Brachyponera chinensis
(Asian needle ant)

Pictures

Picture	Title	Caption	Copyright
Title Adult Caption Brachyponera chinensis (asian needle ant); adult. Copyright ©H. Kohori (harum.koh)/via flickr - CC BY-SA 2.0	Adult	Brachyponera chinensis (asian needle ant); adult.	©H. Kohori (harum.koh)/via flickr - CC BY-SA 2.0
Title Adult Caption Brachyponera chinensis (asian needle ant); adult, lateral view. Museum set specimen. Note scale. Copyright ©April Nobile/AntWeb.org 2002-2019 - CC BY 4.0	Adult	Brachyponera chinensis (asian needle ant); adult, lateral view. Museum set specimen. Note scale.	©April Nobile/AntWeb.org 2002-2019 - CC BY 4.0
Title Adult Caption Brachyponera chinensis (asian needle ant); adult, dorsal view. Museum set specimen. Note scale. Copyright ©April Nobile/AntWeb.org 2002-2019 - CC BY 4.0	Adult	Brachyponera chinensis (asian needle ant); adult, dorsal view. Museum set specimen. Note scale.	©April Nobile/AntWeb.org 2002-2019 - CC BY 4.0
Title Adult Caption Brachyponera chinensis (asian needle ant); adult, head capsule. Museum set specimen. Note scale. Copyright ©April Nobile/AntWeb.org 2002-2019 - CC BY 4.0	Adult	Brachyponera chinensis (asian needle ant); adult, head capsule. Museum set specimen. Note scale.	©April Nobile/AntWeb.org 2002-2019 - CC BY 4.0

Identity

Preferred Scientific Name

• Brachyponera chinensis (Emery, 1895)

Preferred Common Name

• Asian needle ant

Other Scientific Names

• (Brachyponera sinensis) (Misspelled by Imai and Kubota, 1972)
• Brachyponera solitaria Smith 1860
• Euponera chinensis Emery
• Euponera solitaria Emery 1901
• Pachycondyla chinensis (Emery)
• Ponera nigrita subsp. chinensis Emery 1895
• Ponera solitaria Smith 1874

International Common Names

• English: chinese needle ant; giant needle ant

Local Common Names

• Japan: oo-hari-ari

Summary of Invasiveness

Brachyponera chinensis is a temperate ant species native to East Asia, including China, Taiwan, North Korea, South Korea and Japan.

The first records of B. chinensis in the USA were from the states of Georgia, North Carolina and Virginia in 1932, but distribution patterns within these states suggest it may have been present from the early 1900s. It is likely to have been introduced from Japan, probably via shipping ports. This species is now one of the most widespread invasive ant species in the USA, recorded along a continuous range on the East Coast from Florida to Connecticut, west to Arkansas and isolated records from Wisconsin and Washington State. While not aggressive, individuals will sting when disturbed. The sting is painful with pain lasting for a few hours and more rarely leading to urticaria and anaphylaxis in the most serious cases.  

It has naturalized in urban areas and in natural forest, and there is evidence that it is displacing native ant species, such as species of the Aphaenogaster rudis complex, in Georgia and North Carolina. In some areas studied in North Carolina, it may be more abundant than all native species combined, disrupting native ecological communities.

B. chinensis does not disperse plant seeds as efficiently as species of the A. rudis complex, and there is some evidence that this is affecting plant distribution patterns in areas where B. chinensis has displaced native species.     

There is evidence that B. chinensis is also expanding its range into the eastern coast of the Black Sea (Georgia and Southern Russia).

Taxonomic Tree

• Domain: Eukaryota
•     Kingdom: Metazoa
•         Phylum: Arthropoda
•             Subphylum: Uniramia
•                 Class: Insecta
•                     Order: Hymenoptera
•                         Family: Formicidae
•                             Genus: Brachyponera
•                                 Species: Brachyponera chinensis

Notes on Taxonomy and Nomenclature

Brachyponera chinensis was initially described and named by F. Smith in 1874 and has undergone several taxonomic changes, including Ponera solitaria (Smith), Ponera nigrita chinensis (Emery), Euponera chinensis (Emery), Pachycondyla chinensis (Emery), and Brachyponera chinensis (Emery) (Allen, 2017). Although F. Smith undertook the initial identification of P. solitaria, Emery is credited as the author of B. chinensis because chinensis is the junior synonym of solitaria (Brown 1958, Allen 2017).

