Utility of the CLIMEX ‘match climates regional’ algorithm for pest risk analysis: an evaluation with non-native ants in New Zealand

Pest risk analysts frequently ask if the climate of a pest risk analysis area could be suitable for the establishment of an organism of concern. Species distribution models can help to answer this question, but constructing them is technically complex, time consuming, and uninformative for additional non-modelled species. A quicker more broadly applicable approach involves using environmental distance metrics, including climate matching algorithms such as the ‘match climates regional’ function of CLIMEX (CLIMEX-MCR), to generate indices of climatic similarity between different locations without reference to particular species. Several studies have shown that various environmental distance metrics can provide biologically meaningful results. However, the veracity of the CLIMEX-MCR algorithm remains unevaluated, despite its application in numerous published studies. We used CLIMEX-MCR and high resolution New Zealand climate data to measure climatic similarities between New Zealand and the rest of the world. We then tested the veracity of the climatic match estimates by evaluating if their predictions regarding the suitability of New Zealand’s climate for 43 non-native ant species corresponded with empirical observations of those species in New Zealand. Non-native ants that are, or were once, established outdoors in New Zealand had overseas distributions that were climatically well matched with New Zealand. In contrast, species that either are established only indoors in New Zealand, or were observed to temporarily nest outdoors then die in New Zealand, had overseas distributions that were poorly matched. Species that are frequently intercepted at New Zealand’s border, but are not established there, generally also had overseas distributions with low climatic similarities to New Zealand. We also measured climatic similarities between New Zealand’s 13 national parks and the rest of the world. The overseas distributions of the non-native ants showed poor climatic matches with New Zealand’s national parks, which was consistent with the absence of persistent outdoor non-native ant populations in those parks. Our results support the utility of CLIMEX-MCR algorithm for pest risk analysis.     

Modelling the potential geographic distribution of invasive ant species in New Zealand

Despite their economic and environmental impacts, there have been relatively few attempts to model the distribution of invasive ant species. In this study, the potential distribution of six invasive ant species in New Zealand are modelled using three fundamentally different methods (BIOCLIM, DOMAIN, MAXENT). Species records were obtained from museum collections in New Zealand. There was a significant relationship between the length of time an exotic species had been present in New Zealand and its geographic range. This is the first time such a time lag has been described for exotic ant species, and shows there is a considerable time lag in their spread. For example, it has taken many species several decades (40–60 years) to obtain a distribution of 17–25% of New Zealand regions. For all six species, BIOCLIM performed poorly compared to the other two modelling methods. BIOCLIM had lower AUC scores and higher omission error, suggesting BIOCLIM models under-predicted the potential distribution of each species. Omission error was significantly higher between models fitted with all 19 climate variables compared to those models with fewer climate variables for BIOCLIM, but not DOMAIN or MAXENT. Widespread species had a greater commission error. A number of regions in New Zealand are predicted to be climatically suitable for the six species modelled, particularly coastal and lowland areas of both the North and South Islands.     

Inferring historical introduction pathways with mitochondrial DNA: the case of introduced Argentine ants (Linepithema humile) into New Zealand

The threat imposed by invasive species and difficulties associated with control and management places more impetus on trying to prevent their introduction. The identification of introduction pathways is a vital component towards this goal. In this study, we use a genetic marker-based approach to retrospectively investigate the pathway of origin of the invasive Argentine ant ( Linepithema humile ) into New Zealand. We intensively sample the mitochondrial gene cytochrome b , from the entire known range of Argentine ants in New Zealand. No genetic variation was found in New Zealand. In order to identify likely introduction pathways, we use two alternative genetic analyses and suggest that a tcs approach that collapses identical haplotypes and calculates the probability of parsimony is superior to standard phylogenetic tree-building algorithms. A minimum spanning network allowed relationships to be examined among sequences collated from previous international studies. The cytochrome b sequence, when compared to a global database, matched that from an Australian population. That Australia is the potential source of Argentine ants is in agreement with the New Zealand interception record, as goods from Australia have the highest number of interception records of Argentine ants. Our approach can easily be duplicated for other organisms and the methodology can be more widely applied to help aid further efforts to identify the routes of transmission for other invasive species and allow us to efficiently direct our biosecurity monitoring effort.     

