Edge‐mediated reduction of phorid parasitism on leaf‐cutting ants in a Brazilian Atlantic forest

Previous studies have shown that leaf‐cutting ant populations benefit greatly from living in or near the edges of the Brazilian Atlantic forest. One of the mechanisms responsible for this rise in population density is an edge‐mediated increase of pioneer plants, resulting in increased food availability for the ants (i.e., less bottom‐up control). Here, we hypothesized that the release from natural enemies (i.e., less top‐down control) may also contribute to the phenomenon. We investigated whether parasitism of phorid flies on leaf‐cutting ants decreases in colonies located along the forest edge vs. the interior of a large tract of Atlantic forest in northeastern Brazil. For this, we assessed abundance and rates of oviposition attack by phorids in bimonthly intervals over a period of 1 year in 10 adult colonies of Atta cephalotes (L.) (Hymenoptera: Formicidae: Myrmicinae), five at the forest edge and five in the forest interior. The number of phorids attracted by ants at edge colonies was 40% lower than that at interior colonies. The temporal variation in phorid attraction was also significant, with approximately 35% fewer flies in the dry months as compared to the rainy months. As a result of lower phorid abundance, ant workers of edge colonies suffered three times fewer oviposition attacks than those of interior colonies. There was a tendency for fewer attacks during dry months, but the difference in the temporal variation was not significant. Our findings suggest that edge creation contributes to increased leaf‐cutting ant abundance, not only via the attenuation of bottom‐up forces, but also through an environmentally triggered depression of parasitoid abundance/efficiency, possibly because of adverse environmental conditions in edge habitats.     

Effect of phorid fly density on the foraging of Atta vollenweideri leafcutter ants in the field

Leafcutter ants in the genus Atta (Hymenoptera: Formicidae: Attini) are considered major pests of agriculture and forestry in the Neotropics. Phorid flies (Diptera: Phoridae) have been proposed as viable candidates for biological control of ants because of the importance of their trait‐mediated effects on their hosts. However, the impact of different densities of phorid flies has never been assessed in the field. Experiments were conducted by isolating 3‐m sections of Atta vollenweideri Forel foraging trails with tunnels, and sampling ants in trails with 0, 1, or 4 Eibesfeldtphora trilobata Disney female parasitoid flies. Samples were collected every 30 min from these trails. We also collected a sample before introducing the parasitoids and another one 30 min after removing them from the trail. We measured traffic of ants on the trails, weight and type of plant material transported, and the proportion and size of the workers collected. The presence of phorids on the trails reduced the ant traffic and amount of plant material transported into the nests and decreased the proportion of workers on the trails in the size range preferred as hosts by the flies. The effect on worker size, as well as the lag effect recorded after phorids were removed from the tunnels, was more pronounced with four phorids. The presence of phorids also affected the weight of monocotyledon and dicotyledon material transported. Even at the minimum density possible, phorids significantly influenced a key aspect of the colony life, the food intake through foraging. From an applied point of view, our results show that releases of these phorids into the field should not necessarily involve many individuals to reduce foraging by A. vollenweideri, making them potentially useful candidates for biological control of these ants.     

Parasite Lost: Chemical and Visual Cues Used by <Emphasis Type="BoldItalic">Pseudacteon</Emphasis> in Search of <Emphasis Type="BoldItalic">Azteca instabilis</Emphasis>

An undescribed species of phorid fly (genus: Pseudacteon) parasitizes the ant Azteca instabilis F Smith, by first locating these ants through the use of both chemical and visual cues. Experiments were performed in Chiapas, Mexico to examine a) the anatomical source of phorid attractants, b) the specific chemicals produced that attract phorids, and c) the nature of the visual cues used by phorids to locate the ants. We determined that phorid-attracting chemicals were present within the dorsal section of the abdomen, the location of the pygidial gland. Further experiments indicate that a pygidial gland compound, 1-acetyl-2-methylcyclopentane, is at least partially responsible for attracting phorid flies to their host. Finally, although visual cues such as movement were important for host location, size and color of objects did not influence the frequency with which phorids attacked moving targets.     

