Diversity of ant-plant interactions: protective efficacy in Macaranga species with different degrees of ant association

The pioneer tree Macaranga in SE Asia has developed manyfold associations with ants. The genus comprises all stages of interaction with ants, from facultative relationships to obligate myrmecophytes. Only myrmecophytic Macaranga offer nesting space for ants and are associated with a specific ant partner. The nonmyrmecophytic species are visited by a variety of different ant species which are attracted by extrafloral nectaries (EFN) and food bodies. Transitional Macaranga species like M. hosei are colonized later in their development due to their stem structure. Before the colonization by their specific Crematogaster partner the young plants are visited by different ant species attracted by EFN. These nectaries are reduced and food body production starts as soon as colonization becomes possible. We demonstrated earlier that obligate ant partners can protect their Macaranga plants against herbivore damage and vine cover. In this study we focused on nonspecific interactions and studied M. tanarius and M. hosei, representing a non-myrmecophyte and a transitional species respectively. In ant exclusion experiments both M. tanarius and M. hosei suffered significantly higher mean leaf damage than controls, 37% versus 6% in M. hosei, 16% versus 7% in M. tanarius. M. tanarius offers both EFN and food bodies so that tests for different effects of these two food rewards could be conducted. Plants with food bodies removed but with EFN remaining had the lowest mean increase of herbivore damage of all experimental groups. Main herbivores on M. hosei were mites and caterpillars. Many M. tanarius plants were infested by a shootborer. Both Macaranga species were visited by various ant species, Crematogaster spp. being the most abundant. We found no evidence for any specific relationships. The results of this study strongly support the hypothesis that non-specific, facultative associations with ants can be advantageous for Macaranga plants. Food bodies appear to have lower attractive value for opportunistic ants than EFN and may require a specific dietary adaptation. This is also indicated by the fact that food body production in the transitional M. hosei does not start before stem structure allows a colonization by the obligate Crematogaster species. M. hosei thus benefits from facultative association with a variety of ants until it produces its first domatia and can be colonized by its obligate mutualist.     

Mutualism between Maieta guianensis Aubl., a myrmecophytic melastome,and one of its ant inhabitants: ant protection against insect herbivores

Summary The hypothesis that ants (Pheidole minutula) associated with the myrmecophytic melastome Maieta guianensis defend their host-plant against herbivores was investigated in a site near Manaus, Amazonas, Brazil. M. guianensis is a small shrub that produces leaf pouches as ant domatia. Plants whose ants were experimentally removed suffered a significant increase in leaf damage compared with control plants (ants maintained). Ants patrol the young and mature leaves of Maieta with the same intensity, presumably since leaves of both ages are equally susceptible to herbivore attack. The elimination of the associated ant colony, and consequent increase in herbivory, resulted in reduced plant fitness. Fruit production was 45 times greater in plants with ants than in plants without ants 1 year after ant removal.     

Ant species confer different partner benefits on two neotropical myrmecophytes

The dynamics of mutualistic interactions involving more than a single pair of species depend on the relative costs and benefits of interaction among alternative partners. The neotropical myrmecophytes Cordia nodosa and Duroia hirsuta associate with several species of obligately symbiotic ants. I compared the ant partners of Cordia and Duroia with respect to two benefits known to be important in ant-myrmecophyte interactions: protection against herbivores provided by ants, and protection against encroaching vegetation provided by ants. Azteca spp., Myrmelachista schumanni, and Allomerus octoarticulatus demerarae ants all provide the leaves of Cordia and Duroia some protection against herbivores. However, Azteca and Allomerus provide more protection than does Myrmelachista to the leaves of their host plants. Although Allomerus protects the leaves of its hosts, plants occupied by Allomerus suffer more attacks by herbivores to their stems than do plants occupied by other ants. Relative to Azteca or Allomerus, Myrmelachista ants provide better protection against encroaching vegetation, increasing canopy openness over their host plants. These differences in benefits among the ant partners of Cordia and Duroia are reflected in the effect of each ant species on host plant size, growth rate, and reproduction. The results of this study show how mutualistic ant partners can differ with respect to both the magnitude and type of benefits they provide to the same species of myrmecophytic host.     

