publications List

Anna Dornhaus’ original scientific papers

Publications by theme

Overview articles

Benefits and costs of communication

Mechanisms of communication

Division of labor

Cognition

Individual differences

Collective decisions

Spatial patterns

Collective contests

Various

ATTENTION: Website under construction. Note that publications are not sorted by year (you can find a chronological list at Google Scholar). All articles are peer-reviewed unless otherwise marked.

Overview articles

Dornhaus A 2014 ‘Finding food: foraging affects all aspects of an animal’s life’, in: Yasukawa K ‘Animal Behavior, Volume II: Function and Evolution of Animal Behavior’, Praeger Publishers - pdf - textbook chapter for a general audience on what animals do to find food

Dornhaus A, Powell S 2010 ‘Foraging and defence strategies’ In: ‘Ant Ecology’, Eds. L Lach, C Parr, K Abbott, Ant Ecology, Oxford University Press [invited chapter in edited book] - pdf - edited book chapter reviewing communication/recruitment and defense strategies in ants

Dornhaus A, Powell S, Bengston S 2012 ‘Group size and its effects on collective organization’, Annual Review of Entomology 57: 123-141 - pdf - quantitative review on group sizes in insects, and review on how group size does or does not affect collective behavior

Bengston S, Charbonneau D, Dornhaus A 2021, ‘Temnothorax’ entry for Encyclopedia of Social Insects by Starr (Ed.), Springer - pdf - encyclopedia entry about the ant genus ‘Temnothorax’

Dornhaus A, Franks NR 2008 'Individual and collective cognition in ants and other insects (Hymenoptera: Formicidae)' Myrmecol News 11: 215-226 - pdf - review of cognitive/information processing/decision-making skills in social insects; also see next entry and lectures on youtube on this topic

Dornhaus A, in press ‘Aliens Are Likely to Be Smart But Not “Intelligent”: What Evolution of Cognition on Earth Tells Us about Extraterrestrial Intelligence’, In: Vakoch D, editor. Extraterrestrial Intelligence: Cognition and Communication in the Universe. Oxford University Press - pdf - invited book chapter; discussion of how many cognitive skills are common among organisms on Earth, thus presumably ‘easy’ to evolve; but human-level self-reflection/language/altruism is unique, thus possibly ‘hard’ to evolve or not under natural selection at all

Benefits and costs of communication in groups

We often intuitively feel that more information is always better, and that organizations will work more efficiently if more information is shared, but this is often not true for a variety of reasons - as anyone working for a large organization should know. Information has surprising costs: not just producing it, but also opportunity costs of waiting for it, low signal-to-noise ratio if sharing is not selective, too-high uniformity across individuals, difficult/slow updating of information across the group, and other non-obvious consequences of information sharing that are negative for individuals and groups.

Bee Dance and maps

Honey bees (Apis) communicate the locations of food sources - but bumble bees do not. Why?
We demonstrated that benefits of communicating location are not actually common, and appear driven by rare, far away, but plentiful and diverse resources such as mass-flowering trees found in the tropics.

Dornhaus, A, Chittka, L 2004 ‘Why do honey bees dance?’, Behavioral Ecology and Sociobiology 55: 395-401 - pdf - field data in natural areas on increase in nectar collection with vs. without waggle dance location communication show benefit in tropics but not temperate areas

Dornhaus A, Klügl F, Oechslein C, Puppe F, Chittka L 2006 ‘Benefits of recruitment in honey bees: effects of ecology and colony size in an individual-based model’, Behavioral Ecology 17: 336-344 - pdf - simulation model shows strong dependence of waggle dance recruitment benefits on spatial distribution of resources

Dornhaus, A 2002 ‘Significance of honeybee recruitment strategies depending on foraging distance (Hymenoptera : Apidae : Apis mellifera)’, Entomologia Generalis 26: 93-100 - field data on artificial food sources show bees can find nearby food sources in response to recruitment even without waggle dance location information

Donaldson-Matasci MC, Dornhaus A 2012 ‘How habitat affects the benefits of communication in collectively foraging honey bees’, Behavioral Ecology and Sociobiology 66: 583-592 - pdf - field data on a variety of natural habitats show differences in dance benefits best explained by resource diversity

Raine, N E, Ings, T C, Dornhaus, A, Saleh, N, Chittka, L 2006 ‘Adaptation, genetic drift, pleiotropy, and history in the evolution of bee foraging behavior’, Advances in the Study of Behavior 36: 305-354 - pdf - review article on evolution of bee foraging behavior

Bee dance and time

We demonstrated that a variety of (opportunity) costs can arise just from the expectation of communication, and that speed and time delays can drive benefits. Contrary to some expecations, large bee colonies capitalize mostly on having a lot of scouts, not on having a lot of recruits.

