First records of invasion by the myrmicine japanese ant Vollenhovia emeryi W. M. wheeler (Hymenoptera: Formicidae) in the United States

2007·4.9 MB·

by  Daniel S Kjar

PROC. ENTOMOL. SOC. WASH.
109(3), 2007, pp. 596-604
FIRST RECORDS OF INVASION BY THE MYRMICINE JAPANESE ANT
VOLLENHOVIA EMERYIW. M. WHEELER (HYMENOPTERA:
FORMICIDAE) IN THE UNITED STATES
Daniel S. Kjar and Theodore W. Suman
(DSK) Division of Mathematics and Natural Sciences, Elmira College, 1 Park
Place, Elmira, NY 14901, U.S.A. (e-mail: [email protected]); (TWS) Department of
Entomology, National Museum of Natural History, Smithsonian Institution,
P.O. Box 37012, Washington, DC 20013-7012, U.S.A.
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Abstract. Vollenhovia emeryi W. M. Wheeler was found at two sites over 2 yr in
the Dyke Marsh Preserve, part of the George Washington Memorial Parkway,
a National Park in Fairfax County, Virginia. This ant was also found at two
locations along the shore of the Potomac River in Maryland: Fort Washington
National Park, Prince George's County, and Glen Echo Park, Montgomery County.
These are the first records for their respective states. A live colony of V. emeryi was
collected from the campus ofGeorgetown University in the District ofColumbia and
is maintained at the Smithsonian Institution. Previous records for this ant in the
United States are: Rock Creek Park, a National Park in the District of Columbia
(1986), and Philadelphia, Pennsylvania (1993). These records indicate that this
species has spread beyond areas ofits initial introduction. This ant species may have
entered the U.S. with imported Japanese cherry trees in the District of Columbia,
and Philadelphia, Pennsylvania.
Key Words: Vollenhovia emeryi W. M.Wheeler, introduced species, alien, Japanese
ant
Introduced ant species are some ofthe Linepithima humile (Mayr) (Argentine
more destructive and expensive pests in ant), an aggressive alien ant introduced
the U.S. as well as worldwide (Crowell into southern California and Louisiana
1968, Porter and Savignano 1990, Cole et from South America in the 1890s, is
al. 1992, Morrison 2002, Breton et al. rapidly spreading into most of the
2003, Hill et al. 2003). However, the southwestern U.S. (Cole et al. 1992,
majority of ant species are not likely to Holway 1998). Solenopsis invicta Buren
be introduced through human activities (red imported fire ant) is one ofthe more
because of habitat preferences and spe- notorious alien invasives in the U.S.
cialization on specific prey. Alien, in- (Porter and Savignano 1990). This ant
vasive ant species tend to be habitat and spread rapidly after its introduction in
prey generahsts, often with polygynous Mobile, Alabama, during the 1930s and
colonies, and with nesting habits that its range extends through most of the
increase their likelihood of transporta- southern U.S. Solenopsis invicta quickly
tion through human trade (Holway et al. decimated local arthropod populations
2002). and has caused human injuries and
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VOLUME NUMBER
109, 3 597
deaths due to its venomous sting. Re- ian forests, have reduced wings (bra-
cently, and for reasons yet unknown, this chypterous), and a high rate of gynan-
ant appears to be decHning behind its dromorphy (a condition where part of
invasion hne in natural habitats, al- the individual is female and the other
though it is dominant and abundant in part male). Monogyne colonies are
disturbed areas (Morrison 2002). Many found in upland areas, nest in wood,
alien ant species become agricultural and have queens with fully-developed
pests by protecting crop-damaging ar- wings (macropterous) and infrequent
thropods and creating mounds in agri- gynandromorphy (Kubota 1984, Kino-
cultural fields. Other aggressive alien ant mura and Yamauchi 1994). Reproduc-
species in the U.S. include: Anoplolepis tives eclose in the fall and overwinter in
gracilipes (Smith), Paratrechina flavipes their nests. Wheeler (1906) described
(Smith), Pheidole megacephala (Fabri- taxonomic details and more recent work
cius), Solenopsis geminata (Fabricius), has focused on queen morphology and
Technomyrmex albipes Smith, and Was- gynandromorphy (Kubota 1984; Kino-
mannia auropunctata (Roger) (Wheeler mura and Yamauchi 1994).
