Larval morphology and host use confirms ecotypic variation in Cactoblastis cactorum (Berg)

Despite their recognized importance in the literature, the contribution of native-range species interactions to invasion success has been inadequately studied. Previous authors have suggested that biases in the sampling of propagules from the native range might influence invasion success, but most contemporary invasion hypotheses focus on the development of novel interactions or a release from native consumers and competitors. When ecotypic variation exists in native host-consumer associations, the specific pattern of sampling across ecotypes could determine invasion success, especially when the genetic diversity among exotic propagules is low. The South American cactus moth, Cactoblastis cactorum (Berg), is an oligophagous consumer whose larvae feed on prickly pear cacti (subfamily Opuntioideae). The moth was collected from a small geographic area along the Argentina-Uruguay border in 1925 and was introduced to multiple continents as a biological control species, which has subsequently invaded North America. Here we show that groups defined by genetic structure in this species’ native range are concordant with distinct patterns of host association and larval morphology. Furthermore, in Florida populations, morphological traits have diverged from those found in the native range, and patterns of host association suggest that strong biases in host preference also occur in invasive populations. The documented history of C. cactorum introductions confirms that multiple attempts were made to export the moth, but that only a single ecotype was exported successfully. Additional work will be necessary to determine whether the observed host biases in North America reflect a rapid adaptation to naïve hosts or a conservation of traits related to specific aspects of the host-consumer association.     

Herbivory-induced extrafloral nectar increases native and invasive ant worker survival

Ascertaining the costs and benefits of mutualistic interactions is important for predicting their stability and effect on community dynamics. Despite widespread designation of the interaction between ants and extrafloral nectaries (EFNs) as a mutualism and over 100 years of studies on ant benefits to plants, the benefits to ants have never been experimentally quantified. The success of invasive ants is thought to be linked to the availability of carbohydrate-rich resources, though reports of invasive ant visits to EFNs are mixed. In two laboratory experiments, we compared worker survival of one native (Iridomyrmex chasei) and two invasive ant species (Linepithema humile and Pheidole megacephala) exposed to herbivorized or non-herbivorized EFN-bearing plants (Acacia saligna) or positive and negative controls. We found that non-herbivorized plants did not produce any measurable extrafloral nectar, and ants with access to non-herbivorized plants had the same survival as ants with access to an artificial plant and water (unfed ants). Ants given herbivorized plants had 7–11 times greater worker survival relative to unfed ants, but there were no differences in survival between native and invasive ants exposed to herbivorized plants. Our results reveal that ants cannot induce A. saligna extrafloral nectar production, but workers of both native and invasive ant species can benefit from extrafloral nectar as much as they benefit from sucrose.     

The influence of extrafloral nectaries on parasitism of an insect herbivore

The extrafloral nectaries of many plants promote ant defense against insect herbivores. We examined the influence of extrafloral nectaries on the levels of parasitism of a generalist insect herbivore, the gypsy moth (Lymantria dispar L.). Larvae and pupae of the moth were collected from trees with and without extrafloral nectaries growing in the same forests in South Korea and reared to evaluate parasitism. More parasitism occurred on plants with extrafloral nectaries in seven of the nine season-long collections at the six sites and in four out of five collecting periods. Parasitism was higher on the four main genera of plants with extrafloral nectaries than on any of five main genera of plants without extrafloral nectaries. There was no difference in parasitoid richness; nine species occurred in each group, eight of which were the same. There was a positive and almost significant correlation between the abundance of plants with extrafloral nectaries and the parasitism of gypsy moth at the sites. Extrafloral nectaries may reduce herbivory by inducing more parasitism of the insect herbivores that attack plants bearing the glands.     

Ant predation on an invasive herbivore: can an extrafloral nectar-producing plant provide associational resistance to <Emphasis Type="Italic">Opuntia</Emphasis> individuals?

