The bench of Justices Ranjana P Desai and N V Ramana had been trying their best to get the couple come to an amicable settlement and had directed the mother to bring the child to the court on October 16 as the man complained that the wife had not allowed him to meet his son since The bench felt that the father, Rajesh Puliyanklath, had a right to meet the child and on October 15 directed Sreeja V to produce the child in court.
On October 16, there was no trace of her son. In the circumstances, we are not inclined to take up this matter. The matter be removed from our Board. Images of experimental fights. Conflict induced by joining host tree canopies of separate colonies a with a tarp to catch casualties. We calculated the cost to victor colonies as the total worker loss and proportion of the initial colony lost. Host trees may be precariously defended by a diminished worker force after fights, and at elevated risk from herbivore damage and attack by other space-limited neighbor colonies. We examined colony response before and after fights to simulated large mammalian herbivore browsing using methods modified from Palmer and Brody Two observers carefully approached each tree and visually identified an isolated branch with new growth and 1 swollen thorn domatium within 15cm of the tip.
Both observers simultaneously raked focal branches 3 times with a leather-gloved left hand, then enveloped the branch with the mitt. After 30s, observers placed the entire mitt, with its attached attacking workers, into a plastic bag. Observers then sealed and froze the bags. After freezing, observers counted the number of workers in each bag.
Beginning 6 days after paired fights concluded, the above procedure was repeated. Changes in mean within-colony response prefight response — postfight response, where response is equal to the mean number of ants on a mitt across trials between controls and experimental fights were analyzed using t -tests in JMP 8. Each tree was photographed in the morning and afternoon 2 and 3 days before the fight and in the morning of the day before the fight.
A total of 5 photographs were taken on days 6—9 following fight initiation day 6 afternoon, day 7 morning and afternoon, day 8 afternoon, day 9 morning. Macrodigital photos always captured the south-facing side of stems and a size-standard ruler. We recorded the number of ant heads visible in the frame from the ground up to 10cm.
We took the mean of the 5 photographs from both the prefight and postfight periods, and compared experimental and control treatments using t -tests in JMP 8. Individuals from 18 experimental colonies ID1—ID9 pairs, Supplementary Table S1 were analyzed using molecular techniques for all sample sizes see Figure 2.
Experimental design and molecular results for 9 experimentally induced fights between single-tree colonies of Crematogaster mimosae. Fights were induced by joining tree canopies together. Proportion of samples assigned by microsatellite genotype to colony X or Y at each sampling period are indicated by the color in bars gray and black, respectively. In fight pairs 8 and 9, we found immature individuals belonging to both Colony X and Y. In fight pair 9, the lightest gray represents offspring of a queen that was not present at fight initiation, and who was likely a daughter of the original Colony X queen.
We infer 3 fight outcomes with respect to colony genetic structure: rejection of all non-kin enemies, adoption of non-kin enemies, and colony fusion. For postfight collections, we collected 5 domatia from each combined 2-tree system on 15 September hereafter 2 mos. For prefight and 2 mos. Individuals were inspected under a dissecting microscope to ensure they had no appendages missing indicating they were alive at the time of collection and not cached casualties or recently killed workers and to exclude body parts of other individuals.
For 8 mos.
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Immature ants collected 8 months after fights are unlikely to have been present at the time of conflict see Supplemental Materials — Supplementary Data for development time estimates. Outcomes where all 8 mos. Alternatively, if genotypes matching both prefight colony samples could be found among the immatures, an incomplete takeover is indicated no losing colony can be identified because both queens survived and continued to contribute to worker production.
Finally, novel genotypes whose maternity did not match reconstructed queens from the prefight sample suggest that a new queen was present within the colony. This could occur through secondary takeover by nonrelatives, or via the emergence of reproductive daughter queens Rubin et al.
We genotyped prefight and 8 mos. For 2 mos. Fragment data were visualized and scored using STRand Version 2. Null alleles and scoring errors were accounted for by incorporating an expected error rate of 0. No a priori relationships were assumed. For each colony, we inferred the number of queens and reconstructed the genotypes of maternal and paternal lineages at each sampling period.
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Between-colony relatedness coefficients for fight pairs were based on data from all 17 microsatellite loci. Because 2 mos. For fight pairs where the molecular analysis did not reveal a distinct victor ID8 and ID9, Figure 2c , the prefight relatedness coefficient used for statistical analysis is the mean of the within-colony relatedness coefficients of the 2 colonies Figure 4.