The worldwide transfer of ants: geographical distribution and ecological invasions

Aim This is the first comprehensive account of the biogeography of ants transferred and at least temporarily established outside their native habitat. Location Using museum and literature records, I established the distributions of transferred ant species. Methods I used taxonomic and functional groups to assess how geographical spread as a transferred species is affected by taxonomy and life history. Results 147 ant species in forty-nine genera have been recorded outside of their native habitat. The proportion of transferred ants is similar to the number of genera and species in each subfamily. The species-rich subfamily Myrmicinae contains nearly 50% of all transferred species, while many of the species-poor subfamilies have absolutely no transferred species. A disproportionate high number of transferred ants originate from the Neotropical and Oriental biogeographic regions. The Pacific Islands are the recipients of the most transferred ant species. Most transferred ants belong to the CRYPTIC, OPPORTUNIST, and GENERALIZED MYRMICINE functional groups, while there are no recorded transfers of army ants or leaf-cutting ants. Both invasive and human commensal ‘tramp’ ant species are nonrandom subsets of transferred ants. Main conclusions ‘Tramp’ species and invasive species tend to have widespread geographical distributions, and share life history characteristics including queen number, nest structure, and foraging behaviour. Combining observations of functional groups and biogeography may lead to a better understanding of the factors contributing to the spread of transferred species.     

Invasive ants carry novel viruses in their new range and form reservoirs for a honeybee pathogen

When exotic animal species invade new environments they also bring an often unknown microbial diversity, including pathogens. We describe a novel and widely distributed virus in one of the most globally widespread, abundant and damaging invasive ants (Argentine ants, Linepithema humile). The Linepithema humile virus 1 is a dicistrovirus, a viral family including species known to cause widespread arthropod disease. It was detected in samples from Argentina, Australia and New Zealand. Argentine ants in New Zealand were also infected with a strain of Deformed wing virus common to local hymenopteran species, which is a major pathogen widely associated with honeybee mortality. Evidence for active replication of viral RNA was apparent for both viruses. Our results suggest co-introduction and exchange of pathogens within local hymenopteran communities. These viral species may contribute to the collapse of Argentine ant populations and offer new options for the control of a globally widespread invader.     

Long-term impact of exotic ants on the native ants of Madeira

Abstract.  1. The earliest exotic records for two notorious invasive ants, the big-headed ant ( Pheidole megacephala ) and the Argentine ant ( Linepithema humile ), both come from the Atlantic islands of Madeira, where the two species underwent population explosions in the 1850s and 1890s respectively. Researchers have long assumed that these invaders spread across all of Madeira and exterminated most or all native ants, despite no research actually documenting such impact.
2. Re-examination of first-hand nineteenth century accounts suggest that P. megacephala and L. humile may never have spread beyond coastal lowland areas, representing < 10% of Madeira's land area. In 2002, native ants dominated most of Madeira; P. megacephala and L. humile were restricted to ≈ 0.3% and ≈ 6% of Madeira's land area respectively.
3. Of the 10 native ant species known from Madeira, only one ( Temnothorax wollastoni ) was not present in 1999–2002 surveys. Although exotic ants may have exterminated T. wollastoni , it seems likely that this species still survives.
4. Thus, even after 150 or more years of residence, P. megacephala and L. humile have come to occupy only a small part of Madeira, and appear to have had little impact.
5. Most of Madeira may be too cool for P. megacephala and perhaps too moist for L. humile to dominate. Also, Madeira's vast natural areas may generally lack weedy vegetation that can support high densities of plant-feeding Hemiptera critical for the ecological dominance of invasive ants. Finally, a dominant native ant, Lasius grandis , inhabiting ≈ 84% of Madeira, may actively exclude P. megacephala and L. humile .     

Relative risk of invasive ants (Hymenoptera: Formicidae) establishing in New Zealand

Abstract

Human‐assisted spread of species poses a major challenge to border security agencies. Ideally, limited resources need to be targeted at species posing the most risk. Climate matching is an important component of assessing risk but often little or no biological information is available to enable detailed modelling. To assess if distribution records alone provide useful establishment predictions we compare the climate in New Zealand and its outlying islands to that found in the current native and introduced ranges of 12 tramp ant species, three of which are already established in New Zealand, using the climate module of BIOSECURE, a risk assessment tool. Eleven species showed a similar general pattern, with mean annual temperature being the climatic variable with the least overlap between the estimated realised niche and the predicted New Zealand niche. The twelfth species, of temperate origin, is less restricted by temperature, but much of New Zealand may have too high a rainfall. The present and future threats posed by tramp ants are discussed in relation to climate limitations.     