The legume Chamaecrista fasciculata attracts ants to its extrafloral nectar (EFN) which can lead to reduced herbivory and increased fecundity for the plant. In Florida, Opuntia stricta and O. humifusa, hosts of the invasive moth Cactoblastis cactorum, are often found growing in close association with C. fasciculata. We tested the hypotheses that O. stricta and O. humifusa individuals have higher ant abundance, lower levels of herbivore damage, and increased growth when growing in close association with C. fasciculata compared with individuals not growing near the plant. We also experimentally placed C. cactorum eggsticks and pupae on Opuntia individuals to see if ant predation of these stages occurred, and if so, whether predation rates were higher on individuals growing close to C. fasciculata. Opuntia plants near C. fasciculata were less likely to be attacked by C. cactorum and had higher ant abundance than plants far from C. fasciculata. Field surveys showed that Opuntia plants near C. fasciculata had a lower proportion of cladodes with C. cactorum damage of any type. Proportions of cladodes with damage from five native herbivores were not significantly different between treatments. In addition, Opuntia individuals growing near C. fasciculata added proportionately more pads during the growing season. We found evidence of ant predation on 15.9% of C. cactorum eggsticks and 17.6% of pupae. In August and October of 2008, there was significantly more evidence of predation on eggs and pupae placed on Opuntia individuals near C. fasciculata. No effect of distance to C. fasciculata was seen in November of 2008, potentially because plants were no longer producing EFN at this time. Our finding that Opuntia plants close to C. fasciculata show reduced herbivory from invasive C. cactorum, but not from the native herbivores examined, suggests that patterns of associational resistance may be influenced by the co-evolutionary history of the organisms in question.     

Infiltration of a facultative ant–plant mutualism by the introduced Argentine ant: effects on mutualist diversity and mutualism benefits

1. Ant–plant mutualisms have been the focus of considerable empirical research, but few studies have investigated how introduced ants affect these interactions. Using 2 years of survey data, this study examines how the introduced Argentine ant [Linepithema humile (Mayr)] differs from native ants with respect to its ability to protect the extrafloral nectary‐bearing coast barrel cactus (Ferocactus viridescens) in Southern California. 2. Eighteen native ant species visited cacti in uninvaded areas, but cacti in invaded areas were primarily visited by the Argentine ant. The main herbivore of the coast barrel cactus present at the study sites is a leaf‐footed bug (Narnia wilsoni). 3. Herbivore presence (the fraction of surveys in which leaf‐footed bugs were present on individual cacti) was negatively related to ant presence (the fraction of surveys in which ants were present on individual cacti). Compared with cacti in uninvaded areas, those in invaded areas were less likely to have herbivores and when they did had them less often. 4. Seed mass was negatively related to herbivore presence, and this relationship did not differ for cacti in invaded areas versus those in uninvaded areas. 5. Although the Argentine ant might provide superior protection from herbivores, invasion‐induced reductions in ant mutualist diversity could potentially compromise plant reproduction. The cumulative number of ant species on individual cacti over time was lower in invaded areas and was associated with a shortened seasonal duration of ant protection and reduced seed mass. These results support the hypothesis that multiple partners may enhance mutualism benefits.     

Geographic variation in a facultative mutualism: consequences for local arthropod composition and diversity

Geographic variation in the outcome of interspecific interactions may influence not only the evolutionary trajectories of species but also the structure of local communities. We investigated this community consequence of geographic variation for a facultative mutualism between ants and wild cotton (Gossypium thurberi). Ants consume wild cotton extrafloral nectar and can protect plants from herbivores. We chose three sites that differed in interaction outcome, including a mutualism (ants provided the greatest benefits to plant fitness and responded to manipulations of extrafloral nectar), a potential commensalism (ants increased plant fitness but were unresponsive to extrafloral nectar), and a neutral interaction (ants neither affected plant fitness nor responded to extrafloral nectar). At all sites, we manipulated ants and extrafloral nectar in a factorial design and monitored the abundance, diversity, and composition of other arthropods occurring on wild cotton plants. We predicted that the effects of ants and extrafloral nectar on arthropods would be largest in the location with the mutualism and weakest where the interaction was neutral. A non-metric multidimensional scaling analysis revealed that the presence of ants altered arthropod composition, but only at the two sites in which ants increased plant fitness. At the site with the mutualism, ants also suppressed detritivore/scavenger abundance and increased aphids. The presence of extrafloral nectar increased arthropod abundance where mutual benefits were the strongest, whereas both arthropod abundance and morphospecies richness declined with extrafloral nectar availability at the site with the weakest ant–plant interaction. Some responses were geographically invariable: total arthropod richness and evenness declined by approximately 20% on plants with ants, and extrafloral nectar reduced carnivore abundance when ants were excluded from plants. These results demonstrate that a facultative ant–plant mutualism can alter the composition of arthropod assemblages on plants and that these community-level consequences vary across the landscape.     

