Obligate chimerism in male yellow crazy ants.

Multicellular organisms typically develop from a single fertilized egg and therefore consist of clonal cells. We report an extraordinary reproductive system in the yellow crazy ant. Males are chimeras of haploid cells from two divergent lineages: R and W. R cells are overrepresented in the males' somatic tissues, whereas W cells are overrepresented in their sperm. Chimerism occurs when parental nuclei bypass syngamy and divide separately within the same egg. When syngamy takes place, the diploid offspring either develops into a queen when the oocyte is fertilized by an R sperm or into a worker when fertilized by a W sperm. This study reveals a mode of reproduction that may be associated with a conflict between lineages to preferentially enter the germ line.

The unusual genetics of invasive ants.

The males of an invasive ant species are chimeras of two distinct genetic lineages.

Basic ecology of ant gardens in a dry-premontane transitional forest / Ecología básica de jardines de hormigas en un bosque transicional seco-premontano

Abstract Introduction: "Ant gardens'' are ant nests located at different heights on trees on which vascular epiphytic plants that have been transported and sown by ants have germinated. Although this mutualistic relationship has been studied in humid tropical ecosystems, information on other tropical and Colombian ecosystems is scarce. Objective: To characterize the distribution, building, diversity, demography and phenology of ant gardens in dry tropical forest. Methods: In January and February, 2018, we identified 170 ant gardens on a 100x5 m transect on the banks of the Quesada River, Colombia, and in adjacent secondary dry premontane transitional forest; we monitored changes, for gardens and plants, in 28 of them, every two weeks (March 2018-February 2019). Results: The gardens, built by Azteca ulei, were aggregated near water bodies; had 10 species of epiphytes and were on 13 tree species. Larger gardens had more epiphyte species. Some epiphytes had a bimodal phenological pattern. Less seedlings become established in the dry season, and less adults remain in the gardens. Garden characteristics benefit both epiphytes and ants. Conclusions: The gardens built by A. ulei have ecological characteristics that favor the germination, establishment, and reproduction of diverse epiphytes in this dry tropical ecosystem, including aggregation near water flows.

Resumen Introducción: Los "jardines de hormigas'' son nidos de hormigas en árboles, que se localizan a diferentes alturas, sobre los que germinan plantas epífitas vasculares que han sido transportadas y sembradas previamente por hormigas. Aunque esta relación mutualista ha sido estudiada en ecosistemas húmedos tropicales, la información en otros ecosistemas tropicales y en Colombia, es escasa. Objetivo: Caracterizar la distribución, construcción, diversidad, demografía y fenología de los hormigueros en bosque tropical seco. Métodos: En enero y febrero de 2018, identificamos 170 hormigueros en un transecto de 100x5 m a orillas del río Quesada, Colombia, y en el bosque secundario premontano seco de transición adyacente; monitoreamos cambios, para jardines y plantas, en 28 de ellos, cada dos semanas (marzo 2018-febrero 2019). Resultados: Los jardines, construidos por Azteca ulei, se agregaron cerca de cuerpos de agua; tenían 10 especies de epífitas y estaban sobre 13 especies arbóreas. Los jardines más grandes tenían más especies epífitas. Algunas epífitas tuvieron un patrón fenológico bimodal. Se establecen menos plántulas en la estación seca y quedan menos adultos en los jardines. Las características del jardín benefician tanto a las epífitas como a las hormigas. Conclusiones: Los jardines construidos por A. ulei tienen características ecológicas que favorecen la germinación, establecimiento y reproducción de diversas epífitas en este ecosistema tropical seco, incluyendo la agregación cerca de cursos de agua.

The pupal moulting fluid has evolved social functions in ants.