Donaldson-Matasci MC, DeGrandi-Hoffman G, Dornhaus A 2013 ‘Bigger is better: honeybee colonies as distributed information-gathering systems’, Animal Behaviour 85: 585-592 - pdf - larger colonies benefit more from location communication not because they recruit more bees to the same food sources, but because they have more scouts to find food sources earlier

Dechaume-Moncharmont, F-X, Dornhaus, A, Houston, A I, McNamara, J M, Collins, E J, Franks, N R 2005 ‘The hidden cost of information in collective foraging’, Proceedings of the Royal Society: Biological Sciences 272: 1689-1695 - pdf - remaining in the nest to be available to be recruited is a cost of communication; longer availability of resource increases benefits of recruitment

Dornhaus A, Collins EJ, Dechaume-Moncharmont F-X, Houston A, Franks NR, McNamara J 2006 ‘Paying for information: partial loads in central place foragers’, Behavioral Ecology and Sociobiology 61: 151-161 - pdf - foragers may even benefit from returning prematurely from rewarding food sources in order to not miss important new information at the hive

Uniformity vs creativity

Generally communication is expected to reduce exploration and increase across-individual consistency, but we find that this depends on the details of behavioral rules.

Donaldson-Matasci M, Dornhaus A 2014 ‘Dance Communication Affects Consistency, but Not Breadth, of Resource Use in Pollen-Foraging Honey Bees’, PLOS One 9: e107527 - analyzing pollen loads from honey bees shows that communicating did not reduce the diversity of pollen collected, but did increase consistency of what was collected across foragers

Lanan M, Dornhaus A, Jones EI, Waser A, Bronstein JL 2012 ‘The trail less traveled: individual decision-making and its effect on group behavior’, PLoS One 7: e47976 - a simulation model of ants foraging on pheromone trails demonstrates that symmetry breaking, and thus consistency of individual resource choices, depends on how non-linear the choice algorithm is, i.e. on details of how choice probability relates to pheromone concentration; larger colonies may have more or less consistent foragers than small colonies depending on these mechanistic details

mechanisms of communication in social insects

A lot of information-sharing in social insects is indirect; it may involve ‘blackboard architecture’ (depositing information in a central place) and/or cues (receivers extract information from behavior of knowledgeable individuals without explicit/evolved signals). Different species communicate different types of information (e.g. bumble bees communicate less information than honey bees).

Bumble bee dance

We demonstrated that successful bumble bee foragers returning to the nest do 3 things that communicate the presence and scent of food sources: deposit food in pots; run around like crazy (=’dance’); and produce a volatile pheromone from tergal glands.

Dornhaus, A., Chittka, L., 1999, ‘Evolutionary origins of bee dances’, Nature 401: 38 - pdf - tweet - first demonstration of ‘dance’ and scent learning based on honeypots in bumble bees (B. terrestris; no info about resource location communicated

Dornhaus, A., Chittka, L., 2001, ‘Food alert in bumblebees (Bombus terrestris): possible mechanisms and evolutionary implications’, Behavioral Ecology and Sociobiology 50: 570-576 - pdf - first demonstration that volatile pheromone involved, and that influx in honeypots is sufficient to achieve recruitment

Dornhaus, A, Brockmann, A, Chittka, L 2003 ‘Bumble bees alert to food with pheromone from tergal gland’, Journal of Comparative Physiology A 189: 47-51 - pdf - volatile recruitment pheromone emitted during bumble bee ‘dance’ is produced in small glands on tergites

Dornhaus, A, Chittka, L 2004 ‘Information flow and regulation of foraging activity in bumble bees’, Apidologie 35: 183-192 - pdf - short review on bumble bee recruitment system

Dornhaus, A, Chittka, L 2005 ‘Bumble bees (Bombus terrestris) store both food and information in honeypots’, Behavioral Ecology 16: 661-666 - pdf - potential foragers monitor both amount and concentration of nectar in honeypots; activation in response to influx is quality-dependent and affected by overall nectar stored, thus serving as ‘information center’

Granero, A M, Guerra Sanz, J M, Ega Gonzalez, F J, Martinez Vidal, J L, Dornhaus, A, Ghani, J, Serrano, A R, Chittka, L 2005 ‘Chemical compounds of the foraging recruitment pheromone in bumblebees’, Naturwissenschaften 92: 371-374 - pdf - recruitment pheromone found to consist of eucalyptol, ocimene and farnesol, with eucalyptol most active

Dornhaus, A, Cameron, S 2003 ‘A scientific note on food alert in Bombus transversalis’, Apidologie 34: 87-88 - pdf - B. transversalis appears to use the same recruitment strategies as B. terrestris

Chittka, L, Dornhaus, A 1999 ‘Comparisons in physiology and evolution, and why bees can do the things they do’, Ciencia al Dia International 2: No 2 - essay in the importance of considering phylogeny and interindividual variation in studying and explaining behavioral variation among species

Honey bees

Honey bees may in fact also use a volatile pheromone to alert other bees to waggle dances or to communicate the presence of successful foragers. And we did not find that giving-up-time was predicted by either nectar volume or waiting time.

Thom C, Dornhaus A 2007 ‘Preliminary report on the use of volatile compounds by foraging honey bees in the hive (Hymenoptera: Apidae: Apis)’ Entomologia generalis 29: 299-304 - pdf - honey bees appear to use a volatile pheromone to activate other foragers similar to bumble bees

Rivera M, Donaldson-Matasci M, Dornhaus A 2015 ‘Quitting time: When do honey bee foragers decide to stop foraging on natural resources?’, Frontiers in Ecology and Evolution 3: 50 - pdf - we do not find that either delay to or duration of trophallaxis predict when foragers give up on resources; contrary to claims that amount of nectar found on a foraging trip and social need for nectar, thought to correlate with trophallaxis duration and delay respectively, should determine this

Ants

Temnothorax ants, similar to bumble bees, used to be thought of as almost solitary foragers; we demonstrated that they can in fact make use of pheromone trails/marks left by others.

