How to coexist with fire ants: The roles of behaviour and cuticular compounds

tBecause territoriality is energetically costly, territorial animals frequently respond less aggressively toneighbours than to strangers, a reaction known as the “dear enemy phenomenon” (DEP). The contrary,the “nasty neighbour effect” (NNE), occurs mainly for group-living species defending resource-basedterritories. We studied the relationships between supercolonies of the pest fire ant Solenopsis saevissimaand eight ant species able to live in the vicinity of its nests plus Eciton burchellii, an army ant predatorof other ants. The workers from all of the eight ant species behaved submissively when confrontedwith S. saevissima (dominant) individuals, whereas the contrary was never true. Yet, S. saevissima weresubmissive towards E. burchellii workers. Both DEP and NNE were observed for the eight ant species, withsubmissive behaviours less frequent in the case of DEP. To distinguish what is due to chemical cues fromwhat can be attributed to behaviour, we extracted cuticular compounds from all of the nine ant speciescompared and transferred them onto a number of S. saevissima workers that were then confronted withuntreated conspecifics. The cuticular compounds from three species, particularly E. burchellii, triggeredgreater aggressiveness by S. saevissima workers, while those from the other species did not

Mechanisms for the Evolution of Superorganismality in Ants

Ant colonies appear to behave as superorganisms; they exhibit very high levels of within-colony cooperation, and very low levels of within-colony conflict. The evolution of such superorganismality has occurred multiple times across the animal phylogeny, and indeed, origins of multicellularity represent the same evolutionary process. Understanding the origin and elaboration of superorganismality is a major focus of research in evolutionary biology. Although much is known about the ultimate factors that permit the evolution and persistence of superorganisms, we know relatively little about how they evolve. One limiting factor to the study of superorganismality is the difficulty of conducting manipulative experiments in social insect colonies. Recent work on establishing the clonal raider ant, Ooceraea biroi, as a tractable laboratory model, has helped alleviate this difficulty. In this dissertation, I study the proximate evolution of superorganismality in ants. Using focussed mechanistic experiments in O. biroi, in combination with comparative work from other ant species, I study three major aspects of ant social behaviour that provide insight into the origin, maintenance, and elaboration of superorganismality. First, I ask how ants evolved to live in colonies, and how they evolved a reproductive division of labour. A comparative transcriptomic screen across the ant phylogeny, combined with experimental manipulations in O. biroi, finds that reproductive ants have higher insulin levels than their non-reproductive nestmates, and that this likely regulates the reproductive division of labour. Using these data, as well as studies of the idiosyncrasies of O. biroi’s life history, I propose a mechanism for the evolution of the first colonies. It is possible that similar mechanisms underlie the evolution of reproductive division of labour in other superorganisms, and of germ-soma separation in nascent multicellular individuals. Second, I ask how ant workers assess colony hunger to regulate their foraging behaviour. I find that workers use larval signals, but not their own nutritional states, to decide how much to forage. In contrast, they use their nutritional states, but not larval signals, to decide how much to eat, suggesting that in at least some ant species, foraging and feeding have been decoupled. This evolution of colony-level foraging regulation has occurred convergently in hymenopteran superorganisms, and is analogous to the evolution of centralised regulation of foraging behaviour in multicellular animals. Finally, I ask how an iconic collective foraging behaviour – the mass raids of army ants – evolved. I find that O. biroi, a relative of army ants, forages collectively in group raids, that these are ancestral to the mass raids of army ants, and that the transition from group to mass raiding correlates with expansion in colony size. I propose that the scaling effects of increasing colony size explain this transition. It is possible that similar principles underlie the evolution of disparate collective behaviours in other animal groups and among cells within developing animals. Together, these studies illuminate the life history of O. biroi, and suggest mechanisms for the evolution of core aspects of cooperative behaviour in ant colonies. I draw comparisons to the evolution of superorganismality in other lineages, as well as to the evolution of multicellularity. I suggest that there may be additional similarities in the proximate evolutionary trajectories of superorganismality and multicellularity

Ecological consequences of colony structure in dynamic ant nest networks

Access to resources depends on an individual’s position within the environment. This is particularly important to animals that invest heavily in nest construction, such as social insects. Many ant species have a polydomous nesting strategy: a single colony inhabits several spatially separated nests, often exchanging resources between the nests. Different nests in a polydomous colony potentially have differential access to resources, but the ecological consequences of this are unclear. In this study, we investigate how nest survival and budding in polydomous wood ant (Formica lugubris) colonies are affected by being part of a multi-nest system. Using field data and novel analytical approaches combining survival models with dynamic network analysis, we show that the survival and budding of nests within a polydomous colony is affected by their position in the nest-network structure. Specifically, we find that the flow of resources through a nest, which is based on its position within the wider nest-network, determines a nest’s likelihood of surviving, and of founding new nests. Our results highlight how apparently disparate entities in a biological system can be integrated into a functional ecological unit. We also demonstrate how position within a dynamic network structure can have important ecological consequences

The evolution of social traits and biodiversity in the ants.