Insect societies are tightly integrated, complex biological systems in which group-level properties arise from the interactions between individuals1-4. However, these interactions have not been studied systematically and therefore remain incompletely known. Here, using a reverse engineering approach, we reveal that unlike solitary insects, ant pupae extrude a secretion derived from the moulting fluid that is rich in nutrients, hormones and neuroactive substances. This secretion elicits parental care behaviour and is rapidly removed and consumed by the adults. This behaviour is crucial for pupal survival; if the secretion is not removed, pupae develop fungal infections and die. Analogous to mammalian milk, the secretion is also an important source of early larval nutrition, and young larvae exhibit stunted growth and decreased survival without access to the fluid. We show that this derived social function of the moulting fluid generalizes across the ants. This secretion thus forms the basis of a central and hitherto overlooked interaction network in ant societies, and constitutes a rare example of how a conserved developmental process can be co-opted to provide the mechanistic basis of social interactions. These results implicate moulting fluids in having a major role in the evolution of ant eusociality.

Aphids and Ants, Mutualistic Species, Share a Mariner Element with an Unusual Location on Aphid Chromosomes.

Aphids (Hemiptera, Aphididae) are small phytophagous insects. The aim of this study was to determine if the mariner elements found in the ant genomes are also present in Aphis fabae and Aphis hederae genomes and the possible existence of horizontal transfer events. Aphids maintain a relationship of mutualism with the ants. The close contact between these insects could favour horizontal transfer events of transposable elements. Myrmar mariner element isolated from Myrmica ruginodis and Tapinoma ibericum ants have also been found in the two Aphis species: A. fabae and A. hederae (Afabmar-Mr and Ahedmar-Mr elements). Besides, Afabmar-Mr could be an active transposon. Myrmar-like elements are also present in other insect species as well as in one Crustacean species. The phylogenetic study carried out with all Myrmar-like elements suggests the existence of horizontal transfer. Most aphids have 2n = 8 with a XX-X0 sex determination system. Their complicated life cycle is mostly parthenogenetic with sexual individuals only in autumn. The production of X0 males, originated by XX females which produce only spermatozoa with one X chromosome, must necessarily occur through specialized cytogenetic and molecular mechanisms which are not entirely known. In both aphid species, the mariner elements are located on all chromosomes, including the X chromosomes. However, on the two X chromosomes, no positive signals are detected in their small DAPI-negative telomere regions. The rDNA sites are located, as in the majority of Aphids species, on one of the telomere regions of each X chromosome. The hybridization patterns obtained by double FISH demonstrate that Afabmar-Mr and Ahedmar-Mr elements do not hybridize at the rDNA sites of their host species. Possible causes for the absence of these transposons in the rDNA genes are discussed, probably related with the X chromosome biology.

Trap rebuilding by Myrmeleon brasiliensis larvae (Neuroptera: Myrmeleontidae) in response to flooding: the effect of body size / Reconstrucción de trampas por larvas de Myrmeleon brasiliensis (Neuroptera: Myrmeleontidae) tras las inundaciones: efecto del tamaño corporal

Abstract Introduction: Immature forms of the antlion Myrmeleon brasiliensis (Neuroptera, Myrmeleontidae) build traps in dry sandy soil to capture prey. Objective: The aim of the present study was to investigate how the waterlogging of the soil due to rain affects the trap rebuilding and relocation behavior of M. brasiliensis of different sizes. Methods: The study was conducted between July and December 2019. Larvae M. brasiliensis were observed and collected from a forest reserve in the municipality of Aquidauana in the state of Mato Grosso do Sul, Brazil. Results: In the natural environment, most larvae rebuilt their traps in the same location seven days after the simulation of rain, with a smaller diameter than that observed prior to the simulation of rain. In the laboratory, the movements of M. brasiliensis larvae and rebuilding of the traps after the waterlogging of the soil was affected by body size. Larger larvae moved more and were more likely to rebuild their traps. Conclusions: The saturation of the soil affects the foraging of M. brasiliensis larvae, which are impeded from rebuilding their traps for a period. In situations of long periods of saturated soil, the mortality rate of the larvae is high and rebuilding of the traps occurs after the soil dries out, but with a smaller trap size. These data suggest that changes in the rainfall pattern can affect the population structure of M. brasiliensis larvae, with the selection of larger individuals in situations of more severe rains. In this process, the smaller larvae are more affected, as their foraging is impeded.

