Behavioral Aspects of Rodent Population Biology

Five approaches to the study of behavior as a factor in thepopulation biology of rodents are outlined. These approachesare: (1) genetics, (2) physiology, (3) chronobiology,(4) behavioralecology and sociobiology, and (5) the use of intra- and inter-specificcomparisons. For each approach selected examples from the recentliterature, from this symposium and from my own work are usedto illustrate the new directions that are emerging for analyzingbehavior as a factor affecting natality, mortality and emigration/immigrationin rodents. Areas where additional investigations are neededare noted including some questions to be tested by future experiments.A variety of new techniques that have been developed or areinnovative applications of existing techniques are incorporatedinto the examples. Most of the material presented concerns fourgenera of rodents, Microtus, Mus, Peromyscus, and Rattus. Thefollowing major conclusions emerge from an overview of theseapproaches: Most recent research on behavior as a factor inrodent population biology has been concerned primarily withnatality. Few investigators have dealt with mortality and onlya limited number have explored emigration/immigration. Fieldand laboratory studies on behavioral aspects of rodent populationbiology have become more integrated than they were several decadesago. Both proximate and ultimate questions about behavior andpopulation dynamics are now being tested by both laboratoryand field methods. In contrast to some earlier approaches thatoften viewed population control as involving a single factor,most emerging conceptual frameworks have, as a major assumption,that population regulation is a multi-factorial process includingmany behavioral components.     

Spatial clumping of food and social dominance affect interference competition among ruddy turnstones

In studying the success of foraging animals, studies of interferencecompetition have put emphasis on effects of competitor density,whereas studies of resource defense have focused on the effectsof the spatial distribution of food within patches. Very fewstudies have looked at both factors simultaneously, that is,determined whether the effects of competitor density on foragingsuccess depend on the spatial distribution of food. We studiedthe behavior and the foraging success of ruddy turnstones (Arenariainterpres) using an experiment in which we varied both the presenceof a competitor and the food distribution. Because turnstonesmay differ strongly in their relative dominance status, we alsoexperimentally varied the foragers' relative dominance status.We found that the presence of a competitor only reduced theforaging success of subordinate birds foraging at the clumpedfood distribution. At this condition, dominant and subordinatebirds differed markedly in their foraging success. Contraryto our expectations, we did not observe more agonistic behaviorat the clumped food distribution. This indicates that the amountof agonistic behavior observed may be a bad indicator of interferenceeffects. These findings have specific implications for modelsof interference competition. Most notably they show that theeffects of competitor density on agonistic behavior and foragingsuccess may well depend on the spatial distribution of foodand the foragers' relative dominance status. Additionally, ourresults suggest that social dominance will not be fully understoodwithout considering long-term processes such as the formationand maintenance of social dominance hierarchies.     

Parasite-induced change in host behavior of a freshwater snail: parasitic manipulation or byproduct of infection?

Host behavioral changes due to parasitism are often assumedto be adaptations of the parasite. However, behavioral effectsof parasites may be a generalized response to parasitism andonly coincidentally beneficial for parasite transmission. Forthis reason, alternatives to the manipulation hypothesis shouldbe tested. Previous work demonstrated that the trematode parasiteMicrophallus sp. influences the behavior of the snail Potamopyrgusantipodarum in a way that may increase the probability of transmission.Here I report work conducted to test alternatives to the manipulationhypothesis. In a field study, the effect of Microphallus onbehavior was compared to that of two other castrating parasitegroups to determine if the behavioral change is simply a byproductof parasitism. Also, the foraging behaviors of infected anduninfected snails were examined in the presence and absenceof food resources to determine if the hunger level of Microphallus-infectedsnails could account for the parasite-induced behavioral change.First, Microphallus-infected snails were found on top of rocksduring the day less often than the two other parasite groups. Thisevidence suggests that the behavioral change caused by Microphallusis specific to Microphallus-infected snails. Second, Microphallus-infectedsnails responded to the lack of food differently from uninfectedsnails. Uninfected snails retreated to safer positions underrocks when the food source was removed from the top of the rocks,while Microphallus-infected snails remained on top of the rockswhere the risk of consumption by the final host is greater.Taken together with previous studies, these results suggestthat infection by Microphallus results in behavior that enhancesparasite transmission.     