A reappraisal of ponerine ant morphological diversity, combined with recent molecular phylogenetic information, has been used to split the genus Pachycondyla into 19 genera, with the Asian needle ant being placed in the genus Brachyponera (B. chinensis: Schmidt and Shattuck, 2014). The genus Brachyponera contains 24 described species and subspecies (Schmidt and Shattuck, 2014).

Individuals of this species belong to a complex which has yet to be delimited (Brown, 1958; Yashiro et al., 2010;Guénard et al., 2018). This has created several problems, where the available information on this species sometimes refers to separate species within the Brachyponera chinensis complex (e.g. data in Gotoh and Ito, 2008 was later found to refer to B. nakasujii (see Yashiro et al., 2010)).

Overall, the exact taxonomic and geographic limits of Brachyponera chinensis in Asia are unknown, and it is possible that no less than 10 cryptic species co-exist in this region (B. Guénard, Hong Kong, personal communication, 2019). Here, we refer throughout to the known information regarding Brachyponera chinensis as described by Guénard et al. (2018). 

Description

A description of B. chinensis is complicated by the fact that the species shows great morphological variation (Yashiro et al. 2010). Also, taxonomic confusion within the B. chinensis species complex has led to inclusion of now separate species in older morphological descriptions. For example, Yashiro et al. (2010) identified the new cryptic species B.nakasujii through a morphological and molecular analysis of Japanese ant populations. This analysis also confirmed that B. chinensis, B. luteipes, and B. nigrita (Emery) are also three distinct species (Yashiro et al., 2010). This should be taken into account when referring to older morphological descriptions of B. chinensis.

The following text is edited from MacGown (2016):

Identification 
Worker: Average sized dark brown to black ant with outer antennae segments, mandibles, and legs a lighter orangish-brown with the workers usually less than 5 mm in total length. Head longer than wide with dense, appressed setae on the head, mandibles, and scape and a few erect setae; eyes small and located on the anterior quarter of the head; mandibles large and triangular with eight to ten teeth of various sizes; clypeus shining with a few long, anteriorly directed, erect setae; prominent frontal lobes covering the antennal insertion point; antennae 12-segmented. Mesosoma with a smooth and shiny mesopleural region while the rest of the mesosoma has dense appressed setae and some erect setae; propodeum strongly depressed below the mesonotum; hind tibia with two spurs, a large, pectinate one and a simple small one. Waist is single-segmented; subpetiolar process with a distinct, acutely angled projection with some erect setae. Gaster elongate and cylindrical with dense, appressed and erect setae; anterior face almost entirely vertical; prominent sting present at apex.

Queen: Dark brown to black ant with outer antennae segments, mandibles and legs being a lighter orangish-brown. Can be up to 6.50mm in length. Head longer than wide and covered with dense, appressed setae as well as the antennal scapes and with erect setae on the mandibles and the ventral surface; eyes ovoid with the posterior edge at the midline of the head; three ocelli present; mandibles well developed and triangular; frontal lobes prominent and covering the antennal insertion point; antennae are 12-segmented. Mesosoma with a mixture of appressed and erect setae and a glabrous, smooth, shining mesopleural region; enlarged with four wings or wing scars; hind tibia with two spurs, a large, pectinate one and a simple small one; posterior edge with a smooth and shining glabrous region anterior to the petiolar node. Waist single-segmented; petiolar node narrows to a point at the apex; subpetiolar process with a distinct, acutely angled projection with some erect setae. Gaster with a mixture of erect and appressed setae; first two tergites longer than the remaining ones; prominent sting present at the apex.