1929, McGlynn 1999, Trager 1984, We give the first record of this ant in
Deyrup 1991, Trager 1991, Wetterer Virginia, and describe previously unpub-
and Porter 2003, Wetterer 2005) lished records of V. emeryi in Maryland,
Some introduced ant species do not Philadelphia, and Washington D.C. We
appear to displace native species or include observations made on a captive
greatly disrupt native ant communities, V. emeryicolony. We hypothesize that V.
and are predominately found in habitats emeryi is spreading along the riparian
altered by human activity. Monomoriwn corridors of Washington, D.C, and
pharaonis (Linnaeus) (Pharaoh ant) lives suggest that gifts of Japanese cherry
in dry, warm habitats, and a colony of trees during the early 20th century may
these 1.5 mm-long ants can travel great have resulted in the introduction of this
distances in crates, luggage, or machin- ant into the U.S. on two separate
ery. Such travel has resulted in this occasions.
species' worldwide distribution. Tetra-
Materials and Methods
rnorium caespitimi (Linnaeus) (pavement
ant) forms large colonies and lives in Study Sites. Dyke Marsh Preserve:
pavement cracks and building founda- Dyke Marsh Preserve (DMP) is part of
tions throughout the eastern U.S. the George Washington Memorial Park-
Stefan P. Cover first found VoUenJw- way (GWMP) in Fairfax County, Vir-
GWMP
via emeryi in the U.S. in 1986 while ginia. The is a national park
sampling for ants in Washington, D.C., bordering the west shore ofthe Potomac
and its presence in North America is River. The DMP is 3.5 km long, 500 m
briefly mentioned by Holldobler and wide at its widest point on an east-west
Wilson (1990). Vollenhovia spp. are transect, and located 15 km south of the
generalists and are distributed through- Ronald Reagan Washington National
DMP
out Asia and the Pacific Islands (Wheeler Airport. The has areas of flood-
1907, Bolton 1995, Terayama and Kino- plain forests, open tidal freshwater
mura 1997). There are no known Vol- marsh, and swamp forests (Johnston
lenhovia species native to the Western 2000; Barrows et al. 2005). All sampling
DMP
Hemisphere. The behavior and natural sites were within the flood-plain
history of V. emeryi is relatively un- forest.
known. In its native habitat, polygyne The flood-plain forest is dominated by
colonies nest in decaying wood in ripar- Liquidambar styraciflua L. (sweetgum)
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PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
598
and a dense under story of Lindera Glover-Archbold Park, part of Rock
benzoin (L.) Blume (spicebush) and Creek Park (a national park). Observa-
Viburnum nioUe Michx. (smooth arrow- tory Hill is a manicured landscape with
wood). Other trees common in the forest Asimina triloba (L.) Dunal (pawpaw).
include Acer negundo L. (boxelder), Acer Ilex spp. (hollies), Metasequoia glyptos-
rubrum L. (red maple), Fraxinus cimeri- troboides Hu and W. C. Cheng (dawn
cana L. (white ash), Liriodendron tulipi- redwood), Prunus cvs., Quercus spp.
fera L. (tulip tree), Nyssa sylvatica (oaks), Robinia pseudoacacia L. (black
Marshall (tupelo), Quercus palustris locust), and other trees.
Miinchh. (pin oak), Quercus phellos L. Site Selection. Dyke Marsh Preserve:
(willow oak), Quercus rubra L. (red oak). As part of a larger study on the ant
Sassafras albiduni (Nutt.) Nees (sassa- community of the DMP forest, 60
fras), and Ulmus cmiericana L. (american random sites were selected within the
elm). DMP using a geographical information
Fort Washington: Fort Washington system (ESRI Inc. 2001), the National
National Park (FWNP) is located Park Service's AlaskaPak extension (Na-
20 km south of Washington, D.C. in tional Park Service 2002), and high-
Prince George's County, Maryland. The
resolution aerial photography with the
current structure dates to 1824, and cooperation ofthe National Park Service
previous forts on this site date to 1808. GIS Coordinator of the GWMP. Sites
The park and surrounding area is greatly were in a predefined area of the forest
impacted by human activities. Trees m
whose borders were at least 5 from
species found around the park are trails or roads. A Trimble® backpack
Quercus sp., L. styraciflua, U. aniericana,
global positioning system (GPS) was
L. tulipifera, Platanus occidentalis L.
used to locate each of the sites in the
(eastern sycamore), and various Prunus
cvs. (cherries). Open areas are dominated forest.