Utilization on extrafloral nectaries and fruit domatia of Canavalia lineata and C. cathartica (Leguminosae) by ants

Utilization of the extrafloral nectaries (EFNs) and fruits of Canavalia lineata and C. cathartica by ants was investigated at 30 sites in Japan. The fruits of C. lineata and C. cathartica were inhabited by five and eight ant species, respectively. Ant nesting periods and their utilization of EFNs differed between C. lineata and C. cathartica. Canavalia lineata flowers once a year, and periods of EFN-utilization and fruit-nesting by ants do not overlap. The fruit-nesting ants on C. lineata seem to invade the plant from the holes made by moth larvae or breaches made by decay. The ants nesting on the fruits of C. lineata may defend the plant against seed herbivores because they feed on moth larvae. Canavalia cathartica flowers several times over a year, and fruits are found throughout the year; therefore, periods of EFN-utilization and fruit-nesting by ants are overlapped. Canavalia cathartica offers year-round nesting sites and food for ants, and therefore may receive a higher defensive effect from ants than C. lineata. Handling editor: Graham Stone.     

Geographic patterns of genetic diversity from the native range of <Emphasis Type="Italic">Cactoblastis cactorum</Emphasis> (Berg) support the documented history of invasion and multiple introductions for invasive populations

Spread of the invasive cactus-feeding moth Cactoblastis cactorum has been well documented since its export from Argentina to Australia as a biocontrol agent, and records suggest that all non-native populations are derived from a single collection in the moth’s native range. The subsequent global spread of the moth has been complex, and previous research has suggested multiple introductions into North America. There exists the possibility of additional emigrations from the native range in nursery stock during the late twentieth century. Here, we present mitochondrial gene sequence data (COI) from South America (native range) and North America (invasive range) to test the hypothesis that the rapid invasive spread in North America is enhanced by unique genetic combinations from isolated portions of the native range. We found that haplotype richness in the native range of C. cactorum is high and that there was 90% lower richness in Florida than in Argentina. All Florida C. cactorum haplotypes are represented in a single, well-defined clade, which includes collections from the reported region of original export from Argentina. Thus, our data are consistent with the documented history suggesting a single exportation of C. cactorum from the eastern region of the native range. Additionally, the presence of geographic structure in three distinct haplotypes within the same clade across Florida supports the hypothesis of multiple introductions into Florida from a location outside the native range. Because the common haplotypes in Florida are also known to occur in the neighboring Caribbean Islands, the islands are a likely source for independent North American colonization events. Our data show that rapid and successful invasion within North America cannot be attributed to unique genetic combinations. This suggests that successful invasion of the southeastern US is more likely the product of a fortuitous introduction into favorable abiotic conditions and/or defense responses of specific Opuntia hosts, rapid adaptation, or a release from native enemies.     

Do invasive ants respond more strongly to carbohydrate availability than co‐occurring non‐invasive ants? A test along an active Anoplolepis gracilipes invasion front

Invasions by non‐native insects can have important ecological impacts, particularly on island ecosystems. However, the factors that promote the success of invaders relative to co‐occurring non‐invasive species remain unresolved. For invasive ants, access to carbohydrate resources via interactions with both extrafloral nectary‐bearing plants and honeydew‐excreting insects may accelerate the invasion process. A first step towards testing this hypothesis is to determine whether invasive ants respond to variation in the availability of carbohydrate resources, and whether this response differs from that of co‐occurring, non‐invasive ants. We investigated the effect of carbohydrate subsidies on the short‐term foraging and hemipteran‐tending behaviours of the invasive ant Anoplolepis gracilipes (Formicidae) and co‐occurring ant species on an extrafloral nectary‐bearing plant by experimentally manipulating carbohydrate levels and tracking ant recruitment. We conducted experiments in 2 years at two sites: one site was invaded by A. gracilipes prior to 2007 and the other became invaded during the course of our study, allowing pre‐ (2007) and post‐invasion (2009) comparisons. Short‐term increases in carbohydrate availability increased the density of A. gracilipes workers on plants by as much as 400% and reduced tending of honeydew‐excreting insects by this species by up to 89%, with similar responses across years. In contrast, ants at the uninvaded site in 2007 showed a weak and non‐significant forager recruitment response. Across all sites, A. gracilipes workers were the only ants that responded to carbohydrate manipulations in 2009. Furthermore, ant–carbohydrate dynamics at a site newly invaded by A. gracilipes quickly diverged from dynamics at uninvaded sites and converged on those of the site with an established invasion. These findings suggest that carbohydrate resources may be particularly important for A. gracilipes invasions, and underscore the importance of species interactions, particularly putative mutualisms, in facilitating exotic species invasions.     