Male: Head, scape, pronotum, mesopleura, and apical portion of gaster yellowish brown; funiculus, propodeum, petiole, and anterior half or more of gaster dark brown; mesoscutum and legs pale yellow. Head small, almost circular looking, but longer than wide with a mixture of short, appressed setae and longer erect setae; eyes large, ovoid and distinctly convex; three ocelli present with dark pigmentation around their bases; mandibles reduced and indistinct with the rest of the mouth parts conspicuous; antennae 13-segmented; antennal scape about twice the length of the first funicular segment, but slightly shorter than the rest of the antennal segments. Mesosoma mostly smooth and shining; enlarged with four wings; propodeum with medial, smooth, shining area anterior to the petiolar node. Waist single-segmented with short and long erect setae; node roughly triangular in the lateral view, broadly rounded apically; subpetiolar process distinct with an acute tooth. Gaster with a mixture of short appressed setae and long erect setae; apical gastral segment coming to a distinct point, almost like a sting.

Distribution

Native to East Asia including Japan and China where it is widespread, and introduced to the southwestern states of the USA in addition to records from Washington and Wisconsin (Guénard et al., 2018; Antmaps, 2019). Expanding into the eastern coast of the Black Sea (Georgia and Southern Russia).

Distribution Table

The distribution in this summary table is based on all the information available. When several references are cited, they may give conflicting information on the status. Further details may be available for individual references in the Distribution Table Details section which can be selected by going to Generate Report.

History of Introduction and Spread

The mild climate and numerous shipping ports in Southeastern USA have contributed to the region being susceptible to the introduction and establishment of exotic ant species (MacGown et al., 2013).

It is believed that B. chinensis may have been introduced to the USA as early as the 1900s, but was first recorded in three Southeastern states (Georgia, North Carolina and Virginia) in 1932 (Smith, 1934; Guénard and Dunn, 2010; Guénard et al., 2018). Plant shipments infested with B. chinensis possibly reached the USA in the early 1900s where the ant became established (Paysen, 2007).

Phylogenetic analysis suggests that B. chinensis was introduced to the USA from the temperate zones of Japan (Yashiro et al., 2010).  

B. chinensis has also been recorded from New Zealand in the 1950s, although no further records of this species have been submitted in the last 60 years (Guénard et al., 2018). There have been reports of B. chinensis from the eastern coast of the Black Sea (Georgia and Southern Russia) between 2006 and 2017 (Guénard et al., 2018); the source of this population is unclear.

Introductions

Risk of Introduction

It is possible that B. chinensis has been accidentally introduced into the southeastern USA several times (Paysen, 2007). Multiple primary introductions may have occurred due to the lack of quarantine measures against B. chinensis, introducing genetic diversity and allowing viable populations to emerge (Paysen, 2007).

As international trade grows, there is a continued chance of introductions.

Habitat

Found in dark, damp, urban environments and disturbed rural environments (Smith, 1934; Mackay and Mackay, 2010). Colonies occur in rotten wood or in soil under stones, logs and debris (Mackay and Mackay, 2010).

B. chinensis build nests in both disturbed and natural areas. They are a temperate species, and are found in deciduous forests, hardwood forests, agricultural land, and in urban and suburban areas such as office parks and backyards (Miner, 2014). Nests are built on the forest floor, often in decaying logs and piles of leaves (Miner, 2014). 

In the USA, B. chinensis is particularly abundant in mature, temperate, hardwood forests including the Great Smoky Mountains National Park on the North Carolina/Tennessee border and several state parks in North Carolina, South Carolina and Alabama.

Colonies tend to establish nests near mature forests in termite galleries within logs or, in more urban areas, under the pavement, lawn ornaments or other man-made structures, for example, railroad ties, bricks and pavers (Paysen 2007, Guénard and Dunn 2010, Pećarević et al. 2010; MacGown et al., 2013; Miner, 2014).

In China, B. chinensis is ubiquitous on rice-paddy dykes (Brown 1958; Paysen, 2007).

On Yakushima Island, Japan, B. chinensis has been found in forests, forest edges, parks and residential areas (Harada et al., 2009).

