An experimental test of Lanchester's models of combat in the neotropical termite Nasutitermes corniger (Blattodea: Termitidae).

Lanchester's models of combat have been invoked to explain the mechanics of group fighting in social animals. Specifically, Lanchester's square law posits that the fighting ability of the group is proportional to the square of the number of combatants. Although used to explain a variety of ecological phenomena, the models have not been thoroughly tested. We tested the Lanchester models using group battles between colonies of the termite Nasutitermes corniger. Our main goals were to determine if mortality rates fit the Lanchester models, and if so, whether the behavioural mechanisms underlying a group's success match those used in deriving the model. We initiated battles between pairs of colonies with different ratios of fighters and recorded deaths over time. We found that the numerically larger army has an advantage, but that the advantage is not as pronounced as predicted by Lanchester's square law. We also video-recorded battles to analyse individual behaviour, which did not support the mechanisms invoked by Lanchester. Instead, the killing power of an individual is increased by the presence of nest-mates, giving the larger group a disproportionate advantage. Although the behavioural mechanisms leading to the advantage may differ, our results still support some of the proposed ecological phenomena.

Genetic Analysis of Invasive Conehead Termites (Blattodea: Termitidae) Reveals a Single Origin for Two Populations in Florida.

In 2001, Nasutitermes corniger (Motschulsky), common name conehead termite, were discovered near a marina in Dania Beach, FL, where the invasive species was probably transported from its native range in Central and South America or the Caribbean. In January 2016, an infestation was found in Pompano Beach, Florida, approximately 21 km north of the Dania Beach population. This study compares variants in seven microsatellite loci across specimens from 11 nests in Dania Beach and 8 nests in Pompano Beach. Results are consistent with all N. corniger in both locations being descendants of a single introduced colony, spreading within Broward County, FL through human transport of infested materials. No more than four alleles were found at any of the seven microsatellite loci analyzed, inferring that a single Queen and King, or multiple sibling reproductives descended from a monogamous pair, headed the colony that arrived in Florida. The potential economic and environmental impacts of this invasive termite are enormous due to its broad diet, including agricultural crops and orchards, native and ornamental plants, natural landscapes, and structures. Conspicuous tunnels and aboveground nests are the key aspects of N. corniger biology that render colonies vulnerable to discovery and control. The now proven ability of N. corniger to establish breeding populations in the United States, to cause extensive property and landscape destruction, and to spread by human transport underscores the need for continued aggressive efforts toward eradication of known infestations as well as quick operational actions the next time invasive N. corniger are discovered.

Self-organizing conflicts: Group assessment and the spatio-temporal dynamics of ant territory battles.

Territorial battles among ants exhibit temporal and spatial patterns that self-organize, arising spontaneously from distributed decisions by large numbers of individuals. We describe agent-based models of inter-group fights in ants and show that two behavioral mechanisms that are rarely quantified have large effects on the dynamics of intraspecific battles; specifically, the pattern of search by unengaged ants, and assessment of relative numbers. In the absence of assessment, recruitment by both colonies rises to steady averages. Alternatively, if ants tend to lay trails only when they detect that their nestmates outnumber opponents, fights can be rapidly resolved as one colony ceases recruiting. If ants tend to lay trails when their nestmates are locally outnumbered, the position of the battle may oscillate. We show that the collective ability of fighting ants to accurately compare group sizes may be high even if each ant has limited perception and memory. However, amplification of small initial numerical advantages can lead to priority effects favoring the first colony to recruit even if it is the smaller colony.

Mechanistic Models of Conflict between Ant Colonies and Their Consequences for Territory Scaling.

Territory size in social insects depends on the rules by which border conflicts are resolved. We present three mechanistic mathematical models of conflict, inspired by the behavior of the pavement ant Tetramorium immigrans, to predict the advantage of larger colonies in pairwise contests and the resulting scaling of territory size with worker force. The models track the number of ants in the nest traveling to and from the boundary or engaged at the boundary. Ants at the boundary base their recruitment response on the relative numbers of ants from the two colonies. With two colonies, our central result is that the larger colony gains a territory disproportionately larger than the ratio of worker forces would indicate. This disproportionate territory control determines the scaling relation of territory size with worker force in a population. In two dimensions, if territory size were proportional to worker force, the slope of the scaling relation between log territory size and log worker force would be 1.0. With disproportionate territories, this slope is larger and can be explicitly approximated in terms of model parameters, and it is steepest when colonies are packed close to each other, when ants run quickly, or when colonies are small. A steeper slope exaggerates the advantage of larger colonies, creating a positive feedback that could amplify the inequality of the worker force distribution.

The Distribution and Habitat Affinities of the Invasive Ant Myrmica rubra (Hymenoptera: Formicidae) in Southern New England.