Comparing the effects of the exotic cactus-feeding moth, Cactoblastis cactorum (Berg) (Lepidoptera: Pyralidae) and the native cactus-feeding moth, Melitara prodenialis (Walker) (Lepidoptera: Pyralidae) on two species of Florida Opuntia

This study examined the effects of the native cactus moth borer, Melitara prodenialis, and the invasive cactus moth borer, Cactoblastis cactorum, on two common cactus species, Opuntia stricta and O. humifusa at coastal and inland locations in central Florida. Opuntia stricta were present only at coastal sites and O. humifusa were present at coastal and inland sites. Throughout the duration of the study, coastal plants were subject to damage solely by C. cactorum and inland plants solely by M. prodenialis. Results showed marginally significantly higher numbers of eggsticks on O. stricta than O. humifusa and significantly higher numbers at coastal sites than at inland sites. There was also significantly higher moth damage on O. stricta than O. humifusa and at coastal sites than inland sites, but not significantly so. However, there was a higher level of plant mortality for O. humifusa than for O. stricta and a significantly higher level of cactus mortality at inland sites when compared to coastal sites. This increased mortality may be due to increased attack by true bugs, Chelinidea vittiger, and by Dactylopius sp., combined with attack by M. prodenialis. Inland plants also tended to be smaller than coastal plants and could be more susceptible to the combined effects of all insects. Further long-term research on coastal cactus survival when attacked and unattacked by Cactoblastis is necessary to fully determine the effects of this moth on Opuntia survival.     

Floral visitation by the Argentine ant reduces pollinator visitation and seed set in the coast barrel cactus, Ferocactus viridescens

Mounting evidence indicates that trade-offs between plant defense and reproduction arise not only from resource allocation but also from interactions among mutualists. Indirect costs of plant defense by ants, for example, can outweigh benefits if ants deter pollinators. Plants can dissuade ants from occupying flowers, but such arrangements may break down when novel ant partners infiltrate mutualisms. Here, we examine how floral visitation by ants affects pollination services when the invasive Argentine ant (Linepithema humile) replaces a native ant species in a food-for-protection mutualism with the coast barrel cactus (Ferocactus viridescens), which, like certain other barrel cacti, produces extrafloral nectar. We compared the effects of floral visitation by the Argentine ant with those of the most prevalent native ant species (Crematogaster californica). Compared to C. californica, the Argentine ant was present in higher numbers in flowers. Cactus bees (Diadasia spp.), the key pollinators in this system, spent less time in flowers when cacti were occupied by the Argentine ant compared to when cacti were occupied by C. californica. Presumably as a consequence of decreased duration of floral visits by Diadasia, cacti occupied by L. humile set fewer seeds per fruit and produced fewer seeds overall compared to cacti occupied by C. californica. These data illustrate the importance of mutualist identity in cases where plants balance multiple mutualisms. Moreover, as habitats become increasingly infiltrated by introduced species, the loss of native mutualists and their replacement by non-native species may alter the shape of trade-offs between plant defense and reproduction.     

