The Ecological Significance of Aphid Cornicles and Their Secretions.

Aphid cornicles are abdominal appendages that secrete an array of volatile and nonvolatile compounds with diverse ecological functions. The emission of alarm pheromones yields altruistic benefits for clone-mates in the aphid colony, which is essentially a superorganism with a collective fate. Secreted droplets also contain unsaturated triglycerides, fast-drying adhesives that can be lethal when smeared on natural enemies but more often impede their foraging efficiency. The longest cornicles have evolved in aphids that feed in exposed locations and are likely used to scent-mark colony intruders. Reduced cornicles are associated with reliance on alternative defenses, such as the secretion of protective waxes or myrmecophily. Root-feeding and gall-forming lifestyles provide protected feeding sites and are associated with an absence of cornicles. In some eusocial gall-formers, soldier morphs become repositories of cornicle secretion used to defend the gall, either as menopausal apterae that defend dispersing alatae or as sterile first instars that dispatch predators with their stylets and use cornicle secretions as a construction material for gall repair. Collectively, the evidence is consistent with an adaptive radiation of derived cornicle functions molded by the ecological lifestyle of the aphid lineage.

Mixed pollination system and floral signals of Paepalanthus (Eriocaulaceae): insects and geitonogamy ensure high reproductive success.

BACKGROUND AND AIMS: Eriocaulaceae exhibit a great variety of floral traits associated with insect (e.g. nectariferous structures) and wind pollination (unisexual flowers, exposed sexual organs and small pollen grains), as well as the 'selfing syndrome' (small flowers, short distance between stigma and anthers, and temporal overlap of male and female phases). Paepalanthus bifidus, P. subtilis and P. tortilis are related species that differ in form, size and colour of floral structures. We aimed to investigate the pollination and reproductive biology of these three species. METHODS: We analysed the floral biology, floral visitors, pollinator behaviour, and the contribution of insects, wind and spontaneous geitonogamy to fruit set. We also evaluated the floral colour and scent of the species. Colour reflectance of capitula of each species was measured and plotted in models of insect vision. Floral scent samples were extracted and the compounds were compared to vegetative scent samples. KEY RESULTS: In all species, the staminate and pistillate flowers are arranged in alternating cycles with a temporal overlap between these phases. Ants were the most frequent floral visitors and were effective pollinators in P. bifidus and P. tortilis, while flies were occasional pollinators in P. tortilis. Floral visitors were not observed in P. subtilis. In all species, fruits were produced by spontaneous geitonogamy, with no evidence of wind pollination. According to the models of insect vision, the colours of the capitula of P. bifidus and P. subtilis are the most inconspicuous for ants and flies. We found no difference between the emission of volatiles of inflorescences and vegetative structures. CONCLUSIONS: This study suggests that ant pollination might be more widespread in Eriocaulaceae than currently assumed. Furthermore, for small monocarpic plants, mixed mating strategies are most favourable, by ensuring reproduction either by outcrossing when pollinators are abundant or by spontaneous geitonogamy when pollinations are scarce/absent.

Nectaries in ferns: their taxonomic distribution, structure, function, and sugar composition.

PREMISE: Extrafloral nectaries have mainly been studied in angiosperms, but have also been reported in 39 fern species. Here we provide a global review of nectaries in ferns and examined their structure, function, and nectar sugar composition in two genera. METHODS: We searched in the literature and living plant collections of botanical gardens for indications of fern nectaries, observed nectar-feeding animals, studied the morphoanatomy in the two genera Aglaomorpha and Campyloneurum, and analyzed the total sugar concentrations and ratios of 16 species. Diurnal nectar release was observed with time-lapse photography. RESULTS: We found evidence for nectaries in 101 species of ferns from 11 genera and 6 families. Most of the nectary-bearing species were tree ferns (Cyatheaceae) and epiphytic ferns of the family Polypodiaceae. Nectaries consisted of cytoplasm-rich parenchyma with large nuclei and an epidermis with or without stomata, were attached to amphiphloic vascular bundles, and released nectar on the lower leaf surface mainly on expanding leaves during the night. Sugar concentrations varied between species (3.8-15.3%) but not between genera, and were sucrose-dominant (3 spp.), sucrose-rich (7), or hexose-rich (3). In the greenhouse, introduced ants, scale insects, and snails fed on the nectar. CONCLUSIONS: The wide taxonomic distribution, variable morphology, locations, and sugar compositions point to multiple evolutionary origins of fern nectaries. Nectar release in young leaves might attract mutualistic ants to protect leaves against herbivores only during this most vulnerable developmental stage. Even ex-situ, fern nectar is a valuable food source because it attracted several opportunistic animal species.