Ant diversity and distribution in Acadia National Park, Maine

Exotic ant species are a primary threat to ant biological diversity, posing a negative impact to native ant communities. In this study, we examine species richness of ants (family Formicidae) in Acadia National Park, ME, as a fundamental step toward understanding the present impact of the exotic species Myrmica rubra on native ant species. Twelve habitat types were sampled, along six transects, with pitfall traps, visual searching, bait traps, and leaf litter extraction, and the aid of 34 volunteers. We report 42 species of ants in Acadia National Park, comprising five subfamilies (Amblyoponinae, Dolichoderinae, Formicinae, Myrmicinae, and Ponerinae) and 15 genera; the cataloged species represents 75% of the species originally recorded in the area by Procter (1946). Our findings suggest M. rubra is currently not a dominant species throughout the entire island. However, where this species has invaded locally, few competing native species coexist. The species Lasius alienus, Formica subsericea, Myrmica detritinodis, Camponotus herculeanus, Formica argentea, Formica aserva, and Tapinoma sessile occurred most often in our survey. We report the ant species Amblyopone pallipes and Dolichoderus mariae as two new records for the state of Maine.     

Ants of Ambon Island – diversity survey and checklist

The present checklist of ants (Hymenoptera: Formicidae) of Ambon is the first comprehensive overview of ant species recorded on the island during the last 150 years. The species list is based on literature and museum collections’ records combined with data from our field survey in 2010. In total, there are 74 ant species and subspecies representing 34 genera and six subfamilies known from Ambon. Five of the species found in undisturbed forest were exotic and indicate the overall habitat degradation on the island. The largest proportion of Ambon ant fauna are species with affinities to the Oriental region and species of Oriental-Austro-Melanesian origin. At least 20% of the species are regional endemics. In comparison to other islands in the region, the Ambon fauna seems more diverse and better sampled; however it is clear that a large part of it still remains to be described.     

New records of ant species from Yunnan,China

As with many other regions of the world, significant collecting, curation, and taxonomic efforts will be needed to complete the inventory of China’s ant fauna. This is especially true for the highly diverse tropical regions in the south of the country, where moist tropical forests harbor high species richness typical of the Southeast Asian region. We inventoried ants in the Xingshuangbanna prefecture, Yunnan, in June 2013, using a variety of methods including Winkler extraction and hand collection to sample ant diversity. We identified 213 species/morphospecies of ants from 10 subfamilies and 61 genera. After identification of 148 valid species of the 213 total species collected, 40 species represent new records for Yunnan province and 17 species are newly recorded for China. This increases the total number of named ant species in Yunnan and China to 447 and 951 respectively. The most common species collected were Brachyponera luteipes and Vollenhovia emeryi. Only one confirmed exotic species Strumigenys membranifera, was collected, although several others were potentially introduced by humans. These results highlight the high biodiversity value of the region, but also underscore how much work remains to fully document the native myrmecofauna.     

Establishment patterns of non-native insects in New Zealand

Insects comprise the majority of non-native animal species established around the world. However, geographic biases in knowledge hamper an overall understanding of biological invasions globally. A dataset of accidentally introduced non-native insect species established in New Zealand was compiled from databases, entomological literature, and examination of specimens in the New Zealand Arthropod Collection. For each non-native species, the first recorded location and first recorded date of detection was obtained. Excluding intentionally introduced species, there are 1477 non-native insect species successfully established in New Zealand across 16 orders, 234 families and 1017 genera. Four orders (Coleoptera, Hemiptera, Hymenoptera and Diptera) contributed 77.5% of all established insect species. Herbivores represented the largest feeding guild (47.7%), comprised of polyphagous (48.3%) or oligophagous (39.7%) species. The majority of these species originated in the Australasian (36.7%) and Palearctic regions (24.8%). Regression trees, using a binary recursive partitioning approach, found the number of international tourist arrivals, exotic vegetation cover, and regional gross domestic product were the main factors explaining spatial patterns of recently established species. Gross domestic product best explained temporal patterns of establishment over the last century. Our findings demonstrate that broad-scale analyses of non-native species have important applications for border biosecurity by providing insight into the extent of invasions. In New Zealand, the current trajectory indicates fewer non-native species are establishing annually, suggesting biosecurity efforts are being effective at reducing rates of establishment.     

Exploitation of Fallen Diaspores by Ants: Are there Ant–Plant Partner Choices?