Habitat List

Biology and Ecology

Genetics

The chromosome number of B. chinensis is 2n=22 (Imai and Kubota, 1972). The karyotype consists of five pairs of metacentric, five pairs of submetacentric and one pair of acrocentric or telocentric chromosomes (Imai and Kubota, 1972).

It appears that B. chinensis do not suffer from the deleterious genetic effects of a population bottleneck when introduced to new areas because sibling mating is a pre-existing behaviour in the native population (Eyer et al., 2018). Rather than being a consequence of the founder effect following introduction, inbreeding between sister queens and their brothers occurs in native populations and may have reduced inbreeding depression through purifying selection of deleterious alleles (Eyer et al., 2018).

Activity Patterns

A study of B. chinensis nests collected from North Carolina, USA, revealed that alate females and males were found within the nests continuously from early April to early September (Fig. 4, Guénard et al., 2018). Brood production was found continuously from early April to early October (n.b. no collections were made between mid-February to mid-March. Fig. 4, Guénard et al., 2018). 

In South Carolina, USA, a similar pattern of activity has been found; workers were found to be active from January to October, with low activity levels from January to March, increasing significantly from April and peaking in July/August (Zungoli and Benson, 2008). From mid-September, activity declined sharply and ceased by the end of October (Zungoli and Benson, 2008). When worker activity declined, ground colonies disappeared, although some colonies were located in stumps and logs in December (Zungoli and Benson, 2008). Activity was documented when mean weekly temperature was 15˚C or above (Zungoli and Benson, 2008). A later study by Allen (2017) showed that worker numbers were highest in May, August and October, with the majority of reproductive male and females produced in July and August.

In China, B. chinensis has been reported t­o forage in the open on tree trunks and on the ground in daylight (Brown, 1958). Foragers are more active on cloudy days compared to sunny days (Mackay and MacKay, 2010). 

Physiology and Phenology

B. chinensis is a temperate ant species which can live at a wide range of altitudes from sea-level up to 1100 m with a few records from higher elevations, including up to 2400 m in Nepal (Fig. 3,Guénard et al., 2018). It is found in disturbed habitats, such as parks, under paving slabs and near housing, but also in undisturbed forest locations, even in its introduced range (e.g. USA) (Guénard et al., 2018). The global range for B. chinensis sensu stricto according to Guénard et al. (2018) is from Washington State, USA, in the north (48°N) to Hong Kong (22°N). Records from further south require verification (e.g. southern China and Vietnam where B. chinensis can be easily confused with the slightly smaller B. obscurans (Guénard et al., 2018)).

Climate modelling has suggested that by 2080 the suitable range for B. chinensis could increase by 210.1% in Europe, 75.1% in Oceania, 74.9% in North America and 62.7% in Asia. Conversely, suitable climatic range is predicted to decrease in Africa by 22.9% (Bertelsmeier et al., 2013).

B. chinensis is not aggressive and generally attempts to flee from human contact, although will sting when pressured. Stings tend to occur when humans accidentally come into contact with the ants or their nests (Rice and Waldvogel, 2017).

Unlike most invasive ants, there is no evidence that B. chinensis use chemical signposts to mark the route to a food source, instead employing a relatively slower process known as tandem carrying (Guénard and Silverman, 2011). This process involves workers being carried individually under the belly of the carrier ant (using the mandibles) to a food resource and then released (Guénard and Silverman, 2011). This may be repeated until there are enough ants to retrieve the food source (Han, 2013). Guénard and Silverman (2011) found that this foraging behaviour was context dependent; tandem carrying occurred three to ten times more often with large non-moveable prey than with small moveable prey. B. chinensis appears to be able to detect its main prey, Reticulitermes termites, using olfactory cues (Bednar and Silverman, 2011).

Population Size and Structure

Colony sizes range from a single nest with 39 individuals up to multiple contiguous nest sites with over 5000 individuals (Zungoli and Benson, 2008).  