Fort Washington: Three areas within
by grass and edge vegetation. Areas the FWNP were sampled during March
along the Potomac River are dominated
through September, 2002; and during
by young A. negundo and U. aniericana.
Most areas along the river were used as May and November, 2003. Sampling in
warfs and contain concrete, drift wood, the southeastern part of the park in-
and other beach debris. cluded open grass areas, young upland
Glen Echo Park: Glen Echo Park woods, gullies, and areas around and on
(GEP) is part of the GWMP on the Bunker B. Sampling in the western part
Maryland side of the Potomac River of the park included public fishing areas
7 km north of Washington, D.C. The near the shoreline and near the light-
area has been used as an amusement house. Sampling in the northeast of the
park since the late 19th century. Cur- park was done in wooded areas, picnic
rently the park is operated by the grounds, and along trails.
National Park Service for education Glen Echo Park: Ant sampling in GEP
and various family oriented activities. occurred as part of a Bioblitz sponsored
The forest around the GEP is second by the National Park Service, The
growth eastern deciduous and primarily Nature Conservancy, and the Virginia
consisting ofA. negundo, L. tulipifera, U. Museum of Natural History on June 24
americana, P. occidentcdis. and 25, 2006. Sampling was done along
Observatory Hill: Observatory Hill, on a short stretch of path leading down to
the Main Campus of Georgetown Uni- a gully near the Clara Barton Parkway
versity, Washington, D.C, is adjacent to (38° 58'02" N, 77° 08'47" W).
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VOLUME NUMBER
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A
Observatory Hill: pitfall trap was Results
palreaacedbeolnowObaselravragteoroyakHiltlreien (aQnueorpceuns Ant Co—llection at the Dyke Marsh
Preserve. Vollenhovia emeryiwas found
pruabwrpaawL.){A.antdrilonbeaa)rata38g°ro5v4e'28o"fN,sma7l7°l at two locations in the DMP: site 33 (38°
04'39" W. The area was open with pieces 46'27"N, 77° 03'01" W); site 58 (38°
46'26" N, 77° 03'01" W). Workers were
of bricks and pavement, loose gravel,
found in pitfall trap samples during both
and grass.
2002 and 2003. One was collected in June
Antcollection. DykeMarshPreserve:
Soil cores (70 X 70 mm) were collected of 2002, and another was collected
from each site in the third week ofJune, August 2002, both at site 33. This site
August, and October in 2002 and 2003. is dominated by the alien vine Ampelop-
Arthropods were extracted from the soil sis brevipedmunculata (Maxim.) Trautv.
with the use of Berlese funnels. A single and is 15 from the banks of the
Potomac River. Other alien plants in
collar and funnel pitfall trap of the
this site include Celastrus orbiculatus
design described in Kjar and Barrows
Thunb. (Asian bittersweet). Clematis
(2004) was used at each study site. The
terniflora Dc. (Asian clematis), Lonicera
pitfall traps were run for 24 h, in the last
japonica Thunb. (Japanese honeysuckle),
week of June, August, and October
during 2002 and 2003. and Rosa multiflora Thunb. (multiflora
Fort Washington: Sites at FWNP were rose). The only native plant in site 33 was
a small Primus serotina Ehrh. (wild black
sampled using Berlese funnels, pitfall
traps, and Winkler extraction (Suman cherry) seedling.
During August 2003, one worker was
2004).
Glen Echo Park: Hand sampling was found in a pitfall sample from site 58,
m
done for 30 minutes along the gully by approximately 30 south ofsite 33. Site
m
turning over stones, searching tree limbs 58 is 15 from the river and located
and leaf litter, and breaking up decom- below a 132-cm-dbh A. rubrum, a 64-cm-
posing tree limbs and stumps. dbh L. styraciflua, and a 51-cm-dbh U.
Observatory Hill: A single pitfall trap americana. The plant community in this
of identical design to the DMP site was site was dominated by the alien vine L.
left for 48 h starting on 14 September japonica, and also contained A. brevipe-
2004. Hand sampling near the pitfall trap dunculata. Native plants in site 58 in-
location was performed 7 June 2005. cluded all three tree species mentioned
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Ant Identification. Ants were identi- above and the monocot vine Smilax
fied using Bolton (1994), Creighton rotundifolia L. (greenbrier).