Temporal variation in the effect of ants on the fitness of myrmecophilic plants: seasonal effect surpasses periodic benefits.

Plants provide extrafloral nectar, which is a food resource taken by ants, especially aggressive species that may act as plant guards. To our knowledge, no study has been conducted to concurrently investigate the fluctuation of plant fitness over its whole reproductive season, recording and comparing both short periods (different samplings during the plant's reproductive season) and the season/pooled data (all fruits produced during the reproductive season). Here, by assigning plants to either ant-present or absent treatments, we investigated the influence of the protective foliage-dwelling ant, Camponotus crassus, on the flower bud and fruit production of four extrafloral nectaried plants (Ancistrotropis firmula, Bionia coriacea, Cochlospermum regium, and Peixotoa tomentosa) throughout their annual reproductive season. Periodic samples in the field revealed a large variation in plant reproduction throughout the season; the increases in buds and fruits were not constantly higher in plants with ants, and in fact, plants without ants had more reproductive structures sometimes. Nonetheless, the examination of the pooled data, i.e., cumulative number of flower buds and fruits produced during the reproductive season, revealed the plants with ants produced more flower buds and fruits (e.g., up to two-fold greater in A. firmula) compared to ant-absent treatments. Our results indicate the effects of ants on plant reproduction are not constant over time, but the net benefits to plants with ants are reflected in increased fruit production. Therefore, the investigations of the benefit of ants on plants should consider the whole plant's reproductive season rather than single samplings within plant reproduction period.

Protective Benefits of Tending Ants to a Critically Endangered Butterfly.

Ants provide protection to various organisms via myrmecophilous relationships. Most notably, ants and several butterfly species are involved in mainly mutualistic interactions. Previous field studies have shown that butterfly larval survival is increased in the presence of tending ants, suggesting that ants are providing protection against insect predation or parasitism. Here, we conducted a series of timed observational trials under laboratory conditions to assess larval survival and ant protection from insect predators for a myrmecophilous lycaenid butterfly. We focused on a critically endangered butterfly, the Miami blue (Cyclargus thomasi bethunebakeri) (Comstock and Huntington) (Lepidoptera: Lycaenidae), and its most common ant associate, the Florida carpenter ant (Camponotus floridanus) (Buckley) (Hymenoptera: Formicidae), to test this assumption of ant protection. We found that ants provide significant protection to Miami blue larvae, with later instar larvae receiving a higher level of protection due to differences in tending frequencies. These results will aid in informing conservation management and future organism reintroductions for this endangered butterfly.

Pollen adaptation to ant pollination: a case study from the Proteaceae.

BACKGROUND AND AIMS: Ant-plant associations are widely diverse and distributed throughout the world, leading to antagonistic and/or mutualistic interactions. Ant pollination is a rare mutualistic association and reports of ants as effective pollinators are limited to a few studies. Conospermum (Proteaceae) is an insect-pollinated genus well represented in the south-western Australia biodiversity hotspot, and here we aimed to evaluate the role of ants as pollinators of C. undulatum. METHODS: Pollen germination after contact with several species of ants and bees was tested for C. undulatum and five co-flowering species for comparison. We then sampled the pollen load of floral visitors of C. undulatum to assess whether ants carried a pollen load sufficient to enable pollination. Lastly, we performed exclusion treatments to assess the relative effect of flying- and non-flying-invertebrate floral visitors on the reproduction of C. undulatum. For this, we measured the seed set under different conditions: ants exclusion, flying-insects exclusion and control. KEY RESULTS: Pollen of C. undulatum, along with the other Conospermum species, had a germination rate after contact with ants of ~80 % which did not differ from the effect of bees; in contrast, the other plant species tested showed a drop in the germination rate to ~10 % following ant treatments. Although ants were generalist visitors, they carried a pollen load with 68-86 % of suitable grains. Moreover, ants significantly contributed to the seed set of C. undulatum. CONCLUSIONS: Our study highlights the complexity of ant-flower interactions and suggests that generalizations neglecting the importance of ants as pollinators cannot be made. Conospermum undulatum has evolved pollen with resistance to the negative effect of ant secretions on pollen grains, with ants providing effective pollination services to this threatened species.