Phorid species from Acromyrmex’s hosts and effect on their survival of two fungi proposed for the control of leafcutter ants

Phorid flies (Diptera: Phoridae, Metopinini) are natural enemies of leafcutter ants (Hymenoptera: Formicidae, Attini), which are among the most important pests in the Neotropical region. These parasitoids lay their eggs inside worker ants, causing the death of the parasitized ants, and the oviposition attacks on foraging workers interfere with the collection of vegetal material used in fungal gardens, thereby affecting the whole colony. However, because of the large number of ants per colony, more than one type of biological control agent is needed to have a significant impact. We collected parasitized leafcutter ants Acromyrmex lundii (Guérin‐Méneville), Acromyrmex heyeri (Forel), and Acromyrmex ambiguus (Emery) from various places in Argentina and reared their parasitoids. We recorded developmental times, host and phorid sizes, and other aspects of the biology of the phorids Apocephalus neivai Borgmeier, Apocephalus noetingerorum Brown & Disney, Myrmosicarius gracilipes Borgmeier, Myrmosicarius catharinensis Borgmeier, Myrmosicarius crudelis Borgmeier, and Myrmosicarius cristobalensis Disney et al. This is the first record of A. ambiguus, an important pest of pine plantations, being a host of phorids. We found the first cases of gregarious parasitoidism of a parasitoid with no‐free pupae: M. catharinensis on A. lundii ants. We also evaluated the effect on pupal survival of two cosmopolitan fungi, Beauveria bassiana (Balsamo) Vuill. and Trichoderma lentiforme (Rehm) Chaverri et al., which are also considered as potential biological control agents of leafcutter ants. Suspensions of conidia were tested on Myrmosicarius and Apocephalus parasitoid pupae of different ages. No negative effect on pupal survival was found under either fungal treatment. Therefore, the use of both natural enemies separately but simultaneously, or the introduction of fungi‐inoculated pupae in the pest habitats, seems to be a promising strategy for the multiple biological control of leafcutter ants.     

Cascading trait‐mediated interactions induced by ant pheromones

Trait‐mediated indirect interactions (TMII) can be as important as density‐mediated indirect interactions. Here, we provide evidence for a novel trait‐mediated cascade (where one TMII affects another TMII) and demonstrate that the mechanism consists of a predator eavesdropping on chemical signaling. Ants protect scale insects from predation by adult coccinellid beetles – the first TMII. However, parasitic phorid flies reduce ant foraging activity by 50% – the second TMII, providing a window of opportunity for female beetles to oviposit in high‐quality microsites. Beetle larvae are protected from ant predation and benefit from living in patches with high scale densities. We demonstrate that female beetles can detect pheromones released by the ant when attacked by phorids, and that only females, and especially gravid females, are attracted to the ant pheromone. As ants reduce their movement when under attack by phorids, we conclude that phorids facilitate beetle oviposition, thus producing the TMII cascade.     

Parasitoid–host relationships when host size varies: the case of Pseudacteon flies and Solenopsis fire ants

Abstract. 1. Phorid flies in the genus Pseudacteon are parasitoids of ants. Variation in host size preferences of four South American and two North American Pseudacteon species on monogyne and polygyne forms of their host Solenopsis species [ S. invicta Buren and S. geminata (F.), respectively] was documented.
2. Monogyne Solenopsis workers were, on average, significantly larger than polygyne workers, and the average size monogyne worker attacked was significantly larger than the average size polygyne worker attacked in four of the six Pseudacteon species.
3. Three South American Pseudacteon species attacked larger than average size workers, whereas one attacked smaller than average size workers, in both monogyne and polygyne forms. Both North American Pseudacteon species attacked larger than average size polygyne workers and smaller than average size monogyne workers.
4. Three Pseudacteon species were reared from eggs to adults in infected ants in the laboratory. The size of the emergent phorid fly was related positively to the size of the host worker ant, with females emerging from larger hosts. Similar patterns were documented for both monogyne and polygyne forms.
5. The mean size of worker host from which phorids emerged did not differ significantly between the monogyne and polygyne forms in the subsample of phorids reared to adults.
6. The observed patterns elucidate factors that may cause variation in Pseudacteon sex ratios, and have implications for biological control efforts of pest Solenopsis species.     