Resumen Introducción: Los insectos inmaduros de hormiga león de la especie Myrmeleon brasiliensis (Neuroptera, Myrmeleontidae), construyen trampas en el suelo arenoso seco para capturar a sus presas. Objetivo: El objetivo de este trabajo fue analizar cómo la inundación del suelo, como la lluvia, afecta al comportamiento de reconstrucción y de desplazamiento de las trampas de M. brasiliensis según las diferentes tallas. Métodos: El estudio fue conducido entre julio y diciembre del 2019. Las larvas fueron observadas y recolectadas en una Reserva Forestal en la ciudad de Aquidauana, Estado de Mato Grosso del Sur, Brasil. Resultados: Observamos que, en el ambiente natural, siete días después de la lluvia simulada, la mayoría de las larvas reconstruyeron a sus trampas en el mismo sitio y con diámetro más pequeño que el observado antes de la lluvia simulada. En el laboratorio fue observado que después de la inundación del suelo, el desplazamiento de las larvas y la reconstrucción de las trampas es afectado por el tamaño de su cuerpo. Así, larvas de M. brasiliensis más grandes se desplazan más y presentan una probabilidad más grande de reconstruir a sus trampas. Conclusión: De esa manera, la inundación del suelo afecta el forrajeo de las larvas M. brasiliensis, que se quedan impedidas por un tiempo durante la reconstrucción de sus trampas. Además, en situaciones de largos periodos de inundación del suelo, las larvas presentan una alta tasa de mortalidad y la reconstrucción de sus trampas ocurre después que se seca el suelo, pero esas trampas suelen ser más pequeñas. Esos datos sugieren que es posible que variaciones en el patrón de lluvia pueden afectar la estructura poblacional de las larvas M. brasiliensis, pues en situaciones de lluvias más fuertes, las larvas más grandes suelen tener más éxito. En este proceso, las larvas más pequeñas se ven más afectadas, ya que se impide su búsqueda de alimento.

Functional composition of ant assemblages in habitat islands is driven by habitat factors and landscape composition.

Fragmented natural habitats within human-transformed landscapes play a key role in preserving biodiversity. Ants as keystone species are essential elements of terrestrial ecosystems; thus, it is important to understand the factors influencing their presence. In a large-scale multi-site study, we surveyed ant assemblages using sweep netting and D-vac sampling on 158 ancient burial mounds preserving grassland habitats in agricultural landscapes in East-Hungary. We asked the following questions: (1) How do habitat factors and landscape composition affect species richness and functional diversity of ants? (2) Which ant traits are affected by habitat factors and landscape composition? Despite their small sizes, mounds as permanent and relatively undisturbed landscape elements could provide safe havens for diverse ant assemblages even in transformed agricultural landscapes. The complex habitat structure of wooded mounds supported high species and functional diversity of ant assemblages. Ant species on wooded mounds had small or medium-sized colonies, enabling the co-existence of more species. The effect of landscape composition on ant assemblages was mediated by habitat factors: steep slopes buffered the negative effect of the cropland matrix and enabled higher ant diversity.

Nano-CT imaging of larvae in the ant Pheidole hyatti reveals coordinated growth of a rudimentary organ necessary for soldier development.