Early learning affects social dominance: interspecifically cross-fostered tits become subdominant

Social dominance influences the outcome of competitive interactionsover limited resources, and may hence be important for individualfitness. Theory thus predicts that its heritability will below and that non-genetic determinants of dominance should prevail.In this field experiment we reciprocally cross-fostered greattits (Parus major) to blue tits (Parus caeruleus) to investigatethe impact of early social experience on dominance status incompetition over food during winter. Controlling for potentialeffects of age, size, sex and site-related dominance, we showthat cross-fostered birds of both species were subdominant toconspecific immigrants, while controls originating from unmanipulatedbroods were dominant to conspecific immigrants. Furthermore,blue tits reared by blue tit parents but with at least one greattit broodmate had lower dominance status relative to conspecificimmigrants than did controls. Although great tits generallydominated blue tits, cross-fostered birds of both species initiatedmarginally more fights against the other species than did theirrespective controls, suggesting faulty species recognition.Since both social parents and broodmates strongly influencethe dominance behavior of offspring later in life, we concludethat social conditions experienced at an early age are crucialfor the determination of subsequent social dominance.     

Dominance and diet are unrelated within a population of invasive crayfish

Laboratory behavioral experiments are an important tool in ecology and evolution, but whether these behaviors reflect the field function of organisms is not always clear. Directly connecting laboratory behaviors to field interactions would increase understanding of a variety of organisms. A recent study proposed using stable isotopes to link laboratory behaviors to field function of individuals, but failed to find any such links within a population of the invasive rusty crayfish Faxonius rusticus (Girard, 1852). Here, we assessed whether methodological decisions around tissue analyzed for stable isotopes, laboratory acclimation time, and timing of primary consumer collection may have affected the result, hypothesizing that more dominant crayfish would have higher trophic positions and tissue with faster turnover rates may exhibit a stronger association between laboratory behavior and recent field function. We tested this relationship using F. rusticus individuals from a single population, and related laboratory dominance to stable isotope-derived trophic position using linear regression. We failed to find a relationship between dominance and trophic position, regardless of our different methodologies. Future studies should consider alternative behaviors that may better relate to function in the field and also investigate whether laboratory behavior and field function are related between, rather than within, populations or species.

     

Testosterone in juvenile and adolescent male chimpanzees (Pan troglodytes): effects of dominance rank, aggression, and behavioral style

Testosterone is a steroid hormone with diverse effects on male reproductive function and behavior. The relationship between testosterone and social behavior such as mating and aggression has been investigated in a variety of primate species, but few such studies have been conducted on chimpanzees, and even fewer on primates during the juvenile and adolescent periods. This study explores the relationship between baseline urinary testosterone and behavioral variables including dominance rank, rates of aggression toward peers, and behavioral style in 16 juvenile and adolescent male chimpanzees (Pan troglodytes) living at the New Iberia Research Center in Louisiana. Behavioral observations and urine collection occurred during four research periods, each a year apart. After correcting for the positive association between testosterone and age, testosterone was positively associated with both dominance rank and rates of aggression directed at others. It was negatively associated with rates of aggression received. Individuals scoring highest in the "mellow" behavioral style component showed higher levels of testosterone than individuals scoring lowest in this component, an effect that may be partially due to the confounding effect of rank. The results of this study suggest that hormonal changes during the period preceding adulthood are not simply programmed physiological processes tied primarily to age-related change, but that important age-independent relationships with behavior also exist.     

Hatchery selection promotes boldness in newly hatched brown trout (Salmo trutta): implications for dominance

By using newly hatched (approximately 2 weeks old) brown trout(Salmo trutta) from six families of wild and six families ofsea-ranched origin (seventh generation), we tested the hypothesesthat (1) the hatchery environment selects for increased boldness,and (2) boldness predicts dominance status. Sea-ranched troutspend their first 2 years in the hatchery before being releasedinto the wild at the onset of seaward migration. Trout werepresented with a novel object (tack) and with food (brine shrimp),and their responses were measured and scored in terms of boldness.Siblings with increasing difference in boldness were then pairedin dyadic contests. Fish of sea-ranged origin were on averagebolder than were fish of wild origin, and bolder individualswere more likely to become dominant regardless of origin. Boldnesswas not related to RNA levels, indicating that bold behaviorwas not a consequence of higher metabolism or growth rate. Neitherwas size a predictor of bold behavior or the outcome of dyadiccontests. These results are consistent with studies on olderlife stages showing increased boldness toward predators in hatchery-selectedfish, which suggests that behavioral consequences of hatcheryselection are manifested very early in life. The concordancebetween boldness and dominance may suggest that these behaviorsare linked in a risk prone-aggressive phenotype, which may bepromoted by hatchery selection. However, we also found significantvariation in behavioral and growth-related traits among families,suggesting that heritable variation has not been exhausted bysea-ranching procedures.     