Cao TT, Dornhaus A 2012 ‘Ants use pheromone markings in emigrations to move closer to food-rich areas’, Insectes sociaux 59: 87-92 - pdf - Temnorthorax rugatulus ants are not entirely individualistic foragers, but instead appear to leave footprint pheromones’ these bias traffic/scouts in other contexts, e.g. emigration to a new nest

Division of labor

The central theme in our work on division of labor is understanding the degree of individual variation in cooperative groups that is adaptive, and dissecting the processes that make it so. Counterintuitively, several of our studies on bumble bees and Temnothorax ants have found that conventional explanations, such as higher individual efficiency of specialists, don’t hold, and that specialization is generally much lower than often portrayed for social insects.
Interestingly, other than the classic ‘division of labor to increase work output’ as proposed by Adam Smith (for humans), we have proposed other processes, both adaptive and non-adaptive, that may lead to division of labor: in particular, the interaction of comparative advantage effects and cost of worker production, and the possibility that ‘unemployed’ workers are actually crucial for effective task allocation. We also demonstrated that inactive workers can make up a high proportion of colony workforce, and we discuss additional explanations, such as fast-fluctuating task demands, traffic congestion, and low worker quality (perhaps because of immaturity).

size variation

Bumble bee workers show extreme size variation within colonies. We showed that this is not a lab artefact, and that it is specific to workers (not queens or males), suggesting it is not a necessary result of a disorganized nest, for example (see Spatial Pattern below). The evolutionary benefit of such size variation however is unclear. Smaller workers appear to store more fat, and this may drive longer survival in some conditions, but generally large workers perform better (and live longer). Incidentally we show that under unpredictable food availability, workers generally store more fat.

Several manuscripts on this topic are in review - these suggest that small workers may be cheap, low quality workers for tasks where high quality workers are not worth their cost.

Kelemen EP, Skyrm K, Dornhaus A 2022, ‘Selection on size variation: more variation in bumble bee workers and in the wild’, Insectes sociaux 69: 93-98 - pdf - field-raised bumble bee colonies produce the same variation in worker body size as lab-raised ones; and variation is much higher in workers than males or queens, suggesting different selection on body size consistency

Couvillon MJ, Jandt J, Duong N, Dornhaus A 2010 ‘Ontogeny of worker body size distribution in bumble bee (Bombus impatiens) colonies’, Ecological Entomology 35: 424-435 - pdf - with the exception of the very first worker generations (not studied here), bumble bee colonies produce the same degree of variation in worker body size throughout ontogeny, and produce varying body sizes even in simultaneously raised workers (contradicting the idea that standing size variation may reflect change in size across generations)

Couvillon MJ, Dornhaus A 2010 ‘Small worker bumble bees (Bombus impatiens) are hardier against starvation than their larger sisters', Insectes sociaux 57: 193-197 - pdf - when deprived of food in the lab, larger workers died earlier than smaller ones (support for the idea that perhaps there is a performance-robustness tradeoff across worker sizes)

Couvillon MJ, Jandt J, Bonds J, Helm BR, Dornhaus A 2011 ‘Percent lipid is associated with body size but not task in the bumble bee Bombus impatiens’, Journal of Comparative Physiology A 197: 1097-1104 - pdf - smaller workers store more fats; larger workers more often forage and smaller ones more often nurse, and ‘nursing’ in social insects is often associated with lipid storage (and foraging with leanness) - but here we found that size, not task, was the better explanation for differences in lipid storage

Jandt J, Dornhaus A 2014 'Bumblebee response thresholds and body size: does worker diversity increase colony performance?', Animal Behavior 87:97-106 - pdf - worker variation did not increase colony performance; instead, average worker preference for a task seems to predict task performance; one of two colony level manipulation studies from our lab showing this same lack of benefit of division of labor (the other is Kelemen et al. 2020, below)

Kelemen EP, Davidowitz G, Dornhaus A, 2020, ‘Size variation does not act as insurance in bumble bees; instead, workers add weight in an unpredictable environment’, Animal Behaviour 170, 99-109 - pdf - found no increased performance at whole-colony level of worker variation under different feeding regimes; unclear whether the result from Couvillon & Dornhaus 2010 was replicated here, but did find that lipid storage plastically responded to predictability of food.

Couvillon MJ, Fitzpatrick G, Dornhaus A 2010 ‘Ambient air temperature does not predict whether small or large workers forage in bumble bees (Bombus impatiens)', Psyche 2010, doi:10.1155/2010/536430 - pdf - tested the hypothesis that larger bees may be more prone to overheating (16deg C vs 36deg C) but did not find this to be the case.