The growth of imaginal discs in holometabolous insects is coordinated with larval growth to ensure the symmetrical and proportional development of the adult appendages. In ants, the differential growth of these discs generates distinct castes-the winged male and queen castes and the wingless worker caste. In the hyperdiverse ant genus Pheidole, the worker caste is composed of two morphologically distinct subcastes: small-headed minor workers and larger, big-headed, soldiers. Although these worker subcastes are completely wingless, soldier larvae develop rudimentary forewing discs that function in generating the disproportionate head-to-body scaling and size of soldiers. It remains unclear, however, how rudimentary forewing discs in soldier larvae are coordinated with other imaginal discs. Here we show, using quantitative nano-CT imaging and three-dimensional analyses, that the increase in the volume of the soldier rudimentary forewing discs is coordinated with larval size as well as with the increase in the volume of the leg and eye-antennal (head) discs. However, relative to larval size, we found that when the rudimentary forewing discs appear during the last larval instar, they are relatively smaller but increase in volume faster than that of the head (eye-antennal) and leg discs. These findings show that the rudimentary wing disc in soldier larvae has evolved novel patterns of inter-organ coordination as compared with other insects to generate the big-headed soldier caste in Pheidole. More generally, our study raises the possibility that novel patterns of inter-organ coordination are a general feature of rudimentary organs that acquire novel regulatory functions during development and evolution.

Tramtrack acts during late pupal development to direct ant caste identity.

A key question in the rising field of neuroepigenetics is how behavioral plasticity is established and maintained in the developing CNS of multicellular organisms. Behavior is controlled through systemic changes in hormonal signaling, cell-specific regulation of gene expression, and changes in neuronal connections in the nervous system, however the link between these pathways is unclear. In the ant Camponotus floridanus, the epigenetic corepressor CoREST is a central player in experimentally-induced reprogramming of caste-specific behavior, from soldier (Major worker) to forager (Minor worker). Here, we show this pathway is engaged naturally on a large genomic scale during late pupal development targeting multiple genes differentially expressed between castes, and central to this mechanism is the protein tramtrack (ttk), a DNA binding partner of CoREST. Caste-specific differences in DNA binding of ttk co-binding with CoREST correlate with caste-biased gene expression both in the late pupal stage and immediately after eclosion. However, we find a unique set of exclusive Minor-bound genes that show ttk pre-binding in the late pupal stage preceding CoREST binding, followed by caste-specific gene repression on the first day of eclosion. In addition, we show that ttk binding correlates with neurogenic Notch signaling, and that specific ttk binding between castes is enriched for regulatory sites associated with hormonal function. Overall our findings elucidate a pathway of transcription factor binding leading to a repressive epigenetic axis that lies at the crux of development and hormonal signaling to define worker caste identity in C. floridanus.

Mutualistic interaction of aphids and ants in pepper, Capsicum annuum and Capsicum frutenscens (Solanaceae) / Interacción mutualista de pulgones y hormigas en pimiento, Capsicum annuum y Capsicum frutenscens (Solanaceae)

Abstract Introduction: Adequate biological identification is fundamental for establishing integrated pest management programs and identifying the trophic and mutualist relationships that can affect pest population dynamics. Aphids are the main pest of pepper Capsicum spp. (Solanaceae) crops in Southwestern Colombia, due to their role as vectors of viruses. However, the identification of aphid species is complex, limiting the investigations performed to address their interactions with other organisms. Ants and aphids present a facultative mutualistic relationship, that promotes the growth of hemipteran colonies, for this reason, the study of the ecological mutualistic association between aphids and ants is important. Objective: The main objective was to discriminate the aphid species present in commercial crops of Capsicum spp., and to identify the ant community that attends the aphid colonies and its effects on the size of the aphid colonies. Methods: Aphid species, and their ant mutualist, were collected from Capsicum annuum and Capsicum frutescens, in the Cauca valley, Southwestern Colombia. We used the DNA barcoding approach to identify aphid species, and the ants were identified by morphology-based taxonomy. To evaluate the effect of ant care on the size and structure of aphid colonies, generalized linear models were calculated using as the response variables the total number of aphids for each colony and the proportion of nymphs. Results: The aphid species that attack pepper crops, are: Aphis gossypii and Myzus persicae (Hemiptera: Aphididae), with A. gossypii being the species that interacts with ants (19 ant species). A. gossypii colonies attended by ants had larger sizes and more nymphs per colony, than those not attended. Conclusions: Although the aphid-ant interaction is not species-specific, it is necessary to consider its role in the propagation of viral diseases in peppers and to determine how this interaction may affect regional biological control strategies.