(1950), the National Museum ofNatural Ant Collection at Fort Washington.
History ant collection, and verified by Twenty-four workers of V. emeryi were
FWNP
Dr. David R. Smith (USDA), and Mr. found in on June 26, 2003. The
Terry P. Nuhn (USDA). Voucher speci- ants were found in debris along the shore
mens are in the arthropod collection of of the Potomac River near the light-
the Laboratory of Entomology and Bio- house. A Berlese funnel was used to
diversity at Georgetown University, extract arthropods from the debris.
Washington, D.C. Specimens collected Ant Collection at Glen Echo Park.
from Fort Washington National Park One V. emeryi worker was collected by
are deposited with the National Park hand at the beginning of the trail
Service under catalog number 13651 (Su- surveyed. The worker was found on the
man 2004). surface of the leaf litter layer.
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600 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
Fig. I. Three images ofa V. emeryiworker, and an image ofthe V. emeryi nest at the U.S. National
Museum ofNatural History. The framesare images of: a, profileview; b, head view; c. dorsalview; d. the
nest ofthe colony taken from Observatory Hill, Georgetown University, Washington, D.C.
Ant Collection on Observatory Hill. disturbed, workers quickly moved all
The 14 September 2004 pitfall-trap sam- exposed eggs and larvae under soil or
ple from Observatory Hill contained 10 within the moistened paper used to
V. emeryi workers along with several transport them. When hand collected,
common ant species: Camponotus peim- workers attempted to move away from
sylvanicus (De Geer), Lasiiis alienus an aspirator and forceps and did not
(Foerster), and Prenolepis imparis (Say). have the 'crouch-and-freeze' behavior of
On 7 June 2005, a colony of V. emeryi such small myrmicines as CypJiomyrmex
was hand-collected from the same area spp. and Temnothorax spp. (Wheeler
of Observatory Hill. The colony had 28 1907, Cole 1940, Fellers 1987). Ifa work-
workers, three dealate (queens that have er was pushed onto her dorsal surface,
lost their wings) females, five larvae, and she struggled to right herself. These ants
10 eggs. The colony's nest was located in have short legs with inflated femora,
soil mixed with bricks, gravel, and wood. although they are relatively elongate for
The nest was 2 cm below the soil surface their sinall overall length of about
with slightly mounded soil around the 2.5 mm (Fig. 1). The 28 workers from
opening. the colony showed no readily visible
Members of the colony collected from variation in coloration or size.
Observatory Hill were slow moving on The live colony was taken to Ted R.
a warm day. After the colony was Schultz and Eugenia Okonski at the
VOLUME NUMBER
109, 3 601
Fig. 2. Imageofabrachypterous V. emeryialatequeen. ThisqueenisfromStefanCover's 1986survey
ofRock Creek Park, Washington, D.C. (Photograph used with permission ofand taken by April Nobile,
Cahfornia Academy ofSciences, www.antweb.org).
National Museum of Natural History 1906, Bolton 1995), and therefore may
and Okonski is maintaining it at the have little problem acclimating from
Museum. After the colony was placed in southern Virginia north to southern
the artificial habitat it moved its larvae, New England along the U.S. East Coast.
DMP
eggs, and queens into the small hose used The June 2002 collection in is
to add water to the porous clay base of the first record of this ant in Virginia,
FWNP
the nest container. Within two weeks the and the June 2003 collection at is
colony moved into the small gap between the first record of this ant in Maryland.
this hose and the plastic wall of the The Observatory Hill collections (Sep-
container, and built a small chamber tember 2004 and June 2005) are the
with two openings from the small second and third collections ofthis ant in
amount of soil that was introduced into the District of Columbia. Previous re-
the container with the colony (Fig. Id). cords of this ant include the first record
Foragers frequently enter and leave the ofthis ant in the U.S. from a 1986 survey
chamber. Queens have not been observed of the ants of Rock Creek Park in
outside of this chamber. Washington, D.C. (Holldobler and Wil-
son 1990); and Philadelphia, Pennsylva-
Discussion nia, in 1993 (King and Green 2005). The
Vollenhovia emeryi is a recently dis- alate reproductive females found in the
covered alien myrmicine ant from Japan U.S. have reduced wings (brachypterous)
and may be spreading across the mid- (Fig. 2), and therefore mating flights,
Atlantic region of the U.S. This ant is especially ones of any distance may not
native to Japan (30^5" N) (Wheeler be possible. In view of this information.