Westling JN, Harrington K, Bengston S, Dornhaus A 2014 ‘Morphological differences between extranidal and intranidal workers in the ant Temnothorax rugatulus, but no effect of body size on foraging distance’, Insectes sociaux 61: 367-369 - pdf - although Temnothorax are often considered ‘monomorphic’, we show they do exhibit quite a bit of size variation among workers, and this may be associated with task. However, the latter did not replicate in Charbonneau et al. 2017, Integrative and Comparative Biol.

see also Jandt & Dornhaus 2009 for first comprehensive analysis of bumble bee division of labor

Inactive workers

It has long been known that many workers in social insect colonies appear ‘inactive’; we demonstrate thoroughly that some individuals are consistently inactive, in field and lab, and across contexts.

Why? Inactive workers may be too immature to work effectively, or young workers may serve as food storage or be more likely to be selfish.
Inactive workers, as a group, can also function as a ‘reserve force’, replacing active workers on demand, as previously hypothesized; however evidence for this is mixed.

Dornhaus A, Holley J-A, Pook VG, Worswick G, Franks NR 2008 'Why do not all workers work? Colony size and workload during emigrations in the ant Temnothorax albipennis', Behavioral Ecology and Sociobiology 63: 43-51 - pdf - some workers work a lot, many only a little (often more than half do not participate in emigrations at all); and this is more pronounced in smaller colonies (perhaps contrary to intuition; this result is correlative); quorum thresholds in emigration seem to be set relative to colony size

Charbonneau D, Dornhaus A 2015 ‘Workers ‘specialized’ on inactivity: Behavioral consistency of inactive workers and their role in task allocation’, Behavioral Ecology and Sociobiology 69: 1459-1472 - pdf - inactivity is the behavioral state that most differentiates workers; differs between workers and colonies and negatively correlates with all other tasks; circadian rhythm does not explain this

Charbonneau D, Hillis N, Dornhaus A 2015 ‘‘Lazy’ in nature: ant colony time budgets show high ‘inactivity’ in the field as well as in the lab’, Insectes sociaux 62: 31-35 - pdf - time budgets in field colonies do not differ from those in lab colonies [paper of the year in Ins soc]

Charbonneau D, Dornhaus A 2015 ‘When doing nothing is something. How task allocation strategies compromise between flexibility, efficiency, and inactive agents’, Journal of Bioeconomics 17: 217-242 - pdf - conceptual review of possible adaptive explanations for large numbers of inactive workers, particularly that of fast-fluctuating workloads

Charbonneau D, Poff C, Nguyen H, Shin MC, Kierstead K and Dornhaus A, 2017. Who Are the “Lazy” Ants? The Function of Inactivity in Social Insects and a Possible Role of Constraint: Inactive Ants Are Corpulent and May Be Young and/or Selfish, Integrative and Comparative Biology, 57(3), pp.649-667 - pdf - systematic test of the predictions of 6 hypotheses for why there are so many inactive workers in ant colonies is consistent with the idea that inactive workers may be young, possibly too immature or corpulent to work effectively

Charbonneau D, Sasaki T and Dornhaus A, 2017. Who needs ‘lazy’ workers? Inactive workers act as a ‘reserve’ labor force replacing active workers, but inactive workers are not replaced when they are removed, PloS one, 12(9): e0184074 - pdf - empirical demonstration of the often proposed idea that inactive workers are a reserve force that can recover colony function if previously active workers are removed; inactive workers are not replaced when removed, indicating that they did not perform some hidden function while ‘inactive’

Jandt JM, Robins NS, Moore RE, Dornhaus A 2012 ‘Individual bumblebees vary in response to disturbance: a test of the defensive reserve hypothesis’, Insectes sociaux 59: 313-321 - pdf - inactive bumble bee workers are not particularly likely to respond to disturbance directly or by increasing guarding behavior afterwards; thus they do not seem to act as ‘reserve’, at least for defense

see also Pinter-Wollman et al. 2012 on the reserve hypothesis

Cognition

If there is one theme of research in animal information processing and decision making, it is that insects qualitatively seem to have (almost?) all the cognitive skills we find in humans (see also youtube lectures on intelligence). We contribute by showing that bumble bees assess reliability of different information channels and use them accordingly (visual vs olfactory, social vs personal).

However, we also show that innate predispositions contribute a large amount to what we think of as ‘intelligent’ behavior; and indeed that information is costly, particularly sampling large sets of stimuli or learning large numbers of cues, and thus may also be ignored altogether to increase speed and thus efficiency of foraging.

Speed vs Accuracy

Accuracy, i.e. using information or even learning, trades off with speed; and bees care about the cost of errors.

Chittka, L, Dyer, A, Bock, F, Dornhaus, A 2003 ‘Bees trade off foraging speed for accuracy’, Nature 424: 388 - pdf - some individuals are consistently faster but more error-prone; if errors are costly, all bees distinguish colors more accurately

also see Franks et al. 2003 below for group-level speed vs accuracy

Complex tasks and innovation

Some tasks improve more with learning than others; and tasks that give no rewards until after learning may depend on innate persistence to ever be learnt.