Cooperation has shaped the evolution of life on Earth. The ants are the most numerically diverse of the eusocial Hymenoptera, and display wide variation in social complexity. This positions the ants as an ideal taxon in which to study social evolution in a comparative framework. Social evolution theory has generated many hypotheses that are testable in ants, however the lack of comprehensive or complete phylogenies, and the decentralised and scattered nature of trait data, has been an obstacle to these types of study. In this thesis I construct a large species-level, and a complete genus-level, phylogeny of the ants, and draw together a large dataset of social traits from the literature in order to test hypotheses concerning the evolution of social traits in the ants. I find evidence that the earliest ant was large bodied, and lived in small highly related colonies. I show that group size is a significant trait in the evolution of sociality in ants, predicting the probability of a species having polymorphic workers, or of being polyandrous. I also show that the change in these traits is correlated between ancestral nodes on the phylogeny. Furthermore, in the Attini, colony size correlates closely with non-reproductive and reproductive division of labour. Together these results cement group size as a driving force of social evolution in the ants, and this has interesting implications for social evolution in general. Finally, I report the first evidence that intermediate colony sizes, the presence of discrete worker castes and polygyny are associated with increased diversification rates in ants. This thesis provides a valuable tool for the study of comparative hypotheses in the ants in the form of a complete genus-level phylogeny, and offers significant evidence to support several key hypotheses in social evolution. Furthermore, these results generate hypotheses regarding the evolution of social traits for future research

The curious case of the camelthorn: competition, coexistence and nest-site limitation in a multispecies mutualism

Myrmecophyte plants house ants in domatia in exchange for protection from herbivores. Ant-myrmecophyte mutualisms exhibit two general patterns due to competition between ants for plant occupancy: i) domatia nest-sites are a limiting resource and ii) each individual plant hosts one ant species at a time. However, individual camelthorn trees (Vachellia erioloba) typically host two to four ant species simultaneously, often coexisting in adjacent domatia on the same branch. Such fine-grain spatial coexistence brings into question the conventional wisdom on ant-myrmecophyte mutualisms. Camelthorn ants appear not to be nest-site limited, despite low abundance of suitable domatia, and have random distributions of nest-sites within and across trees. These patterns suggest a lack of competition between ants for domatia and contrast strongly with other ant-myrmecophyte systems. Comparison of this unusual case with others suggests that spatial scale is crucial to coexistence or competitive exclusion involving multiple ant species. Furthermore, coexistence may be facilitated when co-occurring ant species diverge strongly on at least one niche axis. Our conclusions provide recommendations for future ant-myrmecophyte research, particularly in utilising multispecies systems to further our understanding of mutualism biology

Contextual organismality: Beyond pattern to process in the emergence of organisms

Biologists have taken the concept of organism largely for granted. However, advances in the study of chimerism, symbiosis, bacterial-eukaryote associations, and microbial behavior have prompted a redefinition of organisms as biological entities exhibiting low conflict and high cooperation among their parts. This expanded view identifies organisms in evolutionary time. However, the ecological processes, mechanisms, and traits that drive the formation of organisms remain poorly understood. Recognizing that organismality can be context dependent, we advocate elucidating the ecological contexts under which entities do or do not act as organisms. Here we develop a "contextual organismality" framework and provide examples of entities, such as honey bee colonies, tumors, and bacterial swarms, that can act as organisms under specific life history, resource, or other ecological circumstances. We suggest that context dependence may be a stepping stone to the development of increased organismal unification, as the most integrated biological entities generally show little context dependence. Recognizing that organismality is contextual can identify common patterns and testable hypotheses across different entities. The contextual organismality framework can illuminate timeless as well as pressing issues in biology, including topics as disparate as cancer emergence, genomic conflict, evolution of symbiosis, and the role of the microbiota in impacting host phenotype.John Templeton FoundationVersion of record online: 27 October 2016; published open access.This item from the UA Faculty Publications collection is made available by the University of Arizona with support from the University of Arizona Libraries. If you have questions, please contact us at [email protected]