棉蚜获毒后禁食对其保持并传播黄瓜花叶病毒的影响

采用棉蚜Aphis gossypii 甜瓜Cumumis melo 黄瓜花叶病毒（cucumber mosaic virus, CMV）体系,研究棉蚜获毒后在空气中禁食对其保持并传播黄瓜花叶病毒的影响。结果表明获毒后的禁食时间与棉蚜传毒效率呈负相关。运用EPG (electrical penetration graph)及其即时显示、即时中断技术研究分析棉蚜禁食后的早期传毒行为细节。结果显示：禁食处理没有显著影响电势落差（potential drop,pd）数目及穿刺过程中出现的第一个pd波形前穿刺时间这两个重要指标,但禁食处理能引起pd波的两个亚波形pdⅡ-1和pdⅡ-2持续时间的显著减短。进一步分析未禁食棉蚜传毒作用与pd亚波形的关系,显示传毒可能与pdⅡ-2的持续时间相关（P＝0.06）。因此,pdⅡ-2的持续时间可能是与棉蚜传毒相关的一个行为指标。该研究还建立了新的高效而稳定的获毒方法---5pd获毒法,与传统的5min获毒法相比,获毒效率显著提高。     

The relation between dominance and exploratory behavior is context-dependent in wild great tits

Individual differences in personality affect behavior in novelor challenging situations. Personality traits may be subjectto selection because they affect the ability to dominate others.We investigated whether dominance rank at feeding tables inwinter correlated with a heritable personality trait (as measuredby exploratory behavior in a novel environment) in a naturalpopulation of great tits, Parus major. We provided clumped resourcesat feeding tables and calculated linear dominance hierarchieson the basis of observations between dyads of color-ringed individuals,and we used an experimental procedure to measure individualexploratory behavior of these birds. We show that fast-exploringterritorial males had higher dominance ranks than did slow-exploringterritorial males in two out of three samples, and that dominancerelated negatively to the distance between the site of observationand the territory. In contrast, fast-exploring nonterritorialjuveniles had lower dominance ranks than did slow-exploringnonterritorial juveniles, implying that the relation betweendominance and personality is context-dependent in the wild.We discuss how these patterns in dominance can explain earlierreported effects of avian personality on natal dispersal andfitness.     

Fitness consequences of foraging success in water striders (Gerris remigis; Heteroptera: Gerridae)

Foraging theory's central premise assumes that behavioral actionsthat affect the acquisition of food also ultimately affect fitness.However, very few investigations in behavioral ecology actuallydemonstrate ultimate fitness consequences of any particularbehavior. Many studies focus on short-term estimates of fitnessin animals with considerable life spans. I investigated whetherdifferences in foraging performance, as related to patch choiceand behavioral dominance, show consistent patterns over extendedperiods of an animal's life such that they translate into differencesin reproductive success. In one experiment, I focused on theability of single female water strider (Gerris remigis) to discriminatebetween foraging patches of differing prey abundances in laboratorystreams simulating natural conditions. In another experiment,I focused on competitive interactions within groups of threefemale water striders foraging in laboratory streams where resourceavailability varied with position in a patch. The first experimentshowed that, overall, the individuals detected and preferredto forage in the richer patch, which directly increased theirforaging success and their lifetime fecundity. However, therewas marked variability among individuals in their ability torespond to the differences in food availability, ranging fromno tracking to fast tracking when the patch qualities were switched.In the second experiment, both the foraging position of individualsin the stream and their dominance rank were reasonably consistentover their entire reproductive life. Foraging position and dominancerank were significant predictors of lifetime fecundity, theformer being the better predictor due to an imperfect correlationbetween the two variables. In both experiments, the increasein fecundity was achieved by higher oviposition rates ratherthan by extending the oviposition period or by inducing firstreproduction at an earlier age. [Behav Ecol 1991; 2: 46-55)     