Ants frequently interact with fleshy fallen diaspores (fruits or seeds) not adapted for ant‐dispersal. Such interactions are usually considered as opportunistic, but recent evidence has indicated that these ants may differ in their effects on diaspore survival and plant recruitment. We investigated if partner choices are recognizable among genera of ants and plants, and if ant and plant traits may influence such preferences in cerrado (savanna‐like vegetation) from southeast Brazil. During a 2‐yr period, 521 ant–diaspore interactions were recorded through various methods, yielding 71 ant species and 38 plant species. Exploitation of fallen diaspores was common among several ant genera, and included carnivorous, omnivorous, and fungivorous ants. Contrary to others areas around the world, where true myrmecochory (seed dispersal by ants) is common among shrubs, ants also exploited diaspores from several cerrado trees. Plant life form, diaspore size, and ant body size did not seem to explain the pattern of interactions observed. Two subsets of preferential interactions, however, segregated fungivorous ants from another group composed of carnivorous and omnivorous ants, probably influenced by the chemical composition of the plant diaspores. Omnivorous ants usually remove the fleshy portion of diaspores on spot and probably provide limited benefits to plants. Carnivorous and fungivorous ants usually remove the whole diaspore to the nest. As each of these ant groups may influence the fitness of diaspores in different ways, there are possible subtle pathways for the evolution of partner choices between ants and these non‐myrmecochorous diaspores.     

Large scale unicoloniality: the population and colony structure of the invasive Argentine ant (<Emphasis Type="Italic">Linepithema humile</Emphasis>) in New Zealand

The Argentine ant is native to South America and has spread widely across the globe. In this study, we use genetic analyses and behavioural assays to examine the colony structure of Argentine ants in New Zealand. Diet modification studies were also carried out in order to help identify what factors influence these behavioural assays. There was no aggression observed between any pairings tested across the North Island of New Zealand, though we found that diet manipulations in the laboratory could lead to low levels of aggression between previously amiable Argentine ant nests. The New Zealand population of Argentine ants was characterized by low levels of genetic variation in six microsatellite loci from their nuclear genome. Additionally, the population also lacked significant genetic structuring with no patterns of regional differentiation or isolation by distance. An analysis of molecular variation(AMOVA) found that themajority of genetic variation was present at a nest level (93% of total genetic variance), with little genetic differentiation observed within or between regions (3 – 4% of total genetic variance). No correlation between aggression and genetic relatedness was observed. This evidence suggests that Argentine ants in New Zealand effectively form a unicolonial population, which is likely the result of colonization from a single source population. As far as we know, this is the first country to have an entirely unicolonial population of Argentine ants. Received 21 February 2007; revised 20 May 2007; accepted 22 May 2007.     

Frugivore gape size and the evolution of fruit size and shape in southern hemisphere floras

Abstract The present study uses differences among frugivore faunas of the southern hemisphere landmasses to test whether frugivore characteristics have influenced the evolution of fruit traits. Strong floristic similarities exist among southern landmasses; for example, 75% of New Zealand vascular plant genera also have species in Australia. However, plants in Australia and South America have evolved in the presence of a range of mammalian frugivores, whereas those in New Zealand, New Caledonia and the Pacific Islands have not. In addition, the avian frugivores in New Zealand and New Caledonia are generally smaller than those of Australia. If frugivore characteristics have influenced the evolution of fruit traits, predictable differences should exist between southern hemisphere fruits, particularly fruit size and shape. Fruit dimensions were measured for 77 New Zealand species and 31 Australian species in trans‐Tasman genera. New Zealand fruits became significantly more ellipsoid in shape with increasing size. This is consistent with frugivore gape size imposing a selective pressure on fruit ingestability. This result is not a product of phylogenetic correlates, as fruit length and width scaled isometrically for Australian species in genera shared with New Zealand. Within‐genus contrasts between New Zealand and Australian species in 20 trans‐Tasman genera showed that New Zealand species have significantly smaller fruits than their Australian counterparts. Within‐genus contrasts between New Zealand and South American species in nine genera gave the same result; New Zealand species had significantly smaller fruits than their South American counterparts. No difference was found in fruit size or shape between New Zealand and New Caledonia congeneric species from 12 genera. These results are consistent with the broad characteristics of the frugivore assemblage influencing the evolution of fruit size and shape in related species. The smaller‐sized New Zealand frugivore assemblage has apparently influenced the evolution of fruit size of colonizing taxa sometimes within a relatively short evolutionary timeframe.     

A review of Bahamian ant (Hymenoptera: Formicidae) biogeography

Abstract. The results of recent surveys for ants on Staniel Cay, Exumas (and surrounding small cays) and the northeastern region of Andros are presented. These records are compiled with all previously published collection records of ants from the Bahamas into a single database. A list of all known Bahamian species is presented, in accord with current taxonomy. Distributions within the Bahamas are given, along with the status of each species with reference to its origin (endemic, native, or exotic). At present seventy-five named species of ants are known from the Bahamas, which shares 60% of its ant fauna with Florida and 71% with Cuba. More than one third of the species (37%) are exotics, which appear to exist primarily in areas characterized by human disturbance, and apparently have not invaded the native vegetation to a large degree. Both historical and ecological factors are likely to have played important roles in shaping present-day Bahamian ant species diversity and distribution. The number of species on all major Bahamian islands from which ants have been collected is presented as a function of island area. Most islands have not been very thoroughly surveyed. The species-area relationship for the more thoroughly surveyed islands is relatively flat (z= 0.06). Further collecting in this archipelago will probably lead to the discovery of additional species.     