For each fight pair, individuals were grouped by their sampling origin: Colony X prefight, Colony Y prefight, 2 mos. We then estimated mean within-colony relatedness for each of these sample groups. We report Triadic Maximum Likelihood estimates of relatedness coefficients TrioML because their values are bounded at 0 to 1, rendering interpretation more intuitive.
In contrast to other pairwise relatedness estimators, this measure uses a third reference individual to help minimize error Wang To test for effects of fighting on relatedness within victor colonies through time, we compared the TrioML relatedness values across prefight, 2 mos. Because TrioML relatedness coefficients are bounded at 0 and 1, we used an arcsine square root transformation prior to analysis to better meet the assumption of normality.
Following fights, victorious colony territories nearly doubled in size proportion of domatia prefight vs. Vulnerability following conspecific conflict. Victorious colonies decreased defense after battles. Unmanipulated controls did not show a decrease in defense over the same period.
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Sixteen of the 18 prefight colonies were determined to contain full- and half-sib workers produced by a single queen, and the remaining 2 prefight colonies ID were inferred to include workers produced by 2 queens. Average within colony relatedness r TrioML over time for each fight manipulation. Prefight relatedness values are from fight victor colonies, except for ID8 and ID9. For these 2 colonies, where no definitive losing colony could be identified, values are the mean within-colony r TrioML for the 2 colonies.
Victor colonies ID1 and ID2 open symbols contained no losing colony genotypes 2 mos. Solid symbols identify colonies with enemy adoptees losing colony genotypes present at 2 mos. Line symbols identify colonies with no definitive fight victor where colony fusion is inferred both prefight colony genotypes are present in workers at 2 mos.
We infer 3 different categorical outcomes from molecular analysis of fights: complete rejection of non-kin, enemy adoption, and fusion Figure 2c. We therefore infer complete takeovers by 1 colony, without subsequent incorporation of non-kin, in these 2 fight pairs. In the remaining 7 pairs, samples of live workers at 2 months following fights included full siblings with both of the 2 prefight colonies. For these 5 fights then, molecular data suggest a single colony succeeded in conflict, and subsequently adopted the unrelated brood, and perhaps workers, of losing colonies.
Queens in these fights of losing colonies were either no longer present or no longer contributing brood to the colony at 8 months. Surprisingly, for fights ID8 and ID9, genotyped brood at 8 mos. This result indicates that neither colony succeeded in completely overtaking the other, and that both prefight queens were present and producing offspring. The average relatedness for brood found within each colony at 8 mos.
Are territorial fights between C. We found that battle casualties reduce victor colony size by one-third on average and by as much as two-thirds. In contrast to investigations highlighting advantages to combat victory beyond resource acquisition e.
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Previous manipulations of ant abundances on A. The actively growing shoot tips favored by large herbivores are also the site of carbohydrate-rich extrafloral nectar production, which colonies rely on to fuel activity and feed developing larvae Palmer et al. Nest-limited neighbors could discover the diminished resource holding capacity of victors either via territory scouts Palmer et al. Neighbors may then apply the information gained through monitoring to target weakened competitors Clutton-Brock et al.
In a system where colony size underlies competitive success Palmer , we document decreased worker number and defense of hosts by victors.
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We hypothesize that public battles fought to gain territory may subject victor colonies to increased risk of attacks and territory loss Adams ; Marden and Waage Are victorious colonies built back through the adoption of non-kin former enemies? We found that following fights, non-kin former enemies coexist with victorious colony members in shared nests. In 5 of the 9 induced fights, we analyzed genetically, despite prior lethal aggression between competitors, postfight colonies contained live workers that were full siblings with a prefight colony that lost Figure 2b.
We infer that for these 5 colonies, losing queens were either killed or escaped during fights. Because colonies in fighting pairs were estimated to each contain a similar number of brood prior to conflict, adoption of all losing colony brood would provide an instantaneous near-doubling of new workers emerging within the victor colony. Randomly selected C. Pupae can metamorphose into fully formed workers with no tending or nutritional input from adults Supplementary Table S1. Although complete worker development time egg to adult is unavailable for this species, Argentine ants Linepithema humile display pupal development rates that are similar to those observed for C.