Resumen Introducción: La adecuada identificación biológica es fundamental para establecer programas de manejo integrado de plagas e identificar las relaciones tróficas y mutualistas que pueden afectar la dinámica poblacional de insectos plaga. Los áfidos son las principales plagas del ají Capsicum spp. (Solanaceae) en el suroccidente colombiano, debido a su rol como vectores de virus. Sin embargo, su identificación es compleja, y limita las investigaciones que intentan revelar sus interacciones con otros organismos. Las hormigas y los áfidos presentan una relación mutualista facultativa, que promueve el crecimiento de las colonias de los hemípteros, por esta razón, el estudio de la asociación ecológica y mutualista entre áfidos y hormigas es importante. Objetivo: El principal objetivo de esta investigación fue discriminar las especies de áfidos presentes en cultivos comerciales de Capsicum spp., e identificar la comunidad de hormigas que atiende las colonias de áfidos y su efecto en el tamaño de las colonias de áfidos. Métodos: Los áfidos, y las hormigas mutualistas de estos áfidos, se recolectaron de Capsicum annuum y Capsicum frutescens, en el valle del rio Cauca, en el suroccidente colombiano. Se empleó el Código de barras del ADN para identificar las especies de áfidos, y las hormigas se identificaron empleando taxonomía basada en morfología. Para evaluar el efecto que tiene el cuidado de las hormigas sobre el tamaño de las colonias de áfidos, se empleó un modelo lineal generalizado, utilizando como variables de respuesta el número total de áfidos por cada colonia y la proporción de ninfas por colonia. Resultados: Las especies de áfidos que atacan los cultivos de ají, son: Aphis gossypii y Myzus persicae (Hemiptera: Aphididae), siendo A. gossypii la especie que interactúa con hormigas (19 especies). Las colonias de A. gossypii atendidas por hormigas presentan mayor tamaño y número de ninfas, que aquellas desatendidas. Conclusiones: Aunque la interacción áfido-hormiga no es especie específica, es necesario considerar su rol en la propagación de enfermedades virales en plantas cultivadas y determinar cómo esta interacción puede afectar la implementación de estrategias de control biológico.

Ever-increasing viral diversity associated with the red imported fire ant Solenopsis invicta (Formicidae: Hymenoptera).

BACKGROUND: Advances in sequencing and analysis tools have facilitated discovery of many new viruses from invertebrates, including ants. Solenopsis invicta is an invasive ant that has quickly spread worldwide causing significant ecological and economic impacts. Its virome has begun to be characterized pertaining to potential use of viruses as natural enemies. Although the S. invicta virome is the best characterized among ants, most studies have been performed in its native range, with less information from invaded areas. METHODS: Using a metatranscriptome approach, we further identified and molecularly characterized virus sequences associated with S. invicta, in two introduced areas, U.S and Taiwan. The data set used here was obtained from different stages (larvae, pupa, and adults) of S. invicta life cycle. Publicly available RNA sequences from GenBank's Sequence Read Archive were downloaded and de novo assembled using CLC Genomics Workbench 20.0.1. Contigs were compared against the non-redundant protein sequences and those showing similarity to viral sequences were further analyzed. RESULTS: We characterized five putative new viruses associated with S. invicta transcriptomes. Sequence comparisons revealed extensive divergence across ORFs and genomic regions with most of them sharing less than 40% amino acid identity with those closest homologous sequences previously characterized. The first negative-sense single-stranded RNA virus genomic sequences included in the orders Bunyavirales and Mononegavirales are reported. In addition, two positive single-strand virus genome sequences and one single strand DNA virus genome sequence were also identified. While the presence of a putative tenuivirus associated with S. invicta was previously suggested to be a contamination, here we characterized and present strong evidence that Solenopsis invicta virus 14 (SINV-14) is a tenui-like virus that has a long-term association with the ant. Furthermore, based on virus sequence abundance compared to housekeeping genes, phylogenetic relationships, and completeness of viral coding sequences, our results suggest that four of five virus sequences reported, those being SINV-14, SINV-15, SINV-16 and SINV-17, may be associated to viruses actively replicating in the ant S. invicta. CONCLUSIONS: The present study expands our knowledge about viral diversity associated with S. invicta in introduced areas with potential to be used as biological control agents, which will require further biological characterization.