Muth F, Keasar T, Dornhaus A 2015 ‘Trading off short-term costs for long-term gains: how do bumblebees decide to learn morphologically complex flowers?’, Animal Behaviour 101: 191-199 - pdf - bees learned a complex task if they had an innate preference for the flower color; if not, they did not persist enough to be able to overcome initial failure

Leonard AS, Brent J, Papaj DR, Dornhaus A 2013 ‘Floral nectar guide patterns discourage nectar robbing by bumble bees’, PLoS One 8: e55914 - pdf - if you make legitimacy easier, more people stick to it - simple

Barker J, Dornhaus A, Bronstein J, Muth F 2018 ‘Learning about larceny: experience can bias bumble bees to rob nectar’, Behavioral Ecology and Sociobiology 72:68 - pdf - learning may improve efficiency of some tasks more than others; and bees tend to repeat behaviors they know

Individual differences

Individuals differ. But what environments or conditions select for more or less such individual variation? In black widows, individual differences increase in social (high density) situations, and decrease with disturbance (both effects during development).

Interestingly, differences across multiple axes often come together in ‘syndromes’ (such that a difference in one axis predicts differences in another). For individual social insects, this is studied in the context of division of labor (see section above), although often the syndromes may not conform to classic interpretations (e.g. be more along axes of low vs. high quality workers or lean/old vs fat/young workers).

Groups, or colonies, also differ. We show here that ant colonies vary along a life-history-related syndrome that is driven by nest site competition.

colony personalities

Temnothorax ant colonies in habitats with high nest site competition invest more in sexuals (reproduction) and less in growth (new workers), and this correlates with a suite of behavioral traits - a life history syndrome at the colony level.

Bengston SE, Dornhaus A 2014 'Be meek or be bold? A colony-level behavioural syndrome in ants', Proceedings of the Royal Society: Biological Sciences 281: 20140518 - pdf - in colonies from habitats across a large geographic range (Western US), field foraging distance correlates negatively with lab aggression, number of foragers, and threat response: these thus form a syndrome (independent of colony size, no of queens, or no of brood) - with more risk-tolerant (fewer, further foragers, more aggressive) colonies at higher latitudes; activity level in the colony without threat does not correlate.

Bengston SE, Dornhaus A 2015 ‘Latitudinal variation in behaviors linked to risk-tolerance is driven by nest-site competition and spatial distribution in the ant Temnothorax rugatulus’, Behavioral Ecology and Sociobiology 69: 1265-1274 - pdf - of a large number of environmental variables (related to competition, predation, resource avail-ability, or environmental stress), nest-site competition best predicts colony behavior (more competition ~ more risk tolerant)

Bengston S, Shin M, Dornhaus A 2017 ‘Life-history strategy and behavioral type: Risk-tolerance reflects growth rate and energy allocation in ant colonies’, Oikos 126: 556-564 - pdf - more risk-tolerant colonies invest more in reproduction (queens/males) and less in growth (new workers)

black widows

Black widows vary in a low vs high investment in foraging (many sticky threads) vs self-defense (better hideout) syndrome; we show that variation among individuals is driven by juvenile experience, and that social environments increase, but disturbance decreases variation.

DiRienzo N, Johnson C, Dornhaus A 2019 'Juvenile social experience generates differences in behavioral variation but not averages', Behavioral Ecology 30: 455–464 - pdf - juvenile experience of high density increases, juvenile experience of disturbance decreases population-level variation in behavior of black widow spiders

DiRienzo N, Schraft HA, Montiglio PO, Bradley CT, Dornhaus A, 2020, ‘Foraging behavior and extended phenotype independently affect foraging success in spiders’, Behavioral Ecology 31, 1242-1249 - pdf - aggressive behavior and number of gumfooted (sticky foraging) silk lines in the web of a black widow spider both and independently increase foraging success

Schraft H, Bilbrey C, Olenski M, DiRienzo N, Montiglio PO, Dornhaus A 2022 Injected serotonin decreases foraging aggression in black widow spiders (Latrodectus hesperus), but dopamine has no effect, Behavioural Processes 204: 104802 - what the title says

DiRienzo N, Bradley CT, Smith CA, Dornhaus A 2019 'Bringing down the house: male widow spiders reduce the webs of aggressive females more' Behavioral Ecology and Sociobiology 73: 1-10 - pdf - males tear down a portion of the females web; this study shows that this probably mostly serves to reduce female cannibalism (instead of signaling or reducing competition from other males)

Collective decision-making

Groups making decisions may need to integrate information from different individual sources, evaluate it, and reach consensus on an action. Temnothorax ant colonies need to regularly choose and move to new nest sites, and may modulate the decision process based on urgency, collect information even before it becomes relevant, flexibly change decisions even during the process of implementation, and can serve as a model surprisingly closely resembling groups of neurons making decisions in the brain.

choosing a new nest

Presence of other colonies, pheromone marks, or dead ants are taken into account in nest site choices (and all avoided); a large number of traits of nest sites are evaluated, including size and number and width of entrances.