Consequently, feeding losing colony brood to egg-laying queens or to larvae instead of adopting them directly would create a months-long payoff lag. Adoption of abundant non-kin brood is thus a more efficient way in terms of both time and energy of converting losing colony individuals into valuable workers than a potential alternative—cannibalism.
This study represents a rare example of this phenomenon, and the first evidence that non-kin enemy adoption can be triggered experimentally in nature via conflicts between large territorial colonies. Like many other obligate plant ants and cavity nesters, Crematogaster spp. Conspecific usurpation, though difficult to detect in nature, has been predicted Foitzik and Heinze and observed Palmer ; Rudolph K, personal observation. Our findings suggest that non-kin adoption associated with territorial battles could be an overlooked phenomenon, and potentially widespread in ants that engage in conflicts over nest sites or foraging grounds.
Bourke and Franks argue that selection for acceptance of orphaned workers by queen-right colonies should be strong. Such adoption in this case would necessitate that the queen-right colony identifies orphaned workers as such following the departure or death of their queen s. If only brood are adopted, adoption may be facilitated by limited brood odor and associated reduced aggression Vander Meer and Morel Although we infer that 5 victorious colonies adopted non-kin orphans, 2 fights unexpectedly resulted in queen-right colony fusions of mature colonies Figure 2 , a phenomenon reported previously in wild ants only from unicolonial species.
In fights ID8 and ID9, genetic data indicate that both Colony X and Y queens were alive and producing brood within a shared tree canopy 8 months after fights Figure 4. Yet we find that mortal combat with thousands of fatalities transitions to mutual acceptance over the course of hours in C. Not only does aggression toward brood and callow workers cease, but worker versus worker attacks do as well.
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We do not yet know how de-escalation between fighting colonies may proceed. Similar to instances of non-kin adoption, mutual acceptance of non-kin as colony members could result from cue changes odor homogenization through contact or acceptance changes without cue changes change in colony acceptance thresholds.
Regardless of the de-escalation mechanism, ultimate coexistence appears impossible without strong temporal plasticity in templates of recognition or chemical cues among adults. Cue changes might result from the extensive physical contact involved in fighting, where CHC signals are blended are diluted, making non-kin less distinguishable from one another, that is, passive affiliation Kronauer et al.
Information indicating a growing cost of conflict e. Non-kin colony cooperation in times of vulnerability has precedent in ants, for example, joint colony founding Bernasconi and Strassmann and queenless colony fusion Kronauer et al. In the third outcome, 2 victorious colonies did not adopt non-kin at least in numbers detectable in our samples Figure 2 and did not fuse.
If all losing colony workers were eliminated in battle and this brood ejection behavior continued, it could explain the lack of non-kin for ID1 and ID2. The findings of complete rejection, as well as temporary rejection followed by acceptance of non-kin in this study, underscore the tension between the benefits of non-kin adoption and the threat that non-kin could represent to colony cohesion. Further exploration could help determine whether reduced relatedness among nestmates following conflict has unexpected consequences for colony function and the extent of recognition plasticity.
For example, do adoptees perform work within victorious colonies? Our findings represent a rare experimental quantification of the costs and consequences of escalated fighting for victors in their natural environment. Application of molecular analysis to a behavioral study exposed the leakiness of colony boundaries in C. Importantly, this non-kin affiliation occurs within large mature colonies, not recently founded ones Pollock and Rissing A study of conflict in wood ants showed that violent wars fought to expand territory produced casualties that were fed to developing larvae Mabelis Our work points to a different, potentially underappreciated source of profit for colonies that succeed in conflict.
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After costly contests, C. Non-kin adoptees provide victors with an accelerated means to colony size recovery. Supplementary material can be found at Supplementary Data. Many thanks to intrepid field assistants A. Eshwa, A. Inzauri, J. Ekiru, and J. Lemboi and to all the staff of the Mpala Research Centre. Sincerest thanks to M. Stanton, R. Grosberg, and B. Cameron for assistance with molecular work. Manuscript benefited tremendously from comments made by T. Palmer, L. Benedict, N. Tstsui, J. Goheen, J. Brockman, C.
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Sign In. Advanced Search. Article Navigation. Close mobile search navigation Article Navigation. Volume Article Contents. Lay Summary. Spoils of war and peace: enemy adoption and queen-right colony fusion follow costly intraspecific conflict in acacia ants Kathleen P. Oxford Academic. Google Scholar. Jay P.