
and taxonomic limits of V. emeryi.
In 1986, Stefan P. Cover discovered thefirst North American population of V. emeryi inRock Creek Park, Washington DC. Since then,additional North American populations havebeen reported from Maryland, Pennsylvania,
and Virginia (Table 1; Fig. 4). Overall, V. emeryirecords ranged (north to south) from Philadelphia,
Pennsylvania (40.0°N; King & Green 1993) toFort Washington Park, Maryland (38.7°N; Kjar
& Suman 2007). Thus, V. emeryi has a ten times
broader latitudinal spread in Asia (at least 13.5°
range: 29.7°N to 43.2°N) than it has in North
America (1.3° range: 38.7°N to 40.0°N). The
North American records of V. emeryi are all fromlatitudes near the northernmost records in Asia
(Fig. 4). One possible explanation for this pattern
may be that the North American populations of V.
emeryi have a fairly narrow range of temperate
climatic tolerances. In Asia, different V. emeryi
populations may show physiological adaptations
to the local climate. Thus, although the species asa whole is able to live under a great diversity ofclimatic conditions in Asia, any one population
has a much narrower tolerance range. It is
possible that the populations of V. emeryi now
in North America are descended from colonists
from a narrow geographic area and are not

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109

Fig. 3. Vollenhovia emeryi short-winged queen from Cabin John Regional Park, Maryland (photo by Joe MacGown
from www.antweb.org).

Fig. 4. Worldwide distribution records of Vollenhovia emeryi. Records from Taiwan, Thailand, and China may bebased on misidentifications (see text).

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110James K. Wetterer, Benoit Guénard & Douglas B. Booher

adapted to spread much beyond their current
latitudinal range. While it is not clear what its
potential latitudinal limits are in North America,
there appears to be no geographic barriers thatwould prevent V. emeryi in North America fromspreading towards the north, south, and west. The
micro-satellite markers of V. emeryi specimensfrom North America do not match the microsatellite markers of any Asian population that has
been examined so far (D.B. Booher, unpublisheddata, M. Okamoto, pers. comm.), thus thegeographic origins of the exotic North American
populations remain unknown.
Kjar & Suman (2007) reported that all
V. emeryi queens found in North America were S
morph. The L morph queens and males typicallymate during nuptial flights, whereas the S morph
queens and males mate primarily with siblings
within their natal colonies (Okamoto & Ohkawara2010). Kjar & Suman (2007) speculated that any
long-range dispersal would be limited in North
America because S morph V. emeryi do not havenuptial flights. In 2012, however, D. Booher &R.M. Duffield collected long-winged V. emeryi
queens for the first time in North America (Fig.
2), though preliminary genetic analyses indicate
these may be atavistic S morph queens (D.B.
Booher, unpublished data).
Booher & Duffield (pers. obs.) foundthat V. emeryi has become the most common litter
dwelling ant at some sites in Rock Creek Regional
Park and in some riparian forests bordering the
Chesapeake & Ohio Canal in Maryland. Thus, if
V. emeryi does become more widespread in North
America, it is possible that this species could havesignificant ecological impacts in its exotic range.
Kinomura & Yamauchi (1992) described
Vollenhovia nipponica Kinomura & Yamauchi, aworkerless social parasite that lives in the nestsof V. emeryi. In Asia, V. nipponica is now known
from eight prefectures in Japan with a latitudinalrange of 31.4°N to 36.6°N (Terayama andYamauchi 2003), all further south than the known
range of V. emeryi in North America. In 2012, D.
Booher & R.M. Duffield collected V. nipponica
for the first time in North America, at Rock Creek
Regional Park, Maryland. So far, V. nipponica
are only known to parasitize colonies of S morph
V. emeryi. Genetic analyses of native and exotic
populations of V. nipponica and their host V.

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emeryi colonies should prove to be valuable inuntangling the complicated genetic and ecologicalrelations of V. nipponica and V. emeryi.
ACKNOWLEDGMENTS
We thank M. Wetterer for comments on thismanuscript; S. Cover (MCZ) and T. Schultz (SI)for help with their respective ant collections;
W. O’Brien for GIS help; D.P. Wojcik and S.D.
Porter for compiling their valuable FORMIS
bibliography; R. Pasos and W. Howerton of the
FAU library for processing interlibrary loans;
Florida Atlantic University for financial support.
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ASIAN MYRMECOLOGY
A Journal of the International Network for the Study of Asian Ants
Communicating Editor: Dirk Mezger

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