Franks NR, Dornhaus A, Hitchcock G, Guillem R, Hooper J, Webb C 2007 ‘Avoidance of conspecific colonies during nest choice by ants’ Animal Behaviour 73: 525-534 - pdf - if emigrating in the presence of another conspecific colony, ant colonies chose the nest cavity furthest away from it; unless the two colonies fused (with or without casualties); pheromone marks around nests play a role

Franks, N R, Dornhaus, A, Best, C S, Jones, E L 2006 ‘Decision-making by small and large house-hunting ant colonies: one size fits all’, Animal Behaviour 72: 611-616 - pdf - colonies of any size like large nests, perhaps because they grow into them; larger colonies have more scouts and discover more nests, but then use a higher quorum threshold and more reverse tandem runs to recruit transporters

Franks, N R, Dornhaus, A, Metherell, B G, Nelson, T R, Lanfear, S A J, Symes, W 2006 ‘Not everything that counts can be counted: Ants use multiple metrics for a single nest trait’, Proceedings of the Royal Society: Biological Sciences 273: 165-169 - pdf - ants use light level in the nest to help select nests with fewer, narrower entrances, but other metrics are likely also used; no evidence for strict ‘counting’

Franks, N R, Hooper, J, Webb, C, Dornhaus, A 2005 ‘Tomb evaders: house-hunting hygiene in ants’, Biology Letters 1: 190-192 - pdf - ants avoid new nest sites with dead ants in them

Franks, NR, Dornhaus, A 2003 ‘How might individual honeybees measure massive volumes?’, Proceedings B: Biology Letters 270 (Supplement 2): 181-182 - pdf - conceptual paper suggesting that honey bees could use a Buffon’s-Needle-like algorithm to estimate internal area (as ants do) and add 3-dimensional free diameter measurements to estimate volume of a potential nest cavity

Arriving at consensus

The decision-making process in colony emigrations of Temnothorax ants is generally well-studied (see also other publications by NR Franks and S Pratt and others).

Key general contributions here are showing the similarity to known neural decision-making networks in the brain, and how quorum thresholds can modulate between speed and accuracy of decisions.

For ants, we demonstrate learning of information that will only later be relevant (latent learning, and ability to move unprompted in response to opportunity) and flexibility during the decision (changing their collective mind mid-stream).

Dornhaus, A, Franks, NR, Hawkins, RM, Shere, HNS 2004 ‘Ants move to improve – colonies of Leptothorax albipennis emigrate whenever they find a superior nest site’, Animal Behaviour 67: 959-963 - pdf - ants can emigrate from an intact nest to a preferred one, but it takes long and they use a high quorum threshold

Franks NR, Hooper JW, Dornhaus A, Aukett PJ, Hayward AL, Berghoff S 2007 ‘Reconnaissance and latent learning in ants’, Proceedings of the Royal Society: Biological Sciences 274: 1505-1509 - pdf - ants learn about nest properties even before they need a new nest, and remember this information when the need arises later

Dornhaus A, Franks NR 2006 ‘Colony size affects collective decision-making in the ant Temnothorax albipennis’, Insectes sociaux 53: 420-427 - pdf - ants adjust their individual-level behavior to their colony size; colony size effects are not just emergent from higher worker number. It is likely that quorums are set as relative to colony size in the original nest. Learning may have bigger effects in small colonies.

Franks, NR, Dornhaus, A, Fitzsimmons, JP, Stevens, M, 2003, ‘Speed vs Accuracy in Collective Decision-Making’, Proceedings of the Royal Society: Biological Sciences 270: 2457-2463 - pdf - when time is of the essence, ants use a lower quorum threshold, i.e. rely on fewer other individuals and agreement in the group; as a result, collective decisions become faster and less accurate.

Franks NR, Hooper JW, Gumn M, Bridger TH, Marshall JAR, Groß R, Dornhaus A 2007 ‘Moving targets: collective decisions and flexible choices in house-hunting ants’ Swarm Intelligence 1: 81-94 - pdf - ants

Planque R, Dornhaus A, Franks NR, Kovacs T, Marshall JAR 2007 ‘Weighting waiting in collective decision-making’ Behavioral Ecology and Sociobiology 61: 347-356 - pdf - data re-analysis and model show that ants, in collective emigration decisions, are flexible early on and grow more committed (and inflexible) over time, which is an effective strategy

Marshall JAR, Bogacz R, Dornhaus A, Planque R, Kovacs T, Franks NR, 2009 ‘On optimal decision-making in brains and social insect colonies’ Journal of The Royal Society Interface 6: 1065-1074 - pdf -

Marshall JAR, Bogacz R, Planqué R, Dornhaus A, Kovacs T, Franks NR 2011 ‘On optimal decision making in brains and social insect colonies’, in: Seth et al. (eds.) Modelling Natural Action Selection. Cambridge University Press - pdf -

Marshall JAR, Dornhaus A, Franks NR, Kovacs T 2006 ‘Noise, cost and speed-accuracy trade-offs: decision making in decentralised systems’, Journal of The Royal Society Interface 3: 243-254 - pdf -

Franks NR, Dornhaus A, Marshall JAR, Deuchaume-Moncharmont F-X, 2009. ‘The dawn of a golden age in mathematical insect sociobiology’. In: Eds. J. Fewell & J. Gadau, Organization of Insect Societies, Harvard University Press [invited chapter in edited book] - pdf -

Marshall JAR, Kovacs T, Dornhaus AR, et al 2003 ‘Simulating the evolution of ant behaviour in evaluating nest sites’, Lecture Notes in Artificial Intelligence 2801: 643-650 2003 - pdf -

Spatial pattern

Spatial patterning is an important mechanism of organization in social insect colonies, both within and across nests (if polydomous); this creates informational heterogeneity.