Ant pollination of Paepalanthus lundii (Eriocaulaceae) in Brazilian savanna.

BACKGROUND AND AIMS: Ant-plant associations are widely diverse and distributed throughout the world, leading to complex ecological networks. Regarding ant-plant mutualism, ant pollination is a very rare interaction and few studies have shown the role of ants as pollinators. Therefore, we aimed to evaluate the role of ants as effective pollinators of Paepalanthus lundii (Eriocaulaceae) in a Brazilian savanna. METHODS: Fieldwork with experimental manipulation was conducted to evaluate the fitness of P. lundii, considering potential pollinators. For this, we mainly observed the number of seeds produced in different conditions: control, ant exclusion, exclusion of flying insects, and exclusion (entomophily test) of both ants and flying insects. Furthermore, we evaluated all floral visitors throughout the day, stigma receptivity, the numbers of male and female flowers, and patterns of species co-occurrence, which can indicate the presence of different pollinators in the plants at the same time. KEY RESULTS: We observed a relation between seed production and ant visits; Camponotus crassus was the most frequent floral visitor and the most effective pollinator. Also, we observed a statistical difference between the numbers of male and female flowers produced, with a greater number of male flowers. Furthermore, P. lundii presented flowering asynchrony, with 12 different types of maturation sequence, which indicates a cross-pollination system. Lastly, we observed an overlap of the greatest abundance of C. crassus and the time of plant stigmatic receptivity, and a pattern of non co-occurrence of ants, which shows the pollinator role of this ant. CONCLUSIONS: Our data provide evidence that previous generalizations neglecting the importance of ants as pollinators are wrong. Brazilian savanna can reveal a lot about the ant-pollination syndrome, since this environment presents peculiar characteristics related to this association. Thus, this study has great significance for the understanding of the ant-pollination syndrome, and for the understanding of the complex ecological networks present in these dry arid systems.

Resource overlap and dilution effects shape host plant use in a myrmecophilous butterfly.

The effects of consumers on fitness of resource organisms are a complex function of the spatio-temporal distribution of the resources, consumer functional responses and trait preferences, and availability of other resources. The ubiquitous variation in the intensity of species interactions has important consequences for the ecological and evolutionary dynamics of natural populations. Nevertheless, little is known about the processes causing this variation and their operational scales. Here, we examine how variation in the intensity of a consumer-resource interaction is related to resource timing, resource density and abundance of other resources. Using the butterfly consumer Phengaris alcon and its two sequential resources, the host plant Gentiana pneumonanthe and the host ants Myrmica spp., we investigated how butterfly egg-laying depended on focal host plant phenology, density and phenology of neighbouring host plants and host ant abundance. Butterflies preferred plants that simultaneously maximized the availability of both larval resources in time and space, that is, they chose early-flowering plants that were of higher nutritional quality for larvae where host ants were abundant. Both the probability of oviposition and the number of eggs were lower in plant individuals with a high neighbour density than in more isolated plants, and this dilution effect was stronger when neighbours flowered early. Our results show that plant-herbivore interactions simultaneously depend on the spatio-temporal distribution of a focal resource and on the small-scale spatial variation in the abundance of other herbivore resources. Given that consumers have negative effects on fitness and prefer certain timing of the resource organisms, this implies that processes acting at the levels of individuals, populations and communities simultaneously contribute to variation in consumer-mediated natural selection.

Ecological specialization is associated with genetic structure in the ant-associated butterfly family Lycaenidae.

The role of specialization in diversification can be explored along two geological axes in the butterfly family Lycaenidae. In addition to variation in host-plant specialization normally exhibited by butterflies, the caterpillars of most Lycaenidae have symbioses with ants ranging from no interactions through to obligate and specific associations, increasing niche dimensionality in ant-associated taxa. Based on mitochondrial sequences from 8282 specimens from 967 species and 249 genera, we show that the degree of ecological specialization of lycaenid species is positively correlated with genetic divergence, haplotype diversity and an increase in isolation by distance. Nucleotide substitution rate is higher in carnivorous than phytophagous lycaenids. The effects documented here for both micro- and macroevolutionary processes could result from increased spatial segregation as a consequence of reduced connectivity in specialists, niche-based divergence or a combination of both. They could also provide an explanation for the extraordinary diversity of the Lycaenidae and, more generally, for diversity in groups of organisms with similar multi-dimensional ecological specialization.