602 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
it seems likely that V. emeryi was in- southern slope under an oak tree ap-
dependently introduced into Philadel- proximately 1 km from the Potomac
phia and Washington, D.C. River, indicates that this species is not
An intriguing hypothesis on the in- confined to wet habitats as previously
troduction of V. emeryi in the U.S. thought, although the Georgetown colo-
concerns the Japanese gift ofcherry trees ny at the National Museum of Natural
that were planted along the Tidal Basin History showed an obvious preference
by the Jefferson Memorial in Washing- for an extremely moist nest location
ton, D.C, and at Fairmont Park in (Fig. Id). The polygynous nest from an
Philadelphia. The first 2,000 trees in- upland dry area suggests the habitat
tended for Washington, D.C. were im- range of polygyne colonies in the in-
ported in 1910. This first shipment was troduced population should be investi-
destroyed due to the numerous insects gated, as it may differ from what has
and other pests inhabiting them (Nation- been reported for this species in Japan.
al Park Service 2005). After the U.S. Long-distance dispersal by wind is
received a second gift of 3,020 cherry unlikely in this epigaeic ant with puta-
trees from Japan, the trees were planted tively brachypterous reproductives. The
in 1912. More cherry trees were planted presence of this ant in the DMP forest
in 1965 and 1986 around the Tidal Basin and Fort Washington National Park is
and the area around the Washington probably the result ofits rafting in wood.
Monument (National Park Service This ant's preference for wet rotting
2001). The fact that Japan gave 2,000 wood may result in queens' founding
cherry trees to Philadelphia in honor of nests in wood that has been left when
the 150th anniversary (1926) of the U.S. tides go out and picked up again and
Declaration of Independence provides moved down river during subsequent
further support to the cherry tree hy- high tides. Most flooding events would
pothesis. send V. emeryi from the City down-
DMP FWNP
Transport in the root balls of these stream to and with the
trees would be ideal for ants with a wet exception of hurricane associated storm
nesting habitat preference. There were no surges. However, the Potomac River is
exhaustive surveys of the District of tidal in the Washington, D.C, Area and
Columbia's ant fauna after Theodore therefore they could raft up and down
Pergande's (1840-1916) time at the the river depending on conditions. How-
National Museum of Natural History ever, the GEP record of V. emeryi 7 km
until Stefan Cover's work in the mid- north of the city adds support to
1980s. This small ant could have easily a gradual spread and an early introduc-
spread throughout the Washington, tion.
D.C, area unnoticed during this 70-yr VoUenhovia emeryi was found in moist
lapse of myrmecological survey data. stream bottoms in Rock Creek Park.
The potential ecological impact ofthis Rock Creek empties into the Potomac
alien ant is entirely unknown. Cover's River2 km upstream oftheTidal Basin. It
observations in Rock Creek Park did not is possible that V. emeryi traveled along
reveal displacement of native ant species the river banks and the Chesapeake and
by this species, and few ant species in this Ohio Canal spreading throughout Rock
area, with the exception of Ponera Creek Park, Georgetown University, and
pennsylvanica Buckley, share this species' GEP which are all connected directly to
nesting preference of extremely hydric the river and canal by forested park land.
rotting wood. The location of V. emeryi Human introduction of V. emeryi to each
at Georgetown University, a sunny of the five Washington, D.C, area loca-
VOLUME NUMBER
109, 3 603
tions is possible, but the apparently re- Service, Georgetown University, Disco-
stricted regional distribution of this ant verlife.org, Stefan P. Cover for providing
species suggests that it had a single in- the cherry tree hypothesis, David R.
troduction in this city and another in Smith, Terry P. Nuhn, Maya Patel, John
Philadelphia. If this species were com- Pickering, Ted Schultz and his ant lab,
monly introduced by humans through and the National Museum of Natural
agricultural or horticultural products, it History, Smithsonian Institution.
likely would have come to the attention of
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