The capacity of a Myrmica ant nest to support a predacious species of Maculinea butterfly

Summary Caterpillars of Maculinea arion are obligate predators of the brood of Myrmica sabuleti ants. In the aboratory, caterpillars eat the largest available ant larvae, although eggs, small larvae and prepupae are also palatable. This is an efficient way to predate. It ensures that newly-adopted caterpillars consume the final part of the first cohort of ant brood in a nest, before this pupates in early autumn and becomes unavailable as prey. At the same time, the fixed number of larvae in the second cohort is left to grow larger before being killed in late autumn and spring. Caterpillars also improve their feeding efficiency by hibernating for longer than ants in spring, losing just 6% of their weight while the biomass of ant larvae increases by 27%. Final instar caterpillars acquire more than 99% of their ultimate biomass in Myrmica nests, growing from 1.3 mg to an estimated 173 mg. A close correlation was found between the weights of caterpillars throughout autumn and the number of large ant larvae they had eaten. This was used to calculate the number of larvae eaten in spring, allowing both for the loss of caterpillar weight during winter and the increase in the size of their prey in spring. It is estimated that 230 of the largest available larvae, and a minimum nest size of 354 M. sabuleti workers, is needed to support one butterfly. Few wild M. sabuleti nests are this large: on one site, it was estimated that 85% of nests were too small to produce a butterfly, and only 5% could support two or more. This prediction was confirmed by the mortalities of 376 caterpillars in 151 wild M. sabuleti nests there. Mortalities were particularly high in nests that adopted more than two caterpillars, apparently due to scramble competition and starvation in autumn. Survival was higher than predicted in wild nests that adopted one caterpillar. These caterpillars seldom exhaust their food before spring, when there is intense competition among Myrmica for nest sites. Ants often desert their nests in the absence of brood, leaving the caterpillar behind. Vacant nests are frequently repopulated by a neighbouring colony, carrying in a fresh supply of brood. Maculinea arion caterpillars have an exceptional ability to withstand starvation, and sometimes survive to parasitize more than one Myrmica colony. Despite these adaptations, predation is an inefficient way to exploit the resources of a Myrmica nest. By contrast, Maculinea rebeli feeds mainly at a lower trophic level, on the regurgitations of worker ants. Published data show that Myrmica nests can support 6 times more caterpillars of Maculinea rebeli than of M. arion in the laboratory. This is confirmed by field data.     

Food source quality and ant dominance hierarchy influence the outcomes of ant-plant interactions in an arid environment

In this study, we conducted a series of experiments in a population of Vachellia constricta (Fabaceae) in the arid Tehuacan-Cuicatláan valley, Mexico, in order to evaluate if the food source quality and ant dominance hierarchy influence the outcomes of ant-plant interactions. Using an experiment with artificial nectaries, we observed that ants foraging on food sources with higher concentration of sugar are quicker in finding and attacking potential herbivorous insects. More specifically, we found that the same ant species may increase their defence effectiveness according to the quality of food available. These findings indicate that ant effectiveness in plant protection is context-dependent and may vary according to specific individual characteristics of plants. In addition, we showed that competitively superior ant species tend to dominate plants in periods with high nectar activity, emphasizing the role of the dominance hierarchy structuring ant-plant interactions. However, when high sugar food sources were experimentally available ad libitum, the nocturnal and competitively superior ant species, Camponotus atriceps, did not dominate the artificial nectaries during the day possibly due to limitation of its thermal tolerance. Therefore, temporal niche partitioning may be allowing the coexistence of two dominant ant species (Camponotus rubritorax during the day and C. atriceps at night) on V. constricta. Our findings indicate that the quality of the food source, and temporal shifts in ant dominance are key factors which structure the biotic plant defences in an arid environment.     