Molecular phylogenetics of Aspidiotini armored scale insects (Hemiptera: Diaspididae) reveals rampant paraphyly, curious species radiations, and multiple origins of association with Melissotarsus ants (Hymenoptera: Formicidae).

The armored scale insect tribe Aspidiotini comprises many pest species that are globally invasive and economically damaging. The taxonomy of scale insects is based almost solely upon morphological characters of adult females, and little prior work has been done to test the classification of aspidiotines against molecular evidence. To address these concerns, we reconstruct a molecular phylogeny for aspidiotine armored scales that expands greatly upon taxonomic and character representations from previous studies. Our dataset includes 127 species (356 terminal taxa) and four gene regions: 28S, EF-1α, COI-COII, and CAD. Nearly 50% of the species treated are identified as pests and several more may represent emerging pests. Phylogenetic data were analyzed in a Bayesian framework using MC3 iterations. The majority of sampled aspidiotine genera are not monophyletic as currently defined. Monophyly constraints for 'worst offenders' were imposed on the phylogeny and stepping-stone MCMC was performed to calculate marginal likelihood scores. Comparisons of marginal likelihoods from runs with constrained vs. informative priors support the interpretation that pest-rich genera are not monophyletic. We use character mapping to illustrate signal and convergence for selected traits that have been used to define or recognize genera and evaluate consistency and retention indices for these traits. The phylogeny illustrates a pervasive pattern in which extremely polyphagous pests - typically having large populations and wide geographical distributions - are frequently intertwined with range-limited specialists on the phylogeny. Finally, the phylogeny recovers three origins of ant association among the Aspidiotini. The history of ant/diaspidid symbioses involves periods of sustained partner fidelity, spanning multiple speciation events, which have been punctuated by opportunistic switches to novel partners.

Caterpillar seed predators mediate shifts in selection on flowering phenology in their host plant.

Variation in selection among populations and years has important implications for evolutionary trajectories of populations. Yet, the agents of selection causing this variation have rarely been identified. Selection on the time of reproduction within a season in plants might differ both among populations and among years, and selection can be mediated by both mutualists and antagonists. We investigated if differences in the direction of phenotypic selection on flowering phenology among 20 populations of Gentiana pneumonanthe during 2 yr were related to the presence of the butterfly seed predator Phengaris alcon, and if butterfly incidence was associated with the abundance of the butterfly's second host, Myrmica ants. In plant populations without the butterfly, phenotypic selection favored earlier flowering. In populations where the butterfly was present, caterpillars preferentially attacked early-flowering individuals, shifting the direction of selection to favoring later flowering. Butterfly incidence in plant populations increased with ant abundance. Our results demonstrate that antagonistic interactions can shift the direction of selection on flowering phenology, and suggest that such shifts might be associated with differences in the community context.

Ant-lepidopteran associations along African forest edges.

Working along forest edges, we aimed to determine how some caterpillars can co-exist with territorially dominant arboreal ants (TDAAs) in tropical Africa. We recorded caterpillars from 22 lepidopteran species living in the presence of five TDAA species. Among the defoliator and/or nectarivorous caterpillars that live on tree foliage, the Pyralidae and Nymphalidae use their silk to protect themselves from ant attacks. The Notodontidae and lycaenid Polyommatinae and Theclinae live in direct contact with ants; the Theclinae even reward ants with abundant secretions from their Newcomer gland. Lichen feeders (lycaenid; Poritiinae), protected by long bristles, also live among ants. Some lycaenid Miletinae caterpillars feed on ant-attended membracids, including in the shelters where the ants attend them; Lachnocnema caterpillars use their forelegs to obtain trophallaxis from their host ants. Caterpillars from other species live inside weaver ant nests. Those of the genus Euliphyra (Miletinae) feed on ant prey and brood and can obtain trophallaxis, while those from an Eberidae species only prey on host ant eggs. Eublemma albifascia (Erebidae) caterpillars use their thoracic legs to obtain trophallaxis and trophic eggs from ants. Through transfer bioassays of last instars, we noted that herbivorous caterpillars living in contact with ants were always accepted by alien conspecific ants; this is likely due to an intrinsic appeasing odor. Yet, caterpillars living in ant shelters or ant nests probably acquire cues from their host colonies because they were considered aliens and killed. We conclude that co-evolution with ants occurred similarly in the Heterocera and Rhopalocera.