Oviposition site selection in Cactoblastis cactorum (Lepidoptera): constraints and compromises

Summary Oviposition by Cactoblastis cactorum on Opuntia ficus-indica and O. aurantiaca was assessed from the positioning of egg sticks on plants in the field. The number of egg sticks laid on O. ficus-indica plants was affected by: (1) plant size; (2) moth emergence near the plant; (3) cladode condition; and (4) plant conspicuousness. These factors contributed towards the clumping of egg sticks on plants. There was no apparent oviposition preference for one of the two host plant species despite the fact that egg predation was higher and fecundity lower on O. aurantiaca. The selection of a site for oviposition on the host plants was influenced by: (1) cladode condition; (2) height above ground; and (3) shelter from wind during oviposition. Succulent cladodes were the favoured sites for oviposition. The evidence suggests that in C. cactorum, oviposition site selection is largely the net result of a compromise between oviposition behaviour selected for increasing the probability of juvenile survival and oviposition behaviour selected for increasing the probability of laying the full complement of eggs. In addition, environmental and physiological factors such as wind and wing-loading, are thought to place constraints on the range of sites available for oviposition.     

Foraging behavior of Brachygastra lecheguana (Hymenoptera: Vespidae) on Banisteriopsis malifolia (Malpighiaceae): Extrafloral nectar consumption and herbivore predation in a tending ant system

In the Brazilian savanna many plant species bear regular associations with patrolling ants that are aggressive towards insect herbivores. However, not only ants but also several species of predatory wasps are attracted to plants due to the extrafloral nectaries (EFNs). Such wasps feed on both herbivores and plant exudates. In this study we describe the foraging behavior of the social Polistinae wasp Brachygastra lecheguana in the extrafloral nectaried shrub Banisteriopsis malifolia, and investigated the influence of patrolling ants Camponotus blandus on the activity of the wasp. Brachygastra lecheguana fed on the endophytic larvae of Anthonomus (Curculionidae) beetles that developed inside flower buds. The wasp lacerated the bud layers to reach the beetle larvae located at the bud core. The wasp visits to Ba. malifolia were statistically related to the abundance of flower buds and beetles. Ant exclusion experiments revealed that the hunting behavior of B. lecheguana on beetles was not related to the absence of C. blandus. However we found that wasps spent more time consuming extrafloral nectar on branches where ants were excluded. This is the first study reporting extrafloral nectar consumption by B. lecheguana, as well as the predation on herbivores in natural areas. In cerrado vegetation, ants benefit the plant by reducing insect herbivores, and our study provides evidence that the B. lecheguana – Ba. malifolia system represents a potential interaction where the wasp may also benefit the host plant. The value of this wasp species as a plant‐guard is discussed.     

Endangered Cactus Restoration: Mitigating the Non-Target Effects of a Biological Control Agent (Cactoblastis cactorum) in Florida

The moth Cactoblastis cactorum (Berg), the poster child of weed biological control in Australia, has recently invaded the United States and threatens native cacti. Concern is greatest for the endangered semaphore cactus, Opuntia corallicola, of which only two known populations exist in the wild. We made three separate outplantings of O. corallicola, designed to bolster the number of extant cacti and to test the effectiveness of three different treatments to protect the cacti from Cactoblastis. In one outplanting, we tested the associational susceptibility hypothesis and found that cacti planted more than 20 m away from the common prickly pear cactus, Opuntia stricta, which act as a reservoir of Cactoblastis, were just as frequently attacked and killed by Cactoblastis as cacti planted within 5 m. In addition, Cactoblastis attack was greater in the shade than in the sun. In the second outplanting, we minimized the attack from Cactoblastis by using protective cages planted at least 500 m from O. stricta in areas not inhabited by cacti. Cages attracted the attention of local animals, which destroyed the cages and trampled the cacti inside to death. Crown rot caused high mortality in this outplanting. In the third outplanting, again conducted at least 500 m away from O. stricta, fertilization did not reduce crown rot mortality. We suggest that increasing populations of O. corallicola in Florida, by means of outplantings, will remain a challenge because of death from Cactoblastis when planted in areas where cacti normally grow and because of death from crown rot in areas where they do not. Because Cactoblastis is moving rapidly northward and westward and has already reached Charleston, South Carolina, rare cacti in the rest of the U.S. Southeast may be in danger. Eventually, many cactus species in the U.S. South, Southwest, and Mexico will likely be threatened by this moth.     