Unadapted behaviour of native,dominant ant species during the colonization of an aggressive,invasive ant

Among the factors driving the invasive success of non-indigenous species, the “escape opportunity” or “enemy release” hypothesis argues that an invader’s success may result partly from less resistance from the new competitors found in its introduced range. In this study, we examined competitive interactions between the little fire ant Wasmannia auropunctata (Roger) and ant species of the genus Pheidole in places where both are native (French Guiana) and in places where only species of Pheidole are native (New Caledonia). The experimental introduction of W. auropunctata at food resources monopolized by the Pheidole species induced the recruitment of major workers only for the Guianian Pheidole species, which were very effective at killing Wasmannia competitors. In contrast, an overall decrease in the number of Pheidole workers and no recruitment of major workers were observed for the New Caledonian species, although the latter were the only ones able to kill the Wasmannia workers. These results emphasize the inappropriate response of native dominant New Caledonian species to W. auropunctata and, thus, the importance of enemy recognition and specification in the organization of ant communities. This factor could explain how invasive animal species, particularly ants, may be able to successfully invade species-rich communities.     

Predation by vertebrates and invertebrates on the seeds of five canopy tree species of an Amazonian forest

We studied the pre-germination loss of seeds to invertebrate and vertebrate seed predators of 5 species of Amazonian trees (Astrocaryum macrocalyx—Palmae; Bertholletia excelsa—Lecithydaceae; Calatola venezuelana—Icacinaceae; Dipteryx micrantha—Leguminosae (Papilionoidae); Hymenaea courbaril-Leguminosae (Caesalpinoidae)). These five species were selected from a large tree flora on several criteria. All possess large (3–10 cm) well-protected seeds that might plausibly be attractive to mammalian seed predators. The reproductive biology of three of the species, or close congeners, had been studied elsewhere in the Neotropics (Astrocaryum, Dipteryx, Hymenaea); one is important to the economy of southeastern Peru (Bertholletia); and one, despite large and apparently edible seeds, appeared to suffer no pre-germination loss to predators (Calatola). We conducted the research in mature forests in the Manu National Park of southeastern Peru where mammal densities are unperturbed by human activities. Densities of adult trees of the five species in our area range from very high (>30 per ha: Astrocaryum) to very low (1 per ha: Hymenaea).Loss of seeds to all causes, and to mammalian seed predators in particular, was determined for seeds placed in 2-square meter mammal exclosures and in open controls located at 10 m (near) and 50 m (far) from a large mature individual of the target species (with minor variations in the design for Astrocaryum and Calatola). The exclosures were of two types: impermeable—designed to exclude all mammals, but not invertebrate seed predators, and semipermeable—designed to admit small (<500 g), but not large mammals. Experimental and control plots were stocked with apparently viable seeds during the dry-wet transition period (October–November) and scored one year later.A significant distance effect (higher predation near vs far from a large conspecific adult) was found in only one of the species (Astrocaryum), the only one to be attacked with high frequency by invertebrate seed predators. The absence of any detectable distance effect attributable to mammals suggests that mammals, over the course of a year, thoroughly search the forest floor for seeds. Invertebrates may thus be responsible for most pre-germination distance (density) effects. With respect to the treatments, we found three qualitatively distinct results: seeds of three species (Astrocaryum, Bertholletia, Dipteryx) were significantly protected by the impermeable, but not semipermeable exclosures, implicating small mammals in seed loss; the seeds of one species (Hymenaea) were significantly protected by exclosures of both types, implicating large mammals; and the seeds of one species (Calatola) exhibited 100% survival, whether or not protected by exclosures.The importance of large mammals as seed predators is generally underestimated in these experiments because semipermeable exclosures may serve as foraging reserves for small mammals. Finally, we noted no relationship between the intensity of mammalian seed predation (as suggested by the survival of unprotected seeds) and the abundance of adults of the five species in the environment. The diversity of results obtained for the five species reveals that large-seeded tropical trees may display a wide range of demographic patterns, and points to the likely importance of post-germination bottlenecks in the population biology of many species, even those that may experience severe pre-germination seed loss.     