Dead ant walking: a myrmecophilous beetle predator uses parasitoid host location cues to selectively prey on parasitized ants.

Myrmecophiles (i.e. organisms that associate with ants) use a variety of ecological niches and employ different strategies to survive encounters with ants. Because ants are typically excellent defenders, myrmecophiles may choose moments of weakness to take advantage of their ant associates. This hypothesis was studied in the rove beetle, Myrmedonota xipe, which associates with Azteca sericeasur ants in the presence of parasitoid flies. A combination of laboratory and field experiments show that M. xipe beetles selectively locate and prey upon parasitized ants. These parasitized ants are less aggressive towards beetles than healthy ants, allowing beetles to eat the parasitized ants alive without interruption. Moreover, behavioural assays and chemical analysis reveal that M. xipe are attracted to the ant's alarm pheromone, the same secretion used by the phorid fly parasitoids in host location. This strategy allows beetles access to an abundant but otherwise inaccessible resource, as A. sericeasur ants are typically highly aggressive. These results are the first, to our knowledge, to demonstrate a predator sharing cues with a parasitoid to gain access to an otherwise unavailable prey item. Furthermore, this work highlights the importance of studying ant-myrmecophile interactions beyond just their pairwise context.

Trait-mediated indirect interactions of ant shape on the attack of caterpillars and fruits.

Mainly owing to their high diversity and abundance, ants are formidable as predators and defenders of foliage. Consequently, ants can exclude both invertebrate and vertebrate activity on plants via direct and indirect interactions as already shown in many previous studies. Here we present empirical evidence that objects resembling ant shape on dummy caterpillars were able to repel visually oriented predators. Moreover, we also show that rubber ants on dummy fruits can repel potential fruit dispersers. Our results have direct implications on the ecological and evolutionary dynamics of interactions in ant-based systems, as ant presence could affect the fitness of its partners. In short, our study highlights the importance of visual cues in interspecific interactions and opens a new way to study the effects of ant presence to test ecological and evolutionary hypotheses.

Chemical disguise of myrmecophilous cockroaches and its implications for understanding nestmate recognition mechanisms in leaf-cutting ants.

BACKGROUND: Cockroaches of the genus Attaphila regularly occur in leaf-cutting ant colonies. The ants farm a fungus that the cockroaches also appear to feed on. Cockroaches disperse between colonies horizontally (via foraging trails) and vertically (attached to queens on their mating flights). We analysed the chemical strategies used by the cockroaches to integrate into colonies of Atta colombica and Acromyrmex octospinosus. Analysing cockroaches from nests of two host species further allowed us to test the hypothesis that nestmate recognition is based on an asymmetric mechanism. Specifically, we test the U-present nestmate recognition model, which assumes that detection of undesirable cues (non-nestmate specific substances) leads to strong rejection of the cue-bearers, while absence of desirable cues (nestmate-specific substances) does not necessarily trigger aggression. RESULTS: We found that nests of Atta and Acromyrmex contained cockroaches of two different and not yet described Attaphila species. The cockroaches share the cuticular chemical substances of their specific host species and copy their host nest's colony-specific cuticular profile. Indeed, the cockroaches are accepted by nestmate but attacked by non-nestmate ant workers. Cockroaches from Acromyrmex colonies bear a lower concentration of cuticular substances and are less likely to be attacked by non-nestmate ants than cockroaches from Atta colonies. CONCLUSIONS: Nest-specific recognition of Attaphila cockroaches by host workers in combination with nest-specific cuticular chemical profiles suggest that the cockroaches mimic their host's recognition labels, either by synthesizing nest-specific substances or by substance transfer from ants. Our finding that the cockroach species with lower concentration of cuticular substances receives less aggression by both host species fully supports the U-present nestmate recognition model. Leaf-cutting ant nestmate recognition is thus asymmetric, responding more strongly to differences than to similarities.

Plant defences against ants provide a pathway to social parasitism in butterflies.