Nutrition

B. chinensis is a predator of termites in its native and introduced ranges, specifically preying on those in the genus Reticulitermes (Suehiro et al., 2017). It does not tend honeydew excreted by hemipterans, as relied upon by many other species of ants. While termites make up the majority of the diet of B. chinensis (estimated to be around 80%), their reliance on termites seems to be more marked in their native range rather than where they have been introduced (Suehiro et al., 2017). Radiocarbon analysis revealed a younger diet age in introduced populations, suggesting that while B. chinensis is still reliant on termites, it also feeds on other invertebrates with younger diet ages such as herbivorous insects; “We found that B. chinensis has markedly changed its prey composition and now relies less on termites in introduced range and more on other non-decomposer invertebrates.” (Suehiro et al., 2017).

B. chinensis has been observed scavenging crane flies, crickets, cockroaches, beetles, grasshoppers, collembolans, spiders, centipedes, earthworms and other dead organisms (Rice and Waldvogel, 2017). B. chinensis also directs other colony members to sugar sources such as apple jelly and scavenges food from bins (Rice and Waldvogel, 2017).

B. chinensis do not occur in habitats lacking Rhinotermitidae and its likely that the year round, high-quality food source provided by termites has contributed to the invasive success of this species in the USA, where subterranean termites are widespread and abundant (Bednar and Silverman, 2011).

Natural Food Sources

Climate

Latitude/Altitude Ranges

Means of Movement and Dispersal

Accidental Introduction

There are several records of B. chinensis being intercepted on imported plants, for example in Germany (Forel, 1900) and New Jersey (AntWeb, 2019 (record CASENT0246023, on Gentiana makinoi shipped from Japan)) (Guénard et al., 2018). B. chinensis appears to have been accidentally introduced to Greenville Zoo, South Carolina, USA via landscaping materials (plants and soil) (Nelder et al., 2006).

Pathway Causes

Pathway Vectors

Plant Trade

Impact Summary

Environmental Impact

In places where B. chinensis replaces native seed dispersing species such as those of the Aphaenogaster rudis complex [Aphaenogaster picea rudis], plant distribution is affected due to the failure of B. chinensis to disperse seed (Rodriguez-Cabal et al., 2012; Warren et al., 2015).

Impact: Biodiversity

A study in North Carolina, USA, found that B. chinensis was more abundant than all native species combined (Guénard and Dunn, 2010). It also found that the diversity and abundance of native ant species (and some other species) were negatively associated with the presence and abundance of B. chinensis.

A strong negative relationship between the abundance of B. chinensis and species of the keystone seed dispersing ant (Aphaenogaster spp.) has been observed, with similar patterns for the small species of the subfamilies Formicinae and Myrmicinae, and litter-foraging ant species (Guénard and Dunn, 2010). Species density for larger species from the genera Camponotus and Formica responded positively for low to medium abundance of B. chinensis, and had similar species density to the control areas even for large densities of B. chinensis (Guénard and Dunn, 2010).

In North Carolina, USA, abundance of the native species complex Aphaenogaster rudis [Aphaenogaster picea rudis] has been found to be strongly negatively correlated with abundance of B. chinensis (Bednar, 2010). It particularly displaces A. rudis from woodland habitats (Bednar, 2010). Displacement was due to direct (aggressive encounters) and indirect effects (dominating food sources, primarily termite nests) (Bednar, 2010). In most aggressive encounters, B. chinensis subdued A. rudis in approximately 60 seconds; it is suspected that B. chinensis venom is more toxic than that of A. rudis (Bednar, 2010).  

B. chinensis has been found to reduce the number of worker ants of A. rudis by 96%, and as a result reduced the number of plant seeds removed from plots by 70% (Rodriguez-Cabal et al., 2012). In the same study, it was found that a locally abundant myrmecochrous plant was 50% less abundant where B. chinensis was present, suggesting that the ant is disrupting ant-seed dispersal mutualisms (Rodriguez-Cabal et al., 2012).    