Variation in the use of orchid extrafloral nectar by ants

Summary The relation between ant-plant specificity and the use of host plants as a resource was investigated in the facultative, myrmecophytic orchid, Caularthron bilamellatum (Rchg.f.) Schult. Using stable isotopes, we determined the portion of the ants' diets derived from host plants. We documented that six ant species inhabiting the orchid: (1) derived nutritional benefit from host orchids, and (2) had species-specific levels of extrafloral nectar use. Proportionate contribution of extrafloral nectar to ant diets ranged from 11 to 48%. These results demonstrate extreme interspecific differences in the nutritional benefits received by ants from host orchids. Interspecific differences in nutritional benefits from orchid nectar may be affected by colony size, nutritional needs, behavioral ecology of the ants, and the abundance of alternate food sources.     

Disentangling the assembly mechanisms of ant cuticular bacterial communities of two Amazonian ant species sharing a common arboreal nest

Bacteria living on the cuticle of ants are generally studied for their protective role against pathogens, especially in the clade of fungus‐growing ants. However, little is known regarding the diversity of cuticular bacteria in other ant host species, as well as the mechanisms leading to the composition of these communities. Here, we used 16S rRNA gene amplicon sequencing to study the influence of host species, species interactions and the pool of bacteria from the environment on the assembly of cuticular bacterial communities on two phylogenetically distant Amazonian ant species that frequently nest together inside the roots system of epiphytic plants, Camponotus femoratus and Crematogaster levior. Our results show that (a) the vast majority of the bacterial community on the cuticle is shared with the nest, suggesting that most bacteria on the cuticle are acquired through environmental acquisition, (b) 5.2% and 2.0% of operational taxonomic units (OTUs) are respectively specific to Ca. femoratus and Cr. levior, probably representing their respective core cuticular bacterial community, and (c) 3.6% of OTUs are shared between the two ant species. Additionally, mass spectrometry metabolomics analysis of metabolites on the cuticle of ants, which excludes the detection of cuticular hydrocarbons produced by the host, were conducted to evaluate correlations among bacterial OTUs and m/z ion mass. Although some positive and negative correlations are found, the cuticular chemical composition was weakly species‐specific, suggesting that cuticular bacterial communities are prominently environmentally acquired. Overall, our results suggest the environment is the dominant source of bacteria found on the cuticle of ants.     

Bigger ant colonies reduce herbivory and herbivore residence time on leaves of an ant-plant: Azteca muelleri vs. Coelomera ruficornis on Cecropia pachystachya

Summary The effect of defence force size in colonies of the ant Azteca muelleri on the time spent to localize, attack and expel the specialized herbivorous beetle Coelomera ruficornis from Cecropia pachystachya bushes was studied in an area of Atlantic forest in northeastern Brazil. Our results show that Azteca muelleri expel Coelomera ruficornis from Cecropia pachystachya and that the number of ants leaving a colony (defence force size) is negatively correlated with the residence time of an adult beetle on the plant. Colonies with larger defence forces recruited larger numbers of ants, resulting in faster herbivore discovery (r ²=0.80; n=17; P<0.001) and reduced herbivore residence time on a leaf (r ²=0.79 n=23; P<0.001) before being driven off by the ants. We also found a negative and significant relationship between herbivore damage on leaves and ant colony size (r ²=0.28; n=17; P<0.05). We conclude that larger colonies have more individuals available to patrol a plant and recruit defenders toward herbivores. This reduces the time spent to locate and expel susceptible herbivores from the plant. Since the plant probably benefits from reduced herbivory and the plant provides food for the ants, the association between Azteca muelleri and Cecropia pachystachya appears mutualistic.     

Nest site selection by a neotropical swarm-founding wasp: seasonal alternation of nest orientation

Nest orientation in social insects has been intensively studied in warmer and cooler climates, particularly in the northern hemisphere. Previous studies have consistently shown that species subjected to these climatic conditions prefer to select mostly southern locations where the nests can gain direct sunlight. However, very little is known on nest orientation in tropical and subtropical social insects. We studied nest orientations initiated by swarms throughout a year in a Brazilian swarm-founding wasp, Polybia paulista von Ihering (Hymenoptera: Polistinae). Swarms selected various orientations as nest sites, but there was a particular trend in that swarms in the winter period (May–August) preferred to build northward-facing nests. This preference is opposite from that of social wasps observed in the northern hemisphere. Colonies of this species can potentially last for many years with continuous nesting, but nesting activities of colonies during the winter are severely limited due to cool temperature and a shortened day length. Northward-facing nests are warmer through the gain of direct solar heat during the winter period; consequently, choosing northward-facing sites may be advantageous for swarms in terms of a shortened brood development and shortened time needed to increase metabolic rates during warm-up for flight.     