Associations between lycaenid butterflies and ants in Australia

A comprehensive and critical review of all available literature on associations between Australian lycaenid butterflies and ants was undertaken to establish an accurate database of the partners involved. Collections and observations of lycaenids and ants were used to augment this review, resulting in a significant number of newly documented association (and non-association) records. Twenty published records considered to be erroneous or doubtful are noted, with justifications given for their deletion from the association database. In total, 265 different associations between lycaenids and ants, plus 65 non-attendance records are documented for Australia. Nearly 80% of the lycaenid species in Australia, for which the early stages are known, are recorded associating with ants and half of these are obligately ant-associated. Patterns of association are examined from the perspective of both lycaenids and ants, with a focus on ant systematics and ecology. Lycaenids are recorded with five ant subfamilies, including the first record of an association with the Pseudomyrmecinae. The Dolichoderinae, and to some extent the Formicinae, have a disproportionately high percentage of genera that associate with lycaenid butterflies. All ant species that tend lycaenids spend at least some portion of their time foraging on vegetation to collect plant and insect nectar. There is a robust relationship between the competitive status of ants within a community, and their frequency and degree of association with lycaenids. Obligate ant-association is accompanied by a high degree of specificity for ant partner, but two notable exceptions, Ogyris aenone and O. amaryllis are discussed. Facultative myrmecophiles tend to associate with a broad range of ants, although interactions with ecologically dominant ants are less frequent than might be expected based on the abundance of dominant ant species in Australian communities.     

Determinants for the successful establishment of exotic ants in New Zealand

Biological invasions can dramatically alter ecosystems. An ability to predict the establishment success for exotic species is important for biosecurity and conservation purposes. I examine the exotic New Zealand ant fauna for characteristics that predict or determine an exotic species’ ability to establish. Quarantine records show interceptions of 66 ant species: 17 of which have established, 43 have failed to establish, whereas nests of another six are periodically observed but have failed to establish permanently (called ‘ephemeral’ establishment). Mean temperature at the highest latitude and interception variables were the only factors significantly different between established, failed or ephemeral groups. Aspects of life history, such as competitive behaviour and morphology, were not different between groups. However, in a stepwise discriminant analysis, small size was a key factor influencing establishment success. Interception rate and climate were also secondarily important. The resulting classification table predicted establishment success with 71% accuracy. Because not all exotic species are represented in quarantine records, a further discriminant model is described without interception data. Though with less accuracy (65%) than the full model, it still correctly predicted the success or failure of four species not used in the previous analysis. Techniques for improving the prediction accuracy are discussed. Predicting which species will establish in a new area appears an achievable goal, which will be a valuable tool for conservation biology.     

Defensive traits exhibit an evolutionary trade‐off and drive diversification in ants

Evolutionary biologists have long predicted that evolutionary trade‐offs among traits should constrain morphological divergence and species diversification. However, this prediction has yet to be tested in a broad evolutionary context in many diverse clades, including ants. Here, we reconstruct an expanded ant phylogeny representing 82% of ant genera, compile a new family‐wide trait database, and conduct various trait‐based analyses to show that defensive traits in ants do exhibit an evolutionary trade‐off. In particular, the use of a functional sting negatively correlates with a suite of other defensive traits including spines, large eye size, and large colony size. Furthermore, we find that several of the defensive traits that trade off with a sting are also positively correlated with each other and drive increased diversification, further suggesting that these traits form a defensive suite. Our results support the hypothesis that trade‐offs in defensive traits significantly constrain trait evolution and influence species diversification in ants.     

How many and which ant species are being accidentally moved around the world?

Human transportation facilitates the dispersal of exotic ants, but few studies have quantified the magnitude and geography of these movements. We used several non-parametric indexes to estimate the number of species successfully introduced to or established in new regions. We also compared their source biogeographic realms to assess the importance of the geographical origin in determining the likelihood of establishment after introduction. Data on exotic ants derive from studies of three temperate regions. Our results suggest that the numbers of introduced or established ants may be much larger than the numbers so far documented. Ants introduced or established in new regions tend to arrive from the same or neighbouring realms, as would be expected if exotic species tend to match climates and if arrival/establishment is dependent upon higher trade rates from neighbouring countries.