Mating system structure and population density in a polygynous lizard, Sauromalus obesus (= ater)

Increases in population density often are associated with achange in mating system structure in numerous taxa. Typically,male interactions are minimal in extremely low density populations.As density increases, males exhibit territoriality but if densitybecomes too high, the energetic cost of defending a territorywill eventually outweigh the reproductive benefits associatedwith territoriality. Consequently, males in high density populationsmay abandon territoriality and adopt dominance polygyny, lekking behavior, or scramble competition. We investigated the relationshipbetween population density and mating system structure in threepopulations of the chuckwalla, Sauromalus obesus (= ater),near Phoenix, Arizona. Densities in the Phoenix Mountains (2.7chuckwallas/ha) were lower than any population previously studied.In the Santan Mountains (10.9 chuckwallas/ha), densities weresimilar to populations studied in the Mojave Desert, and inthe South Mountains (65 chuckwallas/ha), densities were the highest yet recorded. Male mating behavior was examined by determininghome range overlap and by making direct behavioral observations.Male home range size decreased with increasing population density.There was little overlap in home ranges among males in allthree populations, whereas home ranges of males and femalesconsistently overlapped, indicating that males were strictly territorial. This conclusion was supported by behavioral observationsof interactions among individuals in a natural setting. Thenumber of females wihin male territories was correlated withfood resources (plants) in all three populations. Female homerange size appeard to be related to food resources whereasmale home ranges appeared to be related to female distribution,population density, and geology. The retention of territoriality in spite of high population densities raises new questions aboutthe relationship between density and resource defense.     

Evidence that the parasitic nematode Skrjabinoclava manipulates host Corophium behavior to increase transmission to the sandpiper, Calidris pusilla

We found evidence that a nematode (Skrjabinoclava morrisoni) adaptivelymanipulates the behavior of its intermediate host (the amphipod Corophiumvolutator) to increase its likelihood of transmission to itsfinal host (the semipalmated sandpiper, Calidris pusilla). We foundthat male and female amphipods parasitized by nematodes increasedtheir surface activity in the field during daytime, but notduring nighttime hours. Increased surface activity is knownto increase susceptibility of amphipods to predation by sandpipersduring the day, but not at night, when sandpipers do not feedvisually. Also, as predicted by the manipulation hypothesis,only late-stage (infective) larvae of nematodes were associatedwith behavioral changes of amphipods. We found no evidence thatparasites were associated with other amphipod behaviors in thelaboratory, such as trail complexity, distance traveled, orburrow-probing activity of crawling males as would be expectedif parasitized hosts altered their own behavior. Survivorshipof amphipods was also unaffected by parasitism, which may favorparasite transmission. Thus, behavioral changes of parasitizedhosts were simple, and their expression was context-dependentand related to likelihood of predation. We argue that maturationtimes of nematodes in relation to migration schedules of sandpipers providea narrow window of opportunity and may explain why nematodes manipulateamphipod behavior.     

Predation risk and mating behavior: the responses of moths to bat-like ultrasound

In the presence of predators, animals may reduce or alter theirmating activities. There has been little experimental studyof whether mating behavior varies with the level of predationrisk. Two species of moths, Pseudaletia unipuncta (Noctuidae)and Ostrinia nubilalis (Pyralidae), significantly reduced theirmate-seeking behavior under high levels of simulated predationrisk. Male moths aborted upwind flight in a pheromone plume,and females stopped releasing pheromone in response to soundssimulating the echolocation calls of bats. For O. nubilalis,but not for P. unipuncta, the response varied significantlywith the level of predation risk. Interspecific differencesin behavioral responses likely reflect differences in physiologicalauditory sensitivity and/or behavioral thresholds. Female behavioralresponses to sounds simulating the calls of bats that gleantheir prey from surfaces were significantly weaker than theirresponses to sounds resembling the calls of aerially hawkingbats; these results support the neurophysiological data thatthe calls of gleaning bats are relatively inaudible to moths.This study indicates that some animals can modify their reproductiveactivities in response to auditory cues from predators.     