金城山柔毛淫羊藿的开花物候与生殖特性

采用实地观测的方法,对南充市金城山三个不同生境中柔毛淫羊藿的开花物候特征及其生殖特性进行了研究。结果表明:柔毛淫羊藿花期为3月上旬至4月上旬,其种群、个体、花序和单花的花期分别历时30~41、22~34、9~18和4~8 d。三个生境中柔毛淫羊藿种群的开花物候进程基本相同,均呈单峰集中开花式样,因而能够吸引更多的传粉者访问而达到生殖成功。开花物候指数与坐果数之间的相关分析结果表明,坐果数与始花日期存在显著的负相关关系,与花期持续时间和开花数均呈极显著的正相关关系。揭示了药用植物柔毛淫羊藿的开花物候特征与生殖特性。     

Experimental evaluation of the impacts of two ant species on banana weevil in Uganda

The banana weevil, Cosmopolites sordidus (Germar), is an important pest of bananas. Predatory ants are increasingly being viewed as possible biological control agents of this pest because they are capable of entering banana plants and soil in search of prey. We studied ant predation on banana weevil in Uganda in crop residues and live plants in both laboratory and field experiments. Field studies with live plants used chemical ant exclusion in some plots and ant enhancement via colony transfer in others to measure effects of Pheidole sp. 2 and Odontomachus troglodytes Santschi on plant damage and densities of immature banana weevils.In crop residues, an important pest breeding site, twice as many larvae were removed from ant-enhanced plots as in control plots. In young (2 month) potted suckers held in shade houses, ant ability to reduce densities of banana weevil life stages varied with the weevil inoculation rate. At the lowest density (2 female weevils per pot), densities of eggs, larvae, and pupae were reduced by ants. At higher rates there was no effect. In older suckers (5–11 months) grown in larger containers, banana weevil densities were not affected by ants, but damage levels were reduced. In a field trial lasting a full crop cycle (30 months), we found that the ants tested reduced the density of banana weevil eggs in suckers during the crop, but did not affect larval densities in the sampled suckers. However, most larvae occur in the main banana plants, rather than associated suckers. Nevertheless, levels of damage in mature plants at harvest did not differ between Amdro-treated and ant-enhanced plots, suggesting the ant species studied were not able to provide economic control of banana weevil under our test conditions.     

Spatial patterns of granivorous ant nest abundance and nest site selection in agricultural landscapes of Central Spain

Summary Patterns of abundance and site selection of granivorous ant nests were investigated in extensive cereal croplands of Central Spain. Nest densities and distributions were measured in two consecutive summers (1988 and 1989), together with habitat physiognomy and seed availability. Nest site selection patterns were analysed at two spatial scales (landscape and microhabitat) with respect to habitat physiognomy. Results indicate a very constant and predictable pattern of both nest abundance and nest site selection. Granivorous ant nests were most abundant in shrublands, and shrubby microsites were selected for nest placement. Croplands, and microsites with high covers of bare ground and litter, were avoided. These patterns were consistent between years despite a 1.7-fold increase in shrubland nest densities, that was attributed to the exceptionally dry winter between nest censuses. I suggest that winter survivorship of ant nests in the unploughed landscape units, and periodic ploughing in croplands, may be the main factors constraining granivorous ant densities in the landscape studied.     