Impact of the invasive argentine ant in citrus agroecosystems: effects on the diversity and frequency of native ant species foraging on tree canopy

The invasion of the Argentine ant, Linepithema humile (Mayr) (Hymenoptera, Formicidae) can alter the entire ecosystem with serious impacts on the native community structure (e.g., ant diversity) and processes (e.g., trophic interactions) leading to biodiversity loss and pest outbreaks. Most studies addressing these impacts have been conducted in natural or semi-natural areas, few are those conducted in agricultural ecosystems, such as citrus orchards. These are dominant agricultural ecosystems in Mediterranean landscapes. Furthermore, most studies have been conducted in a short span, not evidencing seasonal fluctuations. In this work, we assessed the ecological impact of the Argentine ant on the native ant communities in citrus orchards, in the region of Algarve, southern Portugal. By using principal response curve, we compared seasonal variation on ant assemblages in invaded and uninvaded citrus orchards foraging on tree canopy from a two-year sampling. The Argentine ant had a marked negative impact on the native ant community foraging on citrus canopy. In the uninvaded orchards, the native ant community had a rich assemblage composed of 16 ant species, in its majority (72%) controlled by the dominant species Lasius grandis Forel, Tapinoma nigerrimum (Nylander) and/or Pheidole pallidula (Nylander). In the invaded orchards, the native ant community was poorer and highly modified, mostly dominated by the Argentine ant (80%). Apparently, the only native ant species not a ected by the presence of the Argentine ant was Plagiolepis pygmaea (Latreille). A significant negative e ect was found between the proportion of infested trees by L. humile and the number of native ant species per orchard. Di erences in the native ant community in the invaded and uninvaded orchards persisted over seasons and years. However, negative impacts were higher in the spring and summer, and less pronounced in the autumn. We discuss implications for citrus pest managementinfo:eu-repo/semantics/publishedVersio

Biological Metaphors for Whiteness: Beyond Merit and Malice

The problem of persistent racial inequality is grounded in a failure of imagination. The general mainstream conception is that unfair racial inequality occurs only when there is intentional racism. Absent conscious racial malice, no racism is seen to exist. The only generally available alternative explanation for racial inequality is the meritocratic system. Viewing the distribution of resources as a product of a generally fair meritocratic system provides a defense against any charge of racism, and justifies the status quo. But in economics, business, computer science, and even biology, observers of complexity are coming to understand how dominant systems can prevail without superior merit, can maintain their position without any conscious guidance or intent, and can be organized without any collusion or direction. Markets, organisms, and ecologies coordinate themselves efficiently and organically, with surprising resilience. Whiteness operates like these other systems. This essay re-imagines Whiteness using images from perhaps unusual sources. Whiteness coalesces through the actions of multitudes of independent individuals, in the same way that slime mold forms when spore cells join together on the forest floor. Racial segregation results from simple self-organizing mathematical algorithms realized in the collective behavior of human beings moving in and out of neighborhoods. Whiteness sustains itself in the same way that cultural practices and self-serving beliefs do, without conscious intent. Whiteness carefully organizes itself in the same way that snowflakes and ants do, without anyone being in charge or giving direction

A Bioeconomic Model of Little Fire Ant Wasmannia auropunctata in Hawaii

Reports were scanned in black and white at a resolution of 600 dots per inch and were converted to text using Adobe Paper Capture Plug-in.Wasmannia auropunctata, known as the Little Fire Ant (LFA), was first detected on the island of Hawai‘i (the Big Island) in 1999. It was most probably introduced through imports of contaminated potted plants from mainland USA. We estimate that LFA has now spread to over 4,000 locations on the Big Island and under current management efforts will spread rapidly inundating the Big Island in 15-20 years. Increased efforts in prevention, detections, and mitigation treatments will suppress existing infestations, reduce rate of spread and decrease long term management costs, damages, and human stings. Benefits from increased management are estimated to be $5 billion savings including$540 million in reduced damages and 2.1 billion fewer sting incidents over 35 years.This research was supported in part by the Tropical and Subtropical Agriculture Research (TSTAR) Program (Award Number 2010-34135-21228), The National institute of Food and Agriculture (NIFA), US Department of Agriculture (USDA