Dominance hierarchy and reproductive conflicts among subordinates in a monogynous queenless ant

In insect societies lacking morphologically specialized breedersand helpers, reproduction is often restricted to behaviorallydominant individuals. Such societies occur in about 100 speciesof ants that have secondarily lost the queen caste. All females,who are morphologically workers, can potentially mate and layeggs but only a few do so, and we demonstrate in Dinoponeraquadriceps that this is regulated by a dominance hierarchy.Six types of agonistic interactions allowed the ranking of 5-10workers in the hierarchy (n = 15 colonies). In particular, alphaand beta had characteristic behavioral profiles and were easily recognized.Only alpha mated, and workers ranking beta to delta sometimes producedunfertilized, male-destined eggs. Natural replacements (n = 19)and experimental removals (n = 15) of alpha demonstrated that betawas the individual most likely to replace alpha, although gammaand more rarely delta sometimes did, and we discuss the conflictthat occurs among high-ranking individuals over who should replacealpha. After such replacements, the new alpha behaved more aggressivelythan the overthrown alpha. Newly emerged workers tended to reachhigh ranks and displaced older high-ranking individuals downthe hierarchy. Low-ranking subordinates often prevented high-rankingindividuals from replacing alpha by biting and holding theirappendages (worker policing), which is consistent with the patternof relatedness associated with monogyny and monandry in D. quadriceps. Weinvestigated the relative importance of chemical communicationand dominance interactions to regulate reproduction. Alpha,beta, and sterile workers have different signatures of cuticularhydrocarbons, and these may provide honest information whichunderpins worker policing by low-ranking individuals.     

Behavioral flexibility in rats and mice: contributions of distinct frontocortical regions

Research examining the contribution of genetics to behavior is increasingly focused on higher order behavioral and cognitive processes including the ability to modify behaviors when environmental demands change. The frontal cortices of mammals, including rodents, subserve a diverse set of behavioral and cognitive functions including motor planning, social behavior, evaluation of expected outcomes and working memory, which may be particularly sensitive to genetic factors and interactions with experience (e.g. stress). Behavioral flexibility is a core attribute of these functions. This review orients readers to the current landscape of the literature on the frontocortical bases of behavioral flexibility in rodent laboratory experiments. Studies are divided into three broad categories: reversal learning, inhibitory learning and set‐shifting. Functional dissociations within the broader scope of behavioral flexibility are reviewed, followed by discussion of the associations between specific components of frontal cortex and specific aspects of relevant behavioral processes. Finally, the authors identify open questions that need to be addressed to better establish the constituents of frontal cortex underlying behavioral flexibility.     

Behavioral Regulatory Mechanisms in the Social Homeostasis of Termites (Isoptera)

Recent progress in the elucidation of the behavioral mechanismsresponsible for maintaining the steady state in a termite colonyis discussed. It is pointed out that various homeostatic mechanismshave common behavioral mechanisms. The mechanisms involved innest repair and construction, foraging, defense, and generalmass movements within nests are explained. An experiment isdescribed in which differential responses to a discrete simplestimulus (heat) are made by two different castes of Nasulitermescorniger and the homeostatic significance emphasized. The factorsin volved in foraging behavior in Reticulitermes are described.An hypothesis for the significance of the behavioral phenomenonof head-banging in termites is put forward: it is consideredto be connected with the maintenance of accelerated activityin a colony after "General Alarm".     

Social Reversal of Sex‐Biased Aggression and Dominance in a Biparental Cichlid Fish (Julidochromis marlieri)

In biparental species, aggression, dominance, and parental care are typically sexually dimorphic. While behavioral dimorphism is often strongly linked to gonadal sex, the environment—either social or ecological—may also influence sex‐biased behavior. In the biparental cichlid fish Julidochromis marlieri, the typical social environment for breeding pairs consists of large females paired with smaller males. While both sexes are capable of providing territory defense and parental care, the larger female provides the majority of defense for the pair, while the smaller male remains in the nest guarding their offspring. We examine the contributions of sex and relative mate size to these sex‐biased behaviors in monogamous J. marlieri pairs. Both female‐larger and male‐larger pairs were formed in the laboratory and were observed for territorial aggression (against conspecifics and heterospecifics), dominance, and parental care. In female‐larger pairs, territorial aggression and intra‐pair dominance were female‐biased, while in male‐larger pairs this bias was reversed. For both pairing types, the presence of an intruder amplified sex differences in territorial aggression, with the larger fish always attacking with greater frequency than its mate. Though less robust, there was evidence for plasticity of sex‐bias for some egg care related behaviors in the inverse direction. Our study suggests that relative mate size strongly influences the sex bias of aggression and dominance in J. marlieri and that this aspect of the social environment can override the influence of gonadal sex on an individual's behavior. The remarkable plasticity of this species makes Julidochromis an exciting model that could be used to address the relationship between proximate and ultimate mechanisms of behavioral plasticity.     