Rainfall facilitates the spread, and time alters the impact, of the invasive Argentine ant

Climate change may exacerbate invasions by making conditions more favorable to introduced species relative to native species. Here we used data obtained during a long-term biannual survey of the distribution of ant species in a 481-ha preserve in northern California to assess the influence of interannual variation in rainfall on the spread of invasive Argentine ants, Linepithema humile, and the displacement of native ant species. Since the survey began in 1993, Argentine ants have expanded their range into 74 new hectares. Many invaded hectares were later abandoned, so the range of Argentine ants increased in some years and decreased in others. Rainfall predicted both range expansion and interannual changes in the distribution of Argentine ants: high rainfall, particularly in summer months, promoted their spread in the summer. This suggests that an increase in rainfall will promote a wider distribution of Argentine ants and increase their spread into new areas in California. Surprisingly, the distribution of two native ant species also increased following high rainfall, but only in areas of the preserve that were invaded by L. humile. Rainfall did not have a negative impact on total native ant species richness in invaded areas. Instead, native ant species richness in invaded areas increased significantly over the 13 years of observation. This suggests that the impact of Argentine ants on naïve ant communities may be most severe early in the invasion process.     

Phylogenetic relationships among species groups of the ant genus Myrmecia

Phylogenetic relationships among the nine species groups of the predominately Australian ant genus Myrmecia were inferred using 38 Myrmecia species and an outgroup using DNA sequences from two nuclear genes (622nt from 28S rRNA and 1907nt from the long-wave opsin gene). Nothomyrmecia macrops was selected as the most appropriate outgroup based on recent reliable studies showing monophyly of Myrmecia with Nothomyrmecia. The four species groups with an occipital carina (those of gulosa, nigrocincta, urens, and picta) were found to form a paraphyletic and basal assemblage out of which the five species groups lacking an occipital carina (those of aberrans, mandibularis, tepperi, cephalotes, and pilosula) arise as a strongly supported monophyletic assemblage. Monophyly was supported for four groups (those of gulosa, nigrocincta, picta, and mandibularis) but the situation is unclear for four others (those of urens, aberrans, tepperi, and pilosula). The aberrans group appears to be basal within the group lacking an occipital carina; a previous suggestion that it is the sister group to the rest of the genus is thus not supported.     

The role of food quality and competition in shaping the seasonal cycle in the reproductive activity of the sycamore aphid

The hypothesis that seasonal changes in sycamore aphid,Drepanosiphum platanoidis (Schr.), recruitment are determined by changes in food quality and aphid population density was tested. There was no clear association between the reproductive activity of the sycamore aphid and the seasonal changes in specific amino acids or groups of amino acids in extracts of sycamore,Acer pseudoplatanus L., leaves. Seasonal changes in reproductive activity tracked the changes in total amino acids of the leaf tissue of the host, but with a short time delay. High numbers of adult aphids appeared to depress reproductive activity. A regression analysis of the results revealed that total amino acids the previous week and current numbers of adult aphids significantly affected sycamore aphid reproductive activity. The results of this analysis support the above hypothesis, that the marked seasonal changes in the total quantity of amino acids in sycamore leaves and intraspecific competition for this resource, through its effect on adult weight, shape the seasonal cycle in the reproductive activity of the sycamore aphid.     

Attachment sites of parasitic lampreys: comparisons among species

Synopsis Distribution of attachment sites varies with the species of lamprey being considered. Large anadromous species (Petromyzon marinus and Lampetra tridentata) tend to attach ventrally, especially near the pectoral fins, while smaller freshwater species in shallow habitats (Ichthyomyzon castaneus and I. unicuspis) and species that feed on muscle tissue (Lampetra ayresi) tend to attach dorsally. Catostomids tend to be attacked on the head and, by the Ichthyomyzon species, on the paired fins. Distribution of attachments by P. marinus in laboratory studies may be affected by tank size. Attachments to the head and pectoral regions are probably associated with greater host mortality rates and may be underrepresented in field samples. Attachments to the pectoral region appear to combine low costs, in terms of handling time prior to feeding, with greater rates of energy intake once feeding has been initiated. Dorsal attachments by species that inhabit relatively shallow rivers and streams may be a compromise to avoid abrasion against the bottom. A consequence of dorsal attachments may be a reduced impact on host populations through prolonged attachments to individual hosts and reduced attack rates.