Hitchhiking behaviour in leaf-cutter ants: An experimental evaluation of three hypotheses

In leaf-cutter ants, small workers often ride or “hitchhike” on leaf fragments carried back to the nest by larger foragers. There are several possible explanations for this unusual behaviour, the main ones being defence against phorid flies, defence against fungal contaminants, and leaf sap obtention. Here we tested these three hypotheses using standardized paired assays with laboratory colonies of Atta sexdens and field colonies of Atta laevigata. For both ant species, the proportion of fragments with hitchhikers increased significantly in response to the presence of fungal contaminants and to leaf sap. The proportion of fragments with hitchhikers also increased in the presence of phorid flies (Neodohrniphora erthali), but only for A. sexdens. In addition, hitchhiker position varied with the treatment applied. In the presence of phorids, hitchhikers frequently moved around the fragment; in the presence of leaf sap, hitchhikers frequently stood at the edge of the fragments, whereas when fragments were inoculated with fungal contaminants, hitchhikers were more often in the centre of the fragment, where the contamination was greatest. Our results strongly suggest that hitchhiking behaviour in Atta has multiple functions. Such behaviour probably evolved as a mechanism of defence, both against potential contaminants of the symbiotic fungus and against ant parasitoids. The obtention of leaf sap by minims seems to be a secondary and probably opportunistic function of this behaviour. Received 7 November 2005; revised 16 March 2006; accepted 6 April 2006.     

Indirect effects of phorid fly parasitoids on the mechanisms of interspecific competition among ants

Indirect effects, which occur when the impact of one species upon another requires the existence of an intermediary species, are apparently very common and may be of greater magnitude than direct effects. Behaviorally mediated indirect effects occur when one species affects the behavior of a second, which in turn affects how that species interacts with a third. I studied behaviorally mediated indirect effects on the mechanisms of competition in two congeneric fire ant species in the presence and absence of parasitoid phorid flies, which parasitized only one ant species. In observational and experimental field studies, the presence of native Texas phorid flies in the genus Pseudacteon decreased food retrieval by their host, Solenopsis geminata (F.), by as much as 50%. In the presence of phorid flies, many S. geminata workers assumed a stationary, curled defensive posture and did not forage. Although the phorid parasitoids had a relatively large effect on exploitative competition, there was no measurable effect on interference competition. Fierce interspecific aggression was observed between S. geminata and S. invicta Buren, and the presence of phorids had no effect on the outcome of these contests. The indirect effects of Pseudacteon parasitoids on Solenopsis fire ant resource retrieval appear to be larger than the direct effect of mortality. Some aspects of the foraging behavior of these Solenopsis species may be, in part, evolutionary adaptations to phorid parasitoid pressure. Because of the relatively large indirect effects, South American Pseudacteon phorids may be promising biocontrol agents of imported fire ants, S. invicta, in the USA. In a laboratory study, a single South American Pseudacteon female was able to significantly decrease food retrieval rates of a North American population of the imported fire ant, S. invicta. Received: 11 May 1998 / Accepted: 18 April 1999     

Quarantining behaviour of ants towards infected aphids as an antifungal mechanism in ant–aphid interactions

The ecological success of social insects, including ants, is tightly connected with their ability to protect themselves and their food resources. In exchange for energy‐rich honeydew, ants protect myrmecophilous aphids from various natural enemies. Fungal infection can have disastrous consequences for both mutualist partners, wherein aphids can be disease vectors. Behavioural responses towards fungus‐infected aphids of ant species in nature have scarcely been studied. Here, we studied the behaviour of honeydew foragers of four ant species – Formica polyctena Foerster, Formica rufa L., Formica pratensis Retzius (Hymenoptera: Formicidae, Formicini), and Lasius niger (L.) (Formicidae, Lasiini) – towards Symydobius oblongus (von Heyden) aphids contaminated with the generalist fungal pathogen Beauveria bassiana (Balsamo‐Crivelli) Vuillemin in the field. Aphid milkers from Formica spp. quickly detected and removed infected aphids from the host plant (Betula pendula Roth., Betulaceae). Neither ant species, the degree of aphid‐milker specialization (medium or high), nor the number of honeydew foragers had significant effects on the behaviour of Formica milkers towards infected aphids. Unlike Formica ants, L. niger usually displayed non‐aggressive behaviour (tolerance, antennation, honeydew collection, grooming). By the immediate removal of infected insects, Formica ants seem to minimize the probability of infection of symbionts as well as themselves. Quarantining behaviour may play an important role in ant–aphid interactions as a preventive antifungal mechanism formed under parasite pressure and thus contributing to the ecological success of ants.     