Understanding the chemical cues and gene expressions that mediate herbivore-host-plant and parasite-host interactions can elucidate the ecological costs and benefits accruing to different partners in tight-knit community modules, and may reveal unexpected complexities. We investigated the exploitation of sequential hosts by the phytophagous-predaceous butterfly Maculinea arion, whose larvae initially feed on Origanum vulgare flowerheads before switching to parasitize Myrmica ant colonies for their main period of growth. Gravid female butterflies were attracted to Origanum plants that emitted high levels of the monoterpenoid volatile carvacrol, a condition that occurred when ants disturbed their roots: we also found that Origanum expressed four genes involved in monoterpene formation when ants were present, accompanied by a significant induction of jasmonates. When exposed to carvacrol, Myrmica workers upregulated five genes whose products bind and detoxify this biocide, and their colonies were more tolerant of it than other common ant genera, consistent with an observed ability to occupy the competitor-free spaces surrounding Origanum. A cost is potential colony destruction by Ma. arion, which in turn may benefit infested Origanum plants by relieving their roots of further damage. Our results suggest a new pathway, whereby social parasites can detect successive resources by employing plant volatiles to simultaneously select their initial plant food and a suitable sequential host.

Rickia wasmannii increases the need for water in Myrmica scabrinodis (Ascomycota: Laboulbeniales; Hymenoptera: Formicidae).

The order Laboulbeniales (Fungi, Ascomycota) is a little-studied group of microscopic ectoparasites of invertebrates, mostly insects. The effects of Laboulbeniales species on their hosts are mostly unknown. Rickia wasmannii Cavara, 1899 is a common Laboulbeniales fungus occurring in Europe and is currently known to be a parasite of at least eight Myrmica ant species. R.wasmannii serves as a good model organism for Laboulbeniales-host interactions, as this species covers the host in a very high density, and infected host individuals can be easily collected in high numbers. The effect of R. wasmannii on the survival rate of its most common host species, Myrmica scabrinodis Nylander 1846, was therefore investigated in a laboratory experiment on an individual level. To enhance the results, environmental stresses were simulated by depriving infected and uninfected workers of water and food. The survival of individuals was recorded hourly until the death of the last individual. Infected specimens were significantly more sensitive to the withdrawal of food and water than uninfected specimens. When we tested for water consumption, we found that infected ants spent more time consuming water than uninfected ants. Therefore, it is possible that infected ants must replace the loss of water. Based on these results, R. wasmannii substantially decreases the chances and time of survival of infected individuals, at least in resource-limited environments, which suggest that R. wasmannii has a negative effect on its host.

An ant-associated mesostigmatid mite in Baltic amber.

Fossil mesostigmatid mites (Acari: Parasitiformes: Mesostigmata) are extremely rare, and specimens from only nine families, including four named species, have been described so far. A new record of Myrmozercon sp. described here from Eocene (ca 44-49 Myr) Baltic amber represents the first-and so far only-fossil example of the derived, extant family Laelapidae. Significantly, modern species of this genus are habitually myrmecophilous and the fossil mite described here is preserved attached to the head of the dolichoderine ant Ctenobethylus goepperti (Mayr, 1868). It thus offers the oldest unequivocal evidence for an ecological association between mesostigmatid mites and social insects in the order Hymenoptera.

Natural history and morphology of the hoverfly Pseudomicrodon biluminiferus and its parasitic relationship with ants nesting in bromeliads.

The syrphid subfamily Microdontinae is characterized by myrmecophily of their immature stages, i.e., they develop in ant nests. Data on natural history of microdontines are scarce, especially in the Neotropics. Based on fieldwork in southern Brazil, this study provided new data on development and ecology of the hoverfly Pseudomicrodon biluminiferus (Hull) (Diptera: Syrphidae) as well as the first morphological descriptions of male genitalia, larvae, and pupa. Immature specimens were specifically found in colonies of the ant species Crematogaster limata Smith (Hymenoptera: Formicidae) found in rosettes of the bromeliad species Aechmea lindenii (E. Morren) Baker (Poales: Bromeliaceae) and A. nudicaulis (L.) Grisebach. Third instar larvae were observed preying on ant larvae, revealing the parasitic nature of P. biluminiferus. In this and several other aspects, the natural history of P. biluminiferus is similar to that of Holarctic microdontine species. Exceptions include: (i) indications that adults of P. biluminiferus outlast the winter months (in contrast to 3(rd)instar larvae in Holarctic species) and (ii) P. biluminiferus' relationship with bromeliads. The importance of bromeliads for this host-parasite system is evaluated in this paper. The single occurrence of another, unidentified microdontine species' pupae in a nest of the ant species Camponotus melanoticus Emery (Hymenoptera: Formicidae) is reported.