Worker sorting

In the first demonstration of an organizational mechanism for creating polymorphic workers in social insect colonies, we demonstrated that nurse bees stick to the center of the nest but brood is spread across the nest, leading to less-fed, smaller pupae in the periphery.

Nonetheless, bumble bees maintain development temperature in all brood across the nest.

Workers of some other tasks are also non-randomly sorted across the nest area.

Jandt JM, Dornhaus A 2009 Spatial organization and division of labor in the bumble bee, Bombus impatiens Animal Behaviour 77: 641-651 - pdf - bumble bees, like some ants, maintain spatial fidelity zones in the nest (small areas where they are predictably found), with brood care workers/smaller workers in smaller zones nearer the nest center, and foragers near the periphery even when inactive; this paper is also the first to demonstrate body size-based division of labor in bumble bees for tasks other than foraging, and shows a weak age differences between tasks

Couvillon MJ, Dornhaus A 2009 Location, location, location: larvae position inside the nest is correlated with adult body size in worker bumble bees (Bombus impatiens), Proceedings of the Royal Society: Biological Sciences 276: 2411-2418 - pdf - larvae nearer the center of the nest are fed more and turn into larger pupae; together with the results above that nurses stick to the center, this provides a mechanism for generating body size variation in concurrently raised brood

Jandt JM, Dornhaus A 2011 Competition and cooperation: bumblebee spatial organization and division of labor may affect worker reproduction late in life, Behavioral Ecology and Sociobiology 65: 2341-2349 - pdf - workers who reproduced after the queen was gone were larger, older, and had been both more inactive and staying away from the queen when she was still around; suggesting inactive workers may gain selfish benefits later even if not immediately

Kelemen E, Dornhaus A 2018 ‘Lower temperatures decrease worker size variation but do not affect fine-grained thermoregulation in bumble bees’, Behavioral Ecology and Sociobiology 72: 170 - pdf - bees maintain high temperatures for brood in all locations, but not honeypots; under higher ambient temperatures, more size variation in brood is produced

Architecture

Since spatial distribution of workers in the nest matters, it is likely that nest cavity shape and built architecture have important roles to play; this new area of research is being explored.

We show that even simple built walls by Temnothorax colonies have repeatable, and thus possibly relevant, traits.

First description of social pseudoscorpion nests in detail.

DiRienzo N, Dornhaus A 2017 ‘Temnothorax rugatulus ant colonies consistently vary in nest structure across time and context’, PLoS One 12: e0177598 - pdf - colonies build walls to encircle the used part of their nest cavity; these are thicker when colonies have more brood, thicker, longer, and heavier with higher ambient humidity, and colonies differ (repeatably) in the traits of walls they build

Chapin KJ, Kittle A, Dornhaus A 2022 Social pseudoscorpion nest architecture provides direct benefits to group members and rivals the efficiency of honey bees, Journal of Arachnology 50: 323-334 - pdf - detailed quantitative descriptions of nests of social pseudoscorpions, which consist of multiple molting chambers or ‘cells’; calculation of the amount of silk saved by cooperation

not yet peer reviewed: Chism GT, Nichols W, Dornhaus A 2022 Nest shape influences colony organization in ants: spatial distribution and connectedness of colony members differs from that predicted by random movement and is affected by nest space, bioRXiv - distribution of colony members differs between tube- and circle-shaped nests and shows non-random distribution, but spatial fidelity zones remain the same

not yet peer reviewed: Chism GT, Dornhaus A 2022 Temnothorax rugatulus ants do not change their nest walls in response to environmental humidity, bioRXiv - external humidity does not have an effect on built nest walls, but ant colonies have preferences for particle sizes and wall density that are consistent between field and lab

Networks

Networks among spatially viscous or unmoving agents show high clustering, and slow information flow along the global network.

More work in this area, particularly with large empirical datasets, is needed.

Blonder B, Dornhaus A 2011 'Time-ordered networks reveal limitations to information flow in ant colonies', PLoS One 6: e20298 - pdf - using a large empirical dataset of ant-ant interactions, we show that indeed spatial viscosity in movement strongly limits information flow in ant colonies

Charbonneau D, Blonder B, Dornhaus A 2013 ‘Social insects: a model system for network dynamics’, in: Holme P and Jari S: Temporal Networks. Springer books - pdf - conceptual review paper showing examples of network analysis in social insect research and calling for more use of such techniques

Polydomy

Multiple nests can be used by ants to effectively exploit a larger area; and ‘outstations’ may be nest-like even without brood in them.

Also, ants make cockroach jerky.