Gut bacteria are essential for normal cuticle development in herbivorous turtle ants.

Across the evolutionary history of insects, the shift from nitrogen-rich carnivore/omnivore diets to nitrogen-poor herbivorous diets was made possible through symbiosis with microbes. The herbivorous turtle ants Cephalotes possess a conserved gut microbiome which enriches the nutrient composition by recycling nitrogen-rich metabolic waste to increase the production of amino acids. This enrichment is assumed to benefit the host, but we do not know to what extent. To gain insights into nitrogen assimilation in the ant cuticle we use gut bacterial manipulation, 15N isotopic enrichment, isotope-ratio mass spectrometry, and 15N nuclear magnetic resonance spectroscopy to demonstrate that gut bacteria contribute to the formation of proteins, catecholamine cross-linkers, and chitin in the cuticle. This study identifies the cuticular components which are nitrogen-enriched by gut bacteria, highlighting the role of symbionts in insect evolution, and provides a framework for understanding the nitrogen flow from nutrients through bacteria into the insect cuticle.

Toxicity and Sublethal Effects of Autumn Crocus (Colchicum autumnale) Bulb Powder on Red Imported Fire Ants (Solenopsis invicta).

Autumn crocus (Colchicum autumnale L.) is a medicinal plant as it contains high concentrations of colchicine. In this study, we reported that the ground powder of autumn crocus bulb is highly toxic to invasive Solenopsis invicta Buren, commonly referred to as red imported fire ants (RIFAs). Ants fed with sugar water containing 5000 mg/L of bulb powder showed 54.67% mortality in three days compared to 45.33% mortality when fed with sugar water containing 50 mg/L of colchicine. Additionally, the effects of short-term feeding with sugar water containing 1 mg/L of colchicine and 100 mg/L of autumn crocus bulb powder were evaluated for RIFAs' colony weight, food consumption, and aggressiveness, i.e., aggregation, grasping ability, and walking speed. After 15 days of feeding, the cumulative colony weight loss reached 44.63% and 58.73% due to the sublethal concentrations of colchicine and autumn crocus bulb powder, respectively. The consumption of sugar water and mealworm (Tenebrio molitor L.) was substantially reduced. The aggregation rates decreased 48.67% and 34.67%, grasping rates were reduced to 38.67% and 16.67%, and walking speed decreased 1.13 cm/s and 0.67 cm/s as a result of the feeding of the two sublethal concentrations of colchicine and autumn crocus bulb powder, respectively. Our results for the first time show that powder derived from autumn crocus bulbs could potentially be a botanical pesticide for controlling RIFAs, and application of such a product could be ecologically benign due to its rapid biodegradation in the environment.

Maternal effect killing by a supergene controlling ant social organization.

Supergenes underlie striking polymorphisms in nature, yet the evolutionary mechanisms by which they arise and persist remain enigmatic. These clusters of linked loci can spread in populations because they captured coadapted alleles or by selfishly distorting the laws of Mendelian inheritance. Here, we show that the supergene haplotype associated with multiple-queen colonies in Alpine silver ants is a maternal effect killer. All eggs from heterozygous queens failed to hatch when they did not inherit this haplotype. Hence, the haplotype specific to multiple-queen colonies is a selfish genetic element that enhances its own transmission by causing developmental arrest of progeny that do not carry it. At the population level, such transmission ratio distortion favors the spread of multiple-queen colonies, to the detriment of the alternative haplotype associated with single-queen colonies. Hence, selfish gene drive by one haplotype will impact the evolutionary dynamics of alternative forms of colony social organization. This killer hidden in a social supergene shows that large nonrecombining genomic regions are prone to cause multifarious effects across levels of biological organization.