In a study in Georgia, USA, B. chinensis colonized artificial nests in deciduous forests, displacing the native A. rudis that shares ecological niche requirements (temperature, moisture and requiring coarse wooden material for nesting colonies) (Warren et al., 2015). In the same study it was found that B. chinensis failed to disperse seed in the same way as A. rudis. Plant aggregation analysis for Asarum arifolium showed that plants were clustered more closely together in the presence of B. chinensis (Warren et al., 2015).

The abundance of B. chinensis is negatively correlated with the native species Paratrechina faisonensis in Fant’s Grove, South Carolina, possibly due to competition for nest sites (Paysen, 2007).  

In Morrisville, North Carolina, USA, B. chinensis was found to displace another invasive ant species, Linepithema humile (Rice and Silverman, 2013).

Threatened Species

Social Impact

B. chinensis has a venomous sting that can cause severe allergic reactions and anaphylactic shock in humans, posing a threat if it moves into areas close to human activity (Fukuzawa et al., 2002; Nelder et al., 2006). It is not an aggressive stinger, with cases occurring when a colony is disturbed or when a winged-female becomes trapped between the clothing and skin (MacGown et al., 2013). Symptoms of stings can include moderate swelling, sweating, light-headedness, severe pain, stinging, and a burning sensation (Guénard et al., 2018) which can last from 30 minutes up to 48 hours (B. Guénard, Hong Kong, personal communication, 2019). Stinging events may be particularly frequent during summer swarming; in one study a peak of stinging activity was found from May to July (Guénard et al., 2018).

The major allergens of B. chinensis are 23 and 25 kDa proteins that belong to the antigen 5 family of proteins (Lee et al., 2009).

B. chinensis can infest homes, school cafeterias and residential kitchens where they have been seen taking food (Rice and Waldvogel, 2017).

Risk and Impact Factors

• Proved invasive outside its native range
• Has a broad native range
• Abundant in its native range
• Highly mobile locally

• Changed gene pool/ selective loss of genotypes
• Damaged ecosystem services
• Ecosystem change/ habitat alteration
• Modification of successional patterns
• Negatively impacts human health
• Reduced native biodiversity
• Threat to/ loss of native species

• Causes allergic responses
• Competition - monopolizing resources
• Interaction with other invasive species
• Predation

• Highly likely to be transported internationally accidentally

Similarities to Other Species/Conditions

Distinguishing B. chinensis from other ant species in Japan:

B. nakasujii is common in Honshu, Shikoku, and Kyushu, Japan, where it is sympatric with B. chinensis (Yashiro et al., 2010). B. nakasujii is easy to distinguish from B. chinensis; the B. nakasujii worker has a proportionally wider petiole and the male has much darker coloration and more well-developed mandibles. The B. nakasujii worker is easily distinguished from another closely related species, B. luteipes, by B. nakasujii having a proportionally wider petiole and a sculptured propodeum (largely smooth in B. luteipes) (Yashiro et al., 2010). Although B. nakasujii and B. chinensis are sympatrically distributed in temperate zones, B. nakasujii is rare in dry and disturbed area, where B. chinensis remains common. Both species have been observed to nest near termite nests frequently and to carry termites back to their nests as food (Yashiro et al., 2010).

Distinguishing B. chinensis from other ant species in the USA, amended from MacGown (2016):

This species is similar in size to average-sized fire ants, but differs by having only only node and by its nesting in wooded habitats. In the Southeast USA, it could be possibly be confused with other ponerine ants (e.g. Hypoponera, Leptogenys, Pachcondyla, and Ponera), but differs by the the promesonotum being distinctly set off from the propodeum, by the very shiny mesopleura, and by large super colonial nesting behaviour. 

A further distinguishing feature from other species in the USA is its inability to cling to smooth surfaces such as glass (Rice and Waldvogel, 2017).

Prevention and Control

Due to the variable regulations around (de)registration of pesticides, your national list of registered pesticides or relevant authority should be consulted to determine which products are legally allowed for use in your country when considering chemical control. Pesticides should always be used in a lawful manner, consistent with the product's label.