Thermal Sensitivity as a Specialization for Prey Capture and Feeding in Snakes

Behavior can be regarded as a result of various processes ofdecision based on the information provided by the sensory organs.In this review the role of the so-called additional heat sense,next to vision, smell and mechanoreception is discussed withrespect to the feeding behavior of snakes. The hierarchy ofthe sensory information in various phases of the feeding behaviordiffers between snakes possessing heat receptors (e.g., speciesof the Crotalinae and Pythoninae) and those without (e.g., speciesof the Viperinae and Colubrinae). Probably depending on theinfluence of ecological demands, visual or chemical cues arethe main information in the behavioral phases before the strikeHowever, in situations with little visual input, e.g., in darkness,rodents' burrows, etc., hunting behavior is guided in the firstplace by radiation of warm objects in Crotalus, Python and Trimeresurusflavoviridis, and by substrate vibrations in Vipera aspis, Pituophismelanoleucus and Boa constrictor. I suggest that in the sensoryhierarchy, heat information functionally replaces the mechanicalinformation which is utilized by snakes without pit organs.Poststrike behavior on the other hand is mainly guided by chemicalcues in all snakes.     

Predicting community structure of ground-foraging ant assemblages with Markov models of behavioral dominance

Although interference competition is a conspicuous component of many animal communities, it is still uncertain whether the competitive ability of a species determines its relative abundance and patterns of association with other species. We used replicated arena tests to quantify behavioral dominance of eight common species of co-occurring ground-foraging ants in the Siskiyou Mountains of southern Oregon. We found that behavior recorded in laboratory assays was an accurate representation of a colony's ability to monopolize resources in the field. We used interaction frequencies from the behavioral tests to estimate transition probabilities in a simple Markov chain model to predict patterns of relative abundance in a metacommunity that is dominated by behavioral interactions. We also tested whether behavioral interactions between each pair of species could be used to predict patterns of species co-occurrence. We found that the Markov model did not accurately predict patterns of observed relative abundance on either the local or the regional scale. However, we did detect a significant negative correlation at the local scale in which behaviorally dominant species occupied relatively few baits. Pairwise behavioral data also did not predict species co-occurrence in any site. Although interference competition is a conspicuous process in ant communities, our results suggest that it may not contribute much to patterns of relative abundance and species co-occurrence in the system studied here. However, the negative correlation between behavioral dominance and bait occupancy at the local scale suggests that competition-colonization trade-offs may be important in resource acquisition and persistence of behaviorally subordinate species.     

Kinship and Dominance Rank Influence the Strength of Social Bonds in Female Geladas (Theropithecus gelada)

In many primates, close social relationships are associated with lower stress, better health, and increased life span. However, individuals do not form bonds indiscriminately; rather, they focus on a few primary partners. This suggests that the identity of the partner may be as important as the bond itself. Although dominance and kinship have repeatedly emerged as salient predictors of female relationships, most of this research comes from species with multimale, multifemale groups and strict dominance hierarchies. Further, kinship was typically determined based on either behavior or on known mother–daughter relationships alone. To understand the generality of previous findings, we use behavioral and genetic sampling to examine whether dominance rank and/or genetic relatedness mediate female social bonds in geladas (Theropithecus gelada) living in the Simien Mountains National Park, Ethiopia. First, we found that, even though females in the same unit are closely related, female geladas still preferentially bond with the closest of these relatives. Second, females that were close kin formed the strongest bonds with females of similar rank to themselves. Finally, rank disparity predicted grooming rates but did not predict whether females were nearest neighbors. This suggests that, in contrast with data from other cercopithecines, spatial proximity among females may be less indicative of strong social bonds for geladas, a species that routinely exhibits a high degree of spatial overlap with extra-unit individuals. Together, these results highlight the importance of combining genetic data with detailed behavioral observations to help us understand how individuals choose and interact with social partners.