Ecological interactions of Pseudacteon parasitoids and Solenopsis ant hosts: environmental correlates of activity and effects on competitive hierarchies

1. Solenopsis (Hymenoptera: Formicidae) fire ants are host to Pseudacteon (Diptera: Phoridae) parasitoids. The activity of S. geminata (F.) hosts and relative abundance of Pseudacteon phorids, along with five environmental variables, were measured at weekly intervals over an 8‐month period at two sites. 2. Pseudacteon relative abundances often varied greatly from week to week, and were only weakly positively correlated with S. geminata activity. 3. A quadratic function of soil temperature was the single best predictor of ant activity at both sites, explaining 32 and 73% of the variation in ant activity. A linear function of soil moisture was the single best predictor of phorid relative abundance at one site (r² = 0.23) whereas no measured variables were significant predictors of phorid relative abundance at the other site. 4. Interspecific interactions at 600 baits were monitored at a third site to document dominance hierarchies and determine whether the presence of Pseudacteon phorids mediated interspecific interactions in their host, S. geminata. 5. Solenopsis geminata was near the top of dominance hierarchies, which did not diverge greatly from a linear pattern. Three species (S. geminata, S. invicta Buren, and Crematogaster laeviuscula Mayr) won the majority of their interspecific interactions and appear to be co‐dominants at this microhabitat‐rich site. 6. Overall, the presence of phorids had no significant effect on the outcome of interspecific contests involving S. geminata and all other ant species grouped together. Phorids may have contributed to some of the S. geminata losses against other co‐dominant species.     

Behavioural and olfactory variations in the leek moth, Acrolepiopsis assectella, after several generations of rearing under diverse conditions

Females of the parasitic phorid Neodohrniphora sp. were collected in the field and released singly inside an observation chamber placed between a laboratory colony of Atta sexdens (L.) and its foraging arena. The number and speed of loaded and unloaded ants returning to the nest, the weight of foragers and their loads, the number of leaf fragments abandoned by ants, and the number of small workers 'hitchhiking' on leaf fragments were measured before phorids were released, while they were in the observation chamber, and after they were removed. Relatively few ants were attacked by Neodohrniphora sp., but the presence of flies prompted outbound ants to return to the nest and caused a significant reduction on the number and mass of foragers. Additionally, the weight of leaf fragments transported by ants was reduced and the number of abandoned fragments increased in response to Neodohrniphora sp. Presence of the parasitoid caused no significant changes in the number of hitchhiking ants. The regular ants' traffic was resumed after phorids were removed, but foraging activity remained below normal for up to three hours. In the field A. sexdens forages mostly at night, but colonies undergo periods of diurnal foraging during which ants are subject to parasitism from several species of phorid flies. Considering that daytime foraging may be necessary for nutritional or metabolical needs, phorids may have a significant impact on their hosts by altering their foraging behavior regardless of the numerical values of parasitism.     

Interactions between ant species increase arrival rates of an ant parasitoid

Behaviours or traits associated with aggression and communication may increase an animal's conspicuousness to predators or parasitoids. Most examples of this come from instances of aggression or communication within a species. We tested whether interspecific encounters between ants enhance the host location success of a parasitoid (Diptera: Phoridae) that attacks ants in the genus Linepithema. At food resources recruited to by Linepithema, parasitoid discovery rates were lower when Linepithema was alone than when other ant species were present. In experimentally controlled encounters, parasitoid discovery rates were elevated when Linepithema confronted an ant species that elicited use of chemicals, but not when it confronted an ant species that primarily elicited physical aggression. These results indicate that phorid parasitoids of Linepithema use the ant's chemicals as host location cues. Because Linepithema is known to abandon food resources in the presence of its phorid parasitoids, its use of chemicals during interspecific encounters may diminish its competitive success when phorids are nearby. Copyright 2003 Published by Elsevier Science Ltd on behalf of The Association for the Study of Animal Behaviour.      