Lanan MC, Dornhaus A, Bronstein J 2011 'The function of polydomy: the ant Crematogaster torosa preferentially forms new nests near food sources and fortifies outstations', Behavioral Ecology and Sociobiology 65: 959-968 - pdf - ants make beef jerky (out of cockroaches); colonies may inhabit multiple nest sites, and if so, often found new nests near valuable locations; and while a ‘nest’ technically is only a cavity with brood in it, ants also inhabit ‘outstations’ (without brood, but with adults and which are fortified, so not just short-term shelters for foragers); workers are preferentially associated with a particular nest even if there is traffic between nests

Collective contests

The primary enemy of ants are other ants, often of the same species: ants are not only competitors for rare nest sites in species that do not build their own nests, but also fight over brood (which is a valuable resource); and ant colonies often completely eliminate other colonies. It is thus likely that strategies for assessing and defending against other ant colonies are as sophisticated as those for, for example, collective foraging. Here we show that defensive resources are adaptively allocated across colony space, and that colonies, probably using interaction rates and assessment of worker speed and aggression, can determine the fighting ability and resources held by other colonies to determine their contest behavior.

Defense allocation

Donaldson-Matasci MC, Powell S, Dornhaus A 2022, ‘Distributing defenses: How resource defendability shapes the optimal response to risk’, The American Naturalist 199: 636-652 - pdf - a general mathematical model to determine the optimal strategy for dividing defenses among assets depending on their value, defendability, and risk of attack; shows that if some assets have little chance of being defended, it is better to give up and not invest in their defense at all

Powell S, Donaldson-Matasci M, Woodrow-Tomizuka A, Dornhaus A 2017 ‘Context-dependent defences in turtle ants: Resource defensibility and threat level induce dynamic shifts in soldier deployment’, Functional Ecology 31: 2287-2298 - pdf - soldier ants (majors which block entrances with their heads) in the turtle ant Cephalotes rohweri are dynamically and adaptively allocated among multiple nests of the same colony

Powell S, Dornhaus A 2013 ‘Soldier-based defences dynamically track resource availability and quality in ants’, Animal Behaviour 85: 157-164 - pdf - see also above paper: soldiers in C. rohweri are adaptively allocated among nests but in a conservative bet-hedging strategy

Enemy assessment

Chapin KJ, Paat VA, Dornhaus A 2022 ‘Brood as booty: the effect of colony size and resource value in social insect contests’, Behavioral Ecology 33: 549-555 - pdf - Temnothorax colonies contest over brood (which can be eaten or raised as ‘slaves’ in the winner colony); we demonstrate that colonies can assess both their opponent’s and their own colony size (fighting ability) and resource value (amount of brood) to determine their contest strategy

Various

...

Methods

Dornhaus A, Smith B, Hristova K, Buckley L 2021, ‘How can we fully realize the potential of mathematical and biological models to reintegrate biology?’, Integrative and Comparative Biology: icab142 - pdf -

Nguyen H, Fasciano T, Charbonneau D, Dornhaus A, Shin MC 2014 ‘Data Association Based Ant Tracking with Interactive Error Correction’; WACV 2014: IEEE Winter Conference on Applications of Computer Vision. (peer reviewed conference paper) - pdf -

Fasciano T, Dornhaus A, Shin MC 2014 ‘Ant Tracking with Occlusion Tunnels’; WACV 2014: IEEE Winter Conference on Applications of Computer Vision. (peer reviewed conference paper) - pdf -

Fletcher M, Dornhaus A, Shin MC 2011 ‘Multiple Ant Tracking with Global Foreground Maximization and Variable Target Proposal Distribution’, WACV '11 Proceedings of the 2011 IEEE Workshop on Applications of Computer Vision WACV p570-576 (peer reviewed conference paper) - pdf -

Fasciano T, Nguyen H, Dornhaus A, Shin MC 2013 ‘Tracking multiple ants in a colony’, Applications of Computer Vision (WACV), 2013 IEEE Workshop pp.534-540 (peer reviewed conference paper) - pdf -

Rice L, Dornhaus A, Shin MC 2015 ‘Efficient Training of Multiple Ant Tracking’, WACV 2015: IEEE Winder Conference on Applications of Computer Vision. Peer-reviewed conference paper - pdf -

Fasciano T, Dornhaus A, Shin MC 2015 ‘Multiple Insect Tracking with Occlusion Sub-tunnels’, WACV 2015: IEEE Winder Conference on Applications of Computer Vision. Peer-reviewed conference paper - pdf -

Blonder B, Wey TW, Dornhaus A, James R, Sih A ‘Temporal dynamics and network analysis’, Methods in Ecology and Evolution 3: 958-972 - pdf -

Ecology

Ferguson HM, Dornhaus A, Beeche A, Borgemeister C, Gottlieb M, Mulla MS, Gimnig JE, Fish D, Killeen GF 2010 ‘Ecology: a prerequisite for Malaria elimination and eradication’, PLoS Medicine 7: e1000303 - pdf -

Jones EI, Dornhaus A 2011 'Predation risk makes bees reject rewarding flowers and reduce foraging activity', Behavioral Ecology and Sociobiology 65: 1505-1511 - pdf -

Huang MH, Dornhaus A 2008 ‘A meta-analysis of ant social parasitism: host characteristics of different parasitism types and a test of Emery's rule’ Ecological Entomology 33: 589-596 - pdf -

Bengston SE, Dornhaus A 2013 ‘Colony size does not predict foraging distance in the ant. Temnothorax rugatulus: a puzzle for standard scaling models’, Insectes sociaux 60: 93-96 - pdf -