Elevated dominance of extrafloral nectary-bearing plants is associated with increased abundances of an invasive ant and reduced native ant richness

Aim Invasive ants can have substantial and detrimental effects on co‐occurring community members, especially other ants. However, the ecological factors that promote both their population growth and their negative influences remain elusive. Opportunistic associations between invasive ants and extrafloral nectary (EFN)‐bearing plants are common and may fuel population expansion and subsequent impacts of invasive ants on native communities. We examined three predictions of this hypothesis, compared ant assemblages between invaded and uninvaded sites and assessed the extent of this species in Samoa. Location The Samoan Archipelago (six islands and 35 sites). Methods We surveyed abundances of the invasive ant Anoplolepis gracilipes, other ant species and EFN‐bearing plants. Results Anoplolepis gracilipes was significantly more widely distributed in 2006 than in 1962, suggesting that the invasion of A. gracilipes in Samoa has progressed. Furthermore, (non‐A. gracilipes) ant assemblages differed significantly between invaded and uninvaded sites. Anoplolepis gracilipes workers were found more frequently at nectaries than other plant parts, suggesting that nectar resources were important to this species. There was a strong, positive relationship between the dominance of EFN‐bearing plants in the community and A. gracilipes abundance on plants, a relationship that co‐occurring ants did not display. High abundances of A. gracilipes at sites dominated by EFN‐bearing plants were associated with low species richness of native plant‐visiting ant species. Anoplolepis gracilipes did not display any significant relationships with the diversity of other non‐native ants. Main conclusions Together, these data suggest that EFN‐bearing plants may promote negative impacts of A. gracilipes on co‐occurring ants across broad spatial scales. This study underscores the potential importance of positive interactions in the dynamics of species invasions. Furthermore, they suggest that conservation managers may benefit from explicit considerations of potential positive interactions in predicting the identities of problematic invaders or the outcomes of species invasions.     

Should aphids attract or repel ants? Effect of rival aphids and extrafloral nectaries on ant–aphid interactions

Among plants and herbivores, two types of conflicts occur in relation to mutualism with ants: one is competition for ant mutualism among myrmecophilous herbivores and plants, and the other is the conflict whether to attract or repel ants between myrmecophiles and nonmyrmecophiles that are damaged by ants. We investigated the extent to which two species of aphids (Megoura crassicauda and Aphis craccivora) and extrafloral nectaries on their host plant (Vicia faba var. minor) interact with one another for their relationships with ants. We designed an experiment where ants can choose to visit seedlings colonized by (1) M. crassicauda, (2) A. cracivora, (3) both aphid species, or (4) neither aphid species. Ants preferred A. craccivora to extrafloral nectaries and avoided tending M. crassicauda. We also analyzed the population growth of each aphid when it coexists with (1) ants, (2) the other aphid species, (3) ants and the other aphid species, or (4) neither of them. Under ant-free conditions, we detected an exploitative competition between the two aphid species. The ants had no significant effect on the population of A. craccivora, whereas they had negative effects on the population growth of M. crassicauda by attacking some individuals. When both aphids coexisted, M. crassicauda suffered ant attack more intensely because A. craccivora attracted more ants than extrafloral nectaries despite ant-repelling by M. crassicauda. On the other hand, the ants benefited A. craccivora by eliminating its competitor. To avoid ant attack, aphids may have been selected either to be more attractive to ants than other sympatric sugar sources or to repel the ants attracted to them. We hypothesize that competition among sympatric sugar sources including rival aphids and extrafloral nectaries is a factor restricting aphids to be myrmecophilous. Received: January 17, 2000 / Accepted: July 4, 2000     

Cactoblastis cactorum (Berg) (Lepidoptera: Pyralidae) use of Opuntia host species in Argentina