Foraging ecology and patterns of diversification in dipteran parasitoids of fire ants in south Brazil

1. At least sixteen species of parasitoid flies in the genus Pseudacteon (family Phoridae) attack fire ants in the Solenopsis saevissima subcomplex in South America. Little is known of behavioural or ecological differences among Pseudacteon parasitoids of fire ants, although their coexistence in multispecies communities would suggest that important differences exist. Seven Pseudacteon species in two separate communities were studied in south-east Brazil. The way in which hosts detect and respond to the presence of parasitoids, attack rates of the parasitoids, and host location behaviour of the parasitoids were examined.
2. Reductions in fire ant recruitment were more closely related to the number of ants attacked along a foraging trail than to the amount of time that a phorid was present.
3. Pseudacteon solenopsidis differed from other phorid species by flying backwards while pursuing ants, by attacking at lower rates than other phorids, and by spending longer around fire ant foraging trails than other phorids before departing. Fire ant recruitment to food often rebounded in the continued presence of P. solenopsidis.
4. In each of the two communities, certain Pseudacteon species appeared frequently at Solenopsis foraging trails, whereas others appeared predominantly at mound disturbances. Two distinct size classes of phorids were present in each community, and the community with the larger ant host species also had a third and larger phorid species. No phorid species from the same community had similar body sizes and similar host location behaviours, although numerous species from different communities shared both of these traits.
5. Heterogeneity in host size and in the ecological circumstances under which hosts are vulnerable to attack appears to have influenced the evolution and perhaps maintenance of diverse Pseudacteon communities.     

Colony-level impacts of parasitoid flies on fire ants

The red imported fire ant is becoming a global ecological problem, having invaded the United States, Puerto Rico, New Zealand and, most recently, Australia. In its established areas, this pest is devastating natural biodiversity. Early attempts to halt fire ant expansion with pesticides actually enhanced its spread. Phorid fly parasitoids from South America have now been introduced into the United States as potential biological control agents of the red imported fire ant, but the impact of these flies on fire ant populations is currently unknown. In the laboratory, we show that an average phorid density of as little as one attacking fly per 200 foraging ants decreased colony protein consumption nearly twofold and significantly reduced numbers of large-sized workers 50 days later. The high impact of a single phorid occurred mainly because ants decreased foraging rates in the presence of the flies. Our experiments, the first (to our knowledge) to link indirect and direct effects of phorids on fire ants, demonstrate that colonies can be stressed with surprisingly low parasitoid densities. We interpret our findings with regard to the more complex fire ant-phorid interactions in the field.     

A single lycaenid caterpillar gets an ant‐constructed shelter and uninterrupted ant attendance

Ant‐lycaenid associations range from mutualism to parasitism and the caterpillars of some species of lycaenids are reported to enter ant nests for shelter, diapause, or pupation. The present study aimed to examine the nature of the association between Euchrysops cnejus (Fabricius) (Lepidoptera: Lycaenidae) and Camponotus compressus (Fabricius) (Hymenoptera: Formicidae) worker ants on the extrafloral nectary‐bearing cowpea plant, Vigna unguiculata (L.) Walp. (Fabaceae). The abundance patterns of the ants and the lycaenid caterpillars together with the spatial patrolling patterns of the ants on the plants revealed that ant abundance increased with the occurrence of the lycaenid caterpillars and the ants preferred the lycaenids over the extrafloral nectar. Camponotus compressus worker ants constructed a shelter at the cowpea plant base after interaction with one or more lycaenid caterpillar(s) and tended the caterpillars and pupae till the emergence of the butterfly. The ant‐constructed shelters (ACSs) inhabited by the minor caste workers (13 ± 1.3 ants per ACS), were utilized by the caterpillars to undergo pupation. The ants confined their activities predominantly to tending the pod‐feeding caterpillars and the solitary pupa within each ACS. It appears that the behavior of the tending worker ants is modulated by the lycaenid vulnerable stages.     