The Eurasian ant Myrmica rubra (L.) (Hymenoptera: Formicidae) was first discovered in North America in the early 1900s in Massachusetts. Populations have since appeared in at least seven states within the United States and in seven Canadian provinces. We conducted a systematic search for the ant across southern New England-the states of Connecticut, Massachusetts, and Rhode Island-where M. rubra is spreading from multiple loci. The species occurs in two large regions in Massachusetts, each spanning approximately 75 km, and in several smaller populations in Massachusetts and Rhode Island. No populations were discovered anywhere in Connecticut or across large expanses of central Massachusetts and northern Rhode Island, despite the presence of apparently favorable habitat. This pattern of distribution suggests a combination of long-distance dispersal by human transport coupled with slow local spread. Resurveys of sites previously known to support M. rubra showed that populations persist for decades. Within invaded areas, M. rubra was strongly associated with particular habitats. Colonies were most prevalent in freshwater wetlands and in moist forests near wetlands and water; they were uncommon in drier forests and were rare in open habitats outside of wetlands. The slow rate of spread over the last 110 yr suggests that the ants do not easily disperse between patches of suitable habitat.

An empirical test of Lanchester's square law: mortality during battles of the fire ant Solenopsis invicta.

Lanchester's models of attrition describe casualty rates during battles between groups as functions of the numbers of individuals and their fighting abilities. Originally developed to describe human warfare, Lanchester's square law has been hypothesized to apply broadly to social animals as well, with important consequences for their aggressive behaviour and social structure. According to the square law, the fighting ability of a group is proportional to the square of the number of individuals, but rises only linearly with fighting ability of individuals within the group. By analyzing mortality rates of fire ants (Solenopsis invicta) fighting in different numerical ratios, we provide the first quantitative test of Lanchester's model for a non-human animal. Casualty rates of fire ants were not consistent with the square law; instead, group fighting ability was an approximately linear function of group size. This implies that the relative numbers of casualties incurred by two fighting groups are not strongly affected by relative group sizes and that battles do not disproportionately favour group size over individual prowess.

Landmarks in territory partitioning: a strategically stable convention?

A convention is a rule based on arbitrary cues that allows quick resolution of potentially protracted disputes. A familiar example is the Bourgeois strategy, in which the second of two animals to discover a resource yields it to the first, even though it may be stronger than its opponent. Here we develop a game-theoretic model to show that neighbors with imperfect information about one another's fighting abilities can be favored to accept a landmark as the designator of a territory boundary, even when the resulting territory is smaller than the one that would have been won through fighting. Thus, the use of landmarks or other mutually obvious solutions can serve as a convention for territory partitioning. For a distribution of fighting ability with low variance and high skew, there is a remarkably high probability that an animal will accept a smaller territory than it would have won through fighting. The analysis provides a possible explanation for the observed use of landmarks as boundary markers by territorial animals in a variety of taxa, including birds, fish, insects, and mammals. The analysis also suggests why territory boundaries are stable, once established, despite changes in characteristics of the residents or the environment.

Territory defense by the ant Azteca trigona: maintenance of an arboreal ant mosaic.

Mosaics of exclusive foraging territories, produced by intra-and interspecific competition, are commonly reported from arboreal ant communities throughout the tropics and appear to represent a recurring feature of community organization. This paper documents an ant mosaic within mangrove forests of Panama and examines the behavioral mechanisms by which one of the common species, Azteca trigona, maintains its territories. Most of the mangrove canopy is occupied by mutually exclusive territories of the ants A. trigona, A. velox, A. instabilis, and Crematogaster brevispinosa. When foraging workers of A. trigona detect workers of these territorial species, they organize an alarm recruitment response using pheromonal and tactile displays. Nestmates are attracted over short distances by an alarm pheromone originating in the pygidial gland and over longer distances by a trail pheromone produced by the Pavan's gland. Recruits are simultaneously alerted by a tactile display. No evidence was found for chemical marking of the territory. Major workers are proportionally more abundant at territory borders than on foraging trails in the interior of the colony. The mechanisms of territory defense in A. trigona are remarkably similar to those of ecologically analogous ants in the Old World tropics.

Chemical interference competition by Monomorium minimum (Hymenoptera: Formicidae).

Workers of Monomorium minimum forage above-ground for dead arthropods. Small particles (<1 mg) are retrieved individually, but larger particles stimulate recruitment and are dissected by groups of workers. The recruitment pheromone originates in the Dufour's gland and the number of ants responding to a trail varies with pheromone concentration. When ants of other species are encountered at food resources, workers of M. minimum gaster-flag and extrude an irritating poison gland secretion from the sting. This chemical interference delays invasion by competitors and prolongs the period during which the colony can dissect and retrieve pieces of the food resource. M. minimum recruits at higher temperatures than sympatric ant species. The probability of interference at food baits rises from 5% to 100% when they become too large for a single worker to carry. The probability of food resource loss is higher for baits of intermediate weight (x=18.1 mg) than for those of low weight (x=0.1 mg) or high weight (x=403.1 mg).