Biological Control

Soldiers of Reticulitermes speratus produce the terpenoid hydrocarbon β-selinene as a defence against predators such as B. chinensis (Iida and Akino, 2016) and investigations have shown that B. chinensis avoids this chemical (Iida and Akino, 2016).

Chemical Control

B. chinensis will scavenge on protein-based insecticide baits, with suitable insecticides including hydramethylnon, s-methoprene [methoprene], indoxacarb and thiamethoxam (Rice and Waldvogel, 2017). It is suggested that starting treatments in April (if B. chinensis activity is observed) may help reduce the impact of the baits on non-target ant species (Rice and Waldvogel, 2017).

Granular bait containing hydramethylnon dispersed in the territory of B. chinensis reduced populations one day after application, and the effects were still apparent 28 days after application (Rice et al., 2012).  

Prey-baiting was used to control B. chinensis in laboratory and field investigations (Buczkowski, 2016; Buczkowski, 2017). Live fipronil-treated termites (Reticulitermes flavipes) were used as bait and were highly effective against B. chinensis, while detrimental effects against non-target species were minimal (Buczkowski, 2017).  

The terpenoid hydrocarbon β-selinene is produced by Reticulitermes speratus as a chemical defence against predators such as B. chinensis (Iida and Akino, 2016); it is therefore a potential chemical control agent.

At the National Zoological Park in Washington DC, USA, granular carbohydrate and protein baits were used to try and control B. chinensis but proved to be largely ineffective (Hiza et al., 2012). B. chinensis control was more effective when chemical sprays were used on foragers, nests and travel routes and when it was repeated the following year (Hiza et al., 2012).

Gaps in Knowledge/Research Needs

More research is needed on the impact of B. chinensis on seed dispersal mutualisms and ecosystem function (Rodriguez-Cabal et al., 2012).

More information on the seasonality of colony structure in the USA is required if an integrated pest management programme is to be developed (Zungoli and Benson, 2008).

References

Allen HR, 2017. Biology and behavior of the Asian Needle Ant, Brachyponera chinensis. DPhil Thesis. Clemson, South Carolina, USA: Clemson University. https://tigerprints.clemson.edu/all_dissertations/2084

AntWeb, 2019. Species: Brachyponera chinensis (Emery, 1895). https://www.antweb.org/description.do?genus=brachyponera&species=chinensis&rank=species&countryName=Japan

Bednar DM, 2010. Pachycondyla (=Brachyponera) chinensis predation on Reticulitermes virginicus and competition with Phaenogaster rudis. Raleigh, North Carolina, USA: North Carolina State University. 65. http://www.lib.ncsu.edu/resolver/1840.16/6363

Bednar, D. M., Silverman, J., 2011. Use of termites, Reticulitermes virginicus, as a springboard in the invasive success of a predatory ant, Pachycondyla (=Brachyponera) chinensis. Insectes Sociaux, 58(4), 459-467. http://www.springerlink.com/content/1420-9098 doi: 10.1007/s00040-011-0163-0

Bertelsmeier, C., Guénard, B., Courchamp, F., 2013. Climate change may boost the invasion of the Asian needle ant. PLoS ONE, 8(10), e75438. http://www.plosone.org/article/info%3Adoi%2F10.1371%2Fjournal.pone.0075438 doi: 10.1371/journal.pone.0075438

Brown WL, 1958. A review of the ants of New Zealand. Acta Hymenopterologica, 1(1), 1-50.

Buczkowski, G., 2016. The Trojan horse approach for managing invasive ants: a study with Asian needle ants, Pachycondyla chinensis. Biological Invasions, 18(2), 507-515. http://link.springer.com/article/10.1007%2Fs10530-015-1023-z doi: 10.1007/s10530-015-1023-z

Buczkowski, G., 2017. Prey-baiting as a conservation tool: selective control of invasive ants with minimal non-target effects. Insect Conservation and Diversity, 10(4), 302-309. http://onlinelibrary.wiley.com/journal/10.1111/(ISSN)1752-4598 doi: 10.1111/icad.12230

Eyer P-A, Matsuura K, Vargo EL, et al., 2018. Inbreeding tolerance as a pre-adapted trait for invasion success in the invasive ant Brachyponera chinensis. Molecular Ecology, 27, 4711– 4724. doi: https://doi.org/10.1111/mec.14910

Forel A, 1900. (Fourmis du Japon. Nids en toile. Strongylognathus Huberi et voi-sins. Fourmilière triple. Cyphomyrmex Wheeleri. Fourmis importées). Bulletin de la Société Entomologique Suisse, 10, 267–287.