Phorid fly parasitism rates vary between individual colonies of the leaf-cutting ant Atta laevigata

1. Phorid flies are important natural enemies of leaf-cutting ants. Female phorids lay eggs in the worker ants' bodies and the parasitoid larvae feed on the host's internal tissues.
2. Here we investigated the parasitism rates by four species of phorids when attacking individual Atta laevigata colonies at three different field sites in order to further understand the host-parasitoid relationship.
3. There were distinct variations in parasitism rates when comparing phorid species attacking individual colonies. In 13 of the colonies sampled, parasitism by Apocephalus attophilus or Eibesfeldtphora erthali was predominant. However, in four of the colonies, Myrmosicarius grandicornis and Apocephalus vicosae were the predominant species.
4. Variations in parasitism rates by the four phorid species were investigated in relation to the size of the potential hosts that had been collected from the foraging trails of each colony.
5. When correlating the size of potential hosts to phorid species, ants with head capsule widths of >2 mm were predominantly parasitized by A. vicosae and E. erthali. Smaller ants, with head capsule widths of 2 mm or less, were predominantly parasitized by A. attophilus and M. grandicornis.
6. The highest parasitism rates by E. erthali were observed in individual colonies that had the highest proportions of large workers on the trails.
7. Although almost half the size of E. erthali, A. attophilus parasitized both large (head capsule width > 2 mm) and small workers (<2 mm). This was related to the fact that between 1 and 16 larvae can develop within an individual host, depending on host size.
8. M. grandicornis parasitized smaller workers (mean head capsule width = 2 mm) as these phorids are themselves small and only one larva develops within each host.
9. This study demonstrated that the parasitism profiles of individual A. laevigata colonies were related to the size of foragers on the trails.

     

Introduction of the fire ant decapitating fly <Emphasis Type="Italic">Pseudacteon obtusus</Emphasis> in the United States: factors influencing establishment in Texas

The introduction and establishment of Pseudacteon obtusus Borgmeier (Diptera: Phoridae) into the United States is an important milestone in the effort to introduce a suite of parasitoids and pathogens as prospective biocontrol agents of the red imported fire ant, Solenopsis invicta Buren (Hymenoptera: Formicidae). P. obtusus is the fourth species of phorid fly to be established in the US and is now present in at least 11 sites in Texas. This species has a greater propensity for attacking foraging worker ants, and to a lesser degree it will attend mound disturbance events. The effectiveness of alternate field release strategies was evaluated across 28 sites in Texas, with establishment success 35 times more likely when no other phorid species were present at the outset of the releases. This result implies that competition between phorid species may interfere with establishment. Therefore release strategies should be directed to locations lacking other naturalized species or to times when abundances of other species are low.     

Biogeography and macroecology of phorid flies that attack fire ants in south-eastern Brazil and Argentina

Aim Saevissima group fire ants, Solenopsis richteri and S. invicta, have become serious pests when introduced from Argentina and Brazil to other continents. In South America, Solenopsis are distributed across a great variety of habitats and climates. In North America, S. invicta, introduced free of phorids, now ranges from coast to coast in the south. Success in introducing particular Pseudacteon as agents for the biological control of fire ants has varied across climatic zones. We aimed at assembling all the information about fire ant phorids from Argentina and Brazil, to estimate their richness and geographical ranges, to perform a climatic analysis for these distributions, to define groups and climate‐based communities, and to test and elucidate Rapoport's biogeographical rule. Location Argentina and Brazil (South America). Methods From field and museum collections and historical records, we developed a database of fire ant‐specific phorids throughout their known geographical range. A total of 123 sites with values for 15 climatic variables were mapped between 10° and 38° SL and between 35° and 65° C WL for the presence/absence of phorids. We calculated species richness across all sites combined, and for each phytogeographical region, using rarefaction curves, and ICE and Mmean estimators. We calculated mid‐latitudinal points, geographical ranges and areas for each species. The correlation between mid‐latitudinal point and ranges/areas was tested against a null model generated from the randomization of the raw distributional data. We used several types of multivariate analyses to distinguish groups of phorids by phytogeographical regions, hosts and climate, to find gradients of climate throughout the studied area, to define phorid communities in terms of their relationships with gradients of climate, and to test a mechanism for Rapoport's rule. Results Richness estimations using ICE and Mmean estimators were similar or higher than the observed values depending on the phytogeographical region. Cluster multivariate analyses based on climatic, phytogeographic and host data revealed distinct groupings of Pseudacton. The ‘cerrado’ group was confined to tropical savanna areas. A more ‘widespread’ group included ‘Chaco’ and ‘Maritime’ subgroups defined by their respective association with extreme temperatures or precipitation. Ordination multivariate analyses showed (1) two climatic gradients throughout the study area: one of temperature and the other of precipitation, and (2) that climatic variables significantly explained the observed assemblages of phorids. Positive and negative signs of the eigenvalues from the main axes of a canonical correspondence analysis allowed us to define eight communities whose geographical distribution resembled that of phytogeographical regions. We found a significant and positive correlation between geographical areas and mid latitudinal points, and furthermore, the Mantel test based on climatic variables suggested a mechanism for Rapoport's rule applying in the case of Pseudacteon. Main conclusions Pseudacteon species with greater mid‐latitudinal points occupy broader geographical areas and confront more stressful environmental conditions. Because the composition of Pseudacteon communities is largely determined by climatic variables, the correspondence between climates at sites of origin vs. sites of release should be an important consideration in choosing specific phorids for biocontrol efforts.     