A central aspect in biology and ecology is to determine the combination of factors that influence the distribution of species. In the case of herbivorous insects, the distribution of herbivorous species is necessarily associated with their host plants, a pattern often referred to as “host use”. Novel interactions that arise during a biological invasion can have important effects on the dynamics of that invasion, especially if it is driven by only a subset of the genetic diversity of the invading species. This is the case of the wellknown South American cactus moth, Cactoblastis cactorum, a successfully used biological control agent of non-native Opuntia cacti in Australia and South Africa, but now threatening unique cactus diversity and agriculture in North America. We studied the patterns of host plant usage by and host plant availability for C. cactorum under field conditions in Argentina, covering the geographical range of the four C. cactorum phylogroups and the recently documented southern distribution. We also assessed female preference and larval performance under laboratory conditions. Cactoblastis cactorum showed a geographical pattern of host use in its native range that was related to host availability. Laboratory assays of female preference showed some degree of preference to oviposit on O. ficus-indica, O. leucotricha and O. quimilo, but it was not positively correlated with the performance of larvae. These findings contribute to the further comprehension of the host use dynamics of C. cactorum in the insects’ native range, and could provide useful information for assessing the risk and future spread of this insect in North America.     

EXTRAFLORAL NECTAR OF FEROCACTUS ACANTHODES (CACTACEAE): COMPOSITION AND ITS IMPORTANCE TO ANTS

Ants are attracted to extrafloral nectaries subtending reproductive organs of Ferocactus acanthodes var. lecontei (Cactaceae) in central Arizona. Extrafloral nectar produced by these glands contained amino acids, sugars, and water. Nectar quality and composition varied temporally in relation to plant reproductive phenology. The number of nectar glands on a barrel cactus did not change significantly, however; the mass of nectar produced per gland increased significantly with immature fruit production. Of the three sugars present in extrafloral nectar (fructose, glucose, and sucrose), only glucose occurred at a higher concentration in June, when immature fruits first appeared on barrel cactus. Amino acid concentration and composition in extrafloral nectar of barrel cactus did not change significantly over time. Ant density on barrel cactus increased significantly from mid-May to mid-June at two field sites. Water availability per nectar gland increased 158% from May to June. Water plays an important role in attracting ants to barrel cacti.     

Ant–plant interaction in the Neotropical savanna: direct beneficial effects of extrafloral nectar on ant colony fitness

Current evidence suggests that ant–plant relationships may influence species composition, abundance, and interactions at the community scale. The main resource that plants offer to ants is extrafloral nectar (EFN) and the major part of published studies shown benefits from ants to plants possessing EFNs. However, the complementary question of whether and how ants benefit from EFNs is rarely addressed. Here, we present the results of a long-term study to demonstrate whether EFN has a positive effect on ant colony fitness. We quantified colony growth rate, survival and the final weight of individuals as measures of benefit derived from EFN. Our results provide clear evidence that EFN can have a significant positive impact on the survivorship, growth and reproduction of the Myrmicinae Cephalotes pusillus. In fact, a diet rich in EFN (providing at least 30 cal per day) resulted in five times more individuals per colony, greater body weights, and more eggs. These results have shed new light on the relationships between ants and EFN-bearing plants such as in tropical and temperate systems. The ant C. pusillus is the first case in which we have firm evidence that EFN improves colony growth and development, corroborating more than 100 years of experimental evidence of benefits to plants in these widespread relationships.     

Fire triggers the activity of extrafloral nectaries,but ants fail to protect the plant against herbivores in a neotropical savanna

Herbivores are attracted to young shoots and leaves because of their tender tissues. However, in extrafloral nectaried plants, young leaves also attract patrolling ants, which may chase or prey on herbivores. We examined this scenario in extrafloral nectaried shrubs of Banisteriopsis malifolia resprouting after fire, which promoted both the aseasonal production of leaves and the activity of extrafloral nectaries (EFNs). Results were compared between resprouting (burned) and unburned control plants. The aggressive ant species Camponotus crassus and the herbivorous thrips Pseudophilothrips obscuricornis were respectively rapidly attracted to resprouting plants because of the active EFNs and their less sclerophyllous leaves. The abundance of these insects was almost negligible in the control (unburned) shrubs. Ants failed to protect B. malifolia, as no thrips were preyed upon or injured by ants in resprouting plants. Consequently, on average, 37 % of leaves from resprouting shrubs had necrosis marks. Upon contact with ants, thrips released small liquid droplets from their abdomen, which rapidly displaced ants from the surroundings. This study shows that P. obscuricornis disrupted the facultative mutualism between C. crassus and B. malifolia, since ants received extrafloral nectar from plants, but were unable to deter herbivore thrips.