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Guénard B, Dunn R R, 2010. A new (old), invasive ant in the hardwood forests of Eastern North America and its potentially widespread impacts. PLoS ONE. e11614. http://www.plosone.org/article/info%3Adoi%2F10.1371%2Fjournal.pone.0011614 DOI:10.1371/journal.pone.0011614

Guénard B, Wetterer JK, MacGown JA, 2018. Global and temporal spread of a taxonomically challenging invasive ant, Brachyponera chinensis (Hymenoptera: Formicidae). In: Florida Entomologist, 101 (4) 649-656.

Harada Y, Matsumoto Y, Maeda S, Oyama A, Yamane S, 2009. Comparison of ant fauna among different habitats of Yaku-shima Island, southern Japan. Bulletin of the Biogeographical Society of Japan. 125-134. http://www.soc.nii.ac.jp/tbsj/index.htm

Mackay WP, Mackay EE, 2010. The systematics and biology of the new world ants of the genus Pachycondyla (Hymenoptera: Formicidae)., Lewiston, New York, USA: Edwin Mellen Press.

Miner A, 2014. Pachycondyla chinensis. In: Animal Diversity Web, Ann Arbor, USA: University of Michigan. https://animaldiversity.org/accounts/Pachycondyla_chinensis/

Nelder M P, Paysen E S, Zungoli P A, Benson E P, 2006. Emergence of the introduced ant Pachycondyla chinensis (Formicidae: Ponerinae) as a public health threat in the Southeastern United States. Journal of Medical Entomology. 43 (5), 1094-1098. http://docserver.esa.catchword.org/deliver/cw/pdf/esa/freepdfs/00222585/v43n5s44.pdf DOI:10.1603/0022-2585(2006)43[1094:EOTIAP]2.0.CO;2

Paysen ES, 2007. Diversity and abundance of ants at forest edges in the Great Smoky Mountains National Park., Clemson, South Carolina, USA: Clemson University. https://tigerprints.clemson.edu/all_dissertations/174

Rice ES, Silverman J, 2013. Propagule pressure and climate contribute to the displacement of Linepithemahumile by Pachycondyla chinensis. In: PLoS One, 8 (2) e56281. https://journals.plos.org/plosone/article?id=10.1371/journal.pone.0056281

Smith MR, 1934. Ponerine ants of the genus Euponera in the United States. In: Annals of the Entomological Society of America, 27 (4) 557–564. https://academic.oup.com/aesa/article-abstract/27/4/557/20127?redirectedFrom=fulltext

Yashiro T, Matsuura K, Guénard B, Terayama M, Dunn R R, 2010. On the evolution of the species complex Pachycondyla chinensis (Hymenoptera: Formicidae: Ponerinae), including the origin of its invasive form and description of a new species. Zootaxa. 39-50. http://www.mapress.com/zootaxa/2010/f/z02685p050f.pdf

Zungoli P A, Benson E P, 2008. Seasonal occurrence of swarming activity and worker abundance of Pachycondyla chinensis (Hymenoptera: Formicidae). In: 6th International Conference on Urban Pests, Budapest, Hungary, 13-16 July 2008 [6th International Conference on Urban Pests, Budapest, Hungary, 13-16 July 2008.], [ed. by Robinson W H, Bajomi D]. UK, UK: International Conference on Urban Pests (ICUP). 51-57.

Contributors

13/12/18 Original text by:

Vicki Cottrell, Consultant, UK

Distribution Maps