The microsporidian pathogen Myrmecomorba nylanderiae: Intracolony transmission and impact upon tawny crazy ant (Nylanderia fulva) colonies

The microsporidian pathogen Myrmecomorba nylanderiae Plowes et al. infects introduced tawny crazy ants (Nylanderia fulva (Mayr)), and constitutes one of the first natural enemies known to attack this invasive ant. We assess how infection is transmitted within colonies and how infection impacts N. fulva colony fragment growth under carbohydrate‐deficient and carbohydrate‐sufficient dietary conditions. Carbohydrate scarcity is a common source of stress for ant colonies. Infected workers efficiently pass infection to developing larvae but all other potential pathways for within colony transmission are rare or non‐viable. For unknown reasons, queens within infected colony fragments are generally uninfected, limiting the role of transovarial transmission in intracolony transmission. In the laboratory, infection by M. nylanderiae primarily impacts the growth of N. fulva colonies by reducing pupal production. Colony growth showed a substantially greater impact under carbohydrate‐deficient conditions implying that the effect of the pathogen may depend on seasonally variable carbohydrate availability. In the colony growth assay, worker mortality did not differ with infection status under either nutrient regime. However, in a longer, direct test of survivorship, infected worker survivorship was significantly lower. Recently, some established N. fulva populations with high prevalence of M. nylanderiae infection have declined precipitously, though it is unknown if M. nylanderiae is a causative agent in these declines. The combination of chronic impacts, presence in North America, and potential association with population declines makes M. nylanderiae a promising prospect for the biological control of N. fulva.     

Ecological interactions among ants in the genus Linepithema, their phorid parasitoids, and ant competitors

Abstract. 1. Many ant species abandon foraging and retreat underground when parasitoids in the dipteran family Phoridae are present. Although the influence of phorids on ant foraging is well documented, their influence on interspecific competition is less studied. This study examined whether phorids influenced the competitive ability of host ants in the genus Linepithema at two sites in Brazil.
2. The phorid Pseudacteon lontrae attacked Linepithema piliferum at one site, while the phorid Pseudacteon pusillus attacked an unknown Linepithema ( Linepithema sp.) at the other site. Phorid parasitoids of Linepithema were far more common than phorids of other ant species.
3. Despite a high abundance of phorids, it was difficult to conclude that they influenced competition. Captures in pitfall traps indicated that host Linepithema were most active during times of day when phorids were inactive.
4. Camponotus rufipes and Brachymyrmex sp., the most common competitors of Linepithema sp. (60% of all interactions), dominated Linepithema sp. during the day regardless of phorids. Remaining ant species could not be evaluated individually because they interacted with Linepithema sp. infrequently.
5. Ectatomma brunneum was the most common competitor of L. piliferum (58% of all interactions). The high abundance of phorids at this site made it impossible to evaluate interactions between E. brunneum and L. piliferum in the absence of phorids.
6. Phorids seldom influenced exploitative competition by causing host Linepithema to abandon the bait when no ant competitors were present.