Biological aspects of Myopopone castanea on it's prey Oryctes rhinoceros larvae.

Ants are social insects with some significant roles in the ecosystem, including acting as predators for various insect pests. Myopopone castanea ants is a predatorfor the larvae of Oryctes rhinoceros pest. The existence of a similar niche of life between M. castanea ants and O. rhinoceros larvae opens an excellent opportunity to utilize these ants as biological agents. The research was conducted to study some aspects biology of M. castanea so that later it can be applied to mass rearing of natural enemies in the laboratory. The study was conducted by maintaining 50 eggs of M. castanea ant. Then, the eggs placed on two pieces of decayed palm oil stem together with twenty individual worker ants and ten individual end instar larvae. It needs five replications for the experiment. The results showed that egg stadia length was 13.8 days. It found five instars within M. castanea ant larvae with varying lengths of each stage. It takes 17.2 days for worker ant pupae to go through stadia pupa and 17.9 days for female ant pupae. The survival rate of M. castanea ant life from eggs until imago is 56.4%, which means that from several groups of eggs laid by queen ants, only about half have succeeded in becoming ant imago.

Comparative Development of the Ant Chemosensory System.

The insect antennal lobe (AL) contains the first synapses of the olfactory system, where olfactory sensory neurons (OSNs) contact second-order projection neurons (PNs). In Drosophila melanogaster, OSNs expressing specific receptor genes send stereotyped projections to one or two of about 50 morphologically defined glomeruli [1-3]. The mechanisms for this precise matching between OSNs and PNs have been studied extensively in D. melanogaster, where development is deterministic and independent of neural activity [4-6]. However, a number of insect lineages, most notably the ants, have receptor gene repertoires many times larger than D. melanogaster and exhibit more structurally complex antennal lobes [7-12]. Moreover, perturbation of OSN function via knockout of the odorant receptor (OR) co-receptor, Orco, results in drastic AL reductions in ants [13, 14], but not in Drosophila [15]. Here, we characterize AL development in the clonal raider ant, Ooceraea biroi. We find that, unlike in Drosophila, ORs and Orco are expressed before the onset of glomerulus formation, and Orco protein is trafficked to developing axon terminals, raising the possibility that ORs play a role during ant AL development. Additionally, ablating ant antennae at the onset of pupation results in AL defects that recapitulate the Orco mutant phenotype. Thus, early loss of functional OSN innervation reveals latent structure in the AL that develops independently of peripheral input, suggesting that the AL is initially pre-patterned and later refined in an OSN-dependent manner. This two-step process might increase developmental flexibility and thereby facilitate the rapid evolution and expansion of the ant chemosensory system.

Long-lived Temnothorax ant queens switch from investment in immunity to antioxidant production with age.

Senescence is manifested by an increase in molecular damage and a deterioration of biological functions with age. In most organisms, body maintenance is traded-off with reproduction. This negative relationship between longevity and fecundity is also evident on the molecular level. Exempt from this negative trait association, social insect queens are both extremely long-lived and highly fecund. Here, we study changes in gene expression with age and fecundity in ant queens to understand the molecular basis of their long lifespan. We analyse tissue-specific gene expression in young founding queens and old fecund queens of the ant Temnothorax rugatulus. More genes altered their expression with age in the fat body than in the brain. Despite strong differences in ovary development, few fecundity genes were differentially expressed. Young founding queens invested in immunity (i.e. activation of Toll signalling pathway) and resistance against environmental and physiological stress (i.e. down-regulation of TOR pathway). Conversely, established older queens invested into anti-aging mechanisms through an overproduction of antioxidants (i.e. upregulation of catalase, superoxide dismutase). Finally, we identified candidate genes and pathways, potentially involved in the association between fertility and longevity in social insects and its proximate basis.

Evolution of hybridogenetic lineages in Cataglyphis ants.

In most social Hymenoptera, a diploid egg develops into either a queen or a worker depending on environmental conditions. Hybridogenetic Cataglyphis ants display a bizarre genetic system, where queen-worker caste determination is primarily determined by genetic factors. In hybridogenetic populations, all workers are F1 hybrids of two distinct lineages, whereas new queens are nearly always pure-lineage individuals produced by clonal reproduction. The distribution and evolutionary history of these hybridogenetic populations have not yet been thoroughly analysed. Here, we studied the phylogeographic distribution of hybridogenetic populations in two closely related Spanish species: Cataglyphis humeya and Cataglyphis velox. Hybridogenesis has been previously documented in a locality of C. velox, but whether this system occurs elsewhere within the range of the two species was yet unknown. Queens and workers from 66 localities sampled across the range of the species were genotyped at 18 microsatellite markers to determine whether queens were produced by parthenogenesis and whether workers were hybrids of divergent lineages. Populations with F1 hybrid workers were identified by combining genetic, geographical and mating assortments data. In most populations of C. velox, workers were found to be hybrids of two divergent lineages. Workers were however produced via random mating in two marginal populations of C. velox, and in all populations studied of its sister species C. humeya. High-throughput sequencing data were obtained to confirm inferences based on microsatellites and to characterize relationships between populations. Our results revealed a complicated history of reticulate evolution that may account for the origin of hybridogenetic lineages in Cataglyphis.

The possible role of ant larvae in the defence against social parasites.

Temporary social parasite ant queens initiate new colonies by entering colonies of host species, where they begin laying eggs. As the resident queen can be killed during this process, host colonies may lose their entire future reproductive output. Selection thus favours the evolution of defence mechanisms, before and after parasite intrusion. Most studies on social parasites focus on host worker discrimination of parasite queens and their offspring. However, ant larvae can also influence brood composition by consuming eggs. This raises the question whether host larvae can aid in preventing colony takeover by consuming eggs laid by parasite queens. To test whether larvae could play a role in anti-parasite defence, we compared the rates at which larvae of a common host species, Formica fusca, consumed eggs laid by social parasite, non-parasite, nest-mate, or conspecific non-nest-mate queens. Larvae consumed social parasite eggs more than eggs laid by a heterospecific non-parasite queen, irrespective of the chemical distance between the egg cuticular profiles. Also, larvae consumed eggs laid by conspecific non-nest-mate queens more than those laid by nest-mate queens. Our study suggests that larvae may act as players in colony defence against social parasitism, and that social parasitism is a key factor shaping discrimination behaviour in ants.

Changes in gene DNA methylation and expression networks accompany caste specialization and age-related physiological changes in a social insect.

Social insects provide systems for studying epigenetic regulation of phenotypes, particularly with respect to differentiation of reproductive and worker castes, which typically arise from a common genetic background. The role of gene expression in caste specialization has been extensively studied, but the role of DNA methylation remains controversial. Here, we perform well replicated, integrated analyses of DNA methylation and gene expression in brains of an ant (Formica exsecta) with distinct female castes using traditional approaches (tests of differential methylation) combined with a novel approach (analysis of co-expression and co-methylation networks). We found differences in expression and methylation profiles between workers and queens at different life stages, as well as some overlap between DNA methylation and expression at the functional level. Large portions of the transcriptome and methylome are organized into "modules" of genes, some significantly associated with phenotypic traits of castes and developmental stages. Several gene co-expression modules are preserved in co-methylation networks, consistent with possible regulation of caste-specific gene expression by DNA methylation. Surprisingly, brain co-expression modules were highly preserved when compared with a previous study that examined whole-body co-expression patterns in 16 ant species, suggesting that these modules are evolutionarily conserved and for specific functions in various tissues. Altogether, these results suggest that DNA methylation participates in regulation of caste specialization and age-related physiological changes in social insects.