Agonistic behaviour of wild eels and depressed survival and growth of farmed eels in mixed rearing experiments.

To contribute to improving eel stocking procedures, the survival, growth and behaviour of farmed Japanese eels reared together with wild individuals were evaluated to learn about possible encounters of farmed- and stocked eels with wild eels. Two experiments were conducted to evaluate the (a) effect of the presence of wild eels on survival and growth of farmed eels in experimental ponds and (b) behavioural differences between wild and farmed eels placed in a small tank. In the survival and growth experiment, significantly lower survival and growth rates of farmed eels reared with wild eels were observed compared with farmed eels reared without wild eels (control). In the behavioural observations, the authors observed significantly higher occupation rates of a refuge and more frequent biting for wild eels, indicating a dominance of wild eels over farmed eels. Because wild and farmed eels used in the current experiments belong to the same genetic population, these differences resulted within one generation through the aquaculture rearing process.

Agonistic behaviour and energy metabolism of bold and shy swimming crabs Portunus trituberculatus.

The metabolism and behaviour of crustaceans are highly flexible, and the inter-individual variation in these traits is evolutionarily and ecologically significant. We analysed the relationships among personality traits (boldness, activity and hesitancy), agonistic behaviour and energy status (glycogen, glucose and lactate) in the swimming crab Portunus trituberculatus The main results were as follows. (1) Boldness was significantly correlated with activity and hesitancy. Bold crabs were more likely to initiate and win a fight. In bold individuals, the frequencies of 'move to', 'cheliped display', 'grasp' and 'contact' were significantly higher than those of shy individuals, whereas the frequency of 'move away' was significantly lower than that of shy individuals. (2) Before fighting, the glucose concentrations in the haemolymph of bold individuals were significantly lower than those of shy individuals, whereas the concentrations of lactate showed the opposite trend. There were no significant differences in glycogen and lactate concentrations in the claw muscle between bold and shy individuals. (3) After fighting, the glucose and lactate concentrations in the haemolymph of both bold and shy individuals were significantly higher than those before fighting. The glucose concentrations in the haemolymph were significantly higher in bold individuals than shy individuals. In addition, bold individuals showed a larger increase in glucose in the haemolymph but a smaller increase in lactate compared with shy individuals. (4) After fighting, the glycogen concentrations in the claw muscle were significantly lower than those before fighting; however, there were no significant differences in the concentrations of lactate in the claw muscle. These results indicated that the agonistic behaviour of the swimming crab is related to its behavioural type. Energy reserves may be one of the factors affecting the personality traits and agonistic behaviour in crabs. These results should lay a foundation for in-depth understanding of the relationships among crustacean personality, agonistic behaviour and metabolic physiology.

The role of skill in animal contests: a neglected component of fighting ability.

What attributes make some individuals more likely to win a fight than others? A range of morphological and physiological traits have been studied intensely but far less focus has been placed on the actual agonistic behaviours used. Current studies of agonistic behaviour focus on contest duration and the vigour of fighting. It also seems obvious that individuals that fight more skilfully should have a greater chance of winning a fight. Here, we discuss the meaning of skill in animal fights. As the activities of each opponent can be disrupted by the behaviour of their rival, we differentiate among ability, technique and skill itself. In addition to efficient, accurate and sometimes precise movement, skilful fighting also requires rapid decision-making, so that appropriate tactics and strategies are selected. We consider how these different components of skill could be acquired, through genes, experiences of play-fighting and of real fights. Skilful fighting can enhance resource holding potential (RHP) by allowing for sustained vigour, by inflicting greater costs on opponents and by minimizing the chance of damage. Therefore, we argue that skill is a neglected but important component of RHP that could be readily studied to provide new insights into the evolution of agonistic behaviour.

Seasonal and daily changes in the behaviour of the adult emu (Dromaius novaehollandiae) under farm conditions.

1. The aim of this study was to present selected elements of emu behaviour in three different seasons (the non-breeding season, the pre-breeding season and the breeding season) and in the morning and afternoon. The study focused on feed, water and sand intake; plumage care; resting; agonistic behaviour and other activities, mostly standing still, aimless walking or observing the surroundings. 2. The troop consisted of 11 emus (5 females and 6 males) aged 12 years, kept in a free-range system. Observations were carried out during a period of 3 weeks in each season, during which 60-min observation sessions took place on 3 randomly chosen days. 3. A significant effect was found of each season on the behaviour patterns related to feeding, sand intake, plumage care, resting and agonistic behaviour. 4. Time of day affected the behaviour associated with the collection of food and water, foraging, pecking and ingesting sand and plumage preening. 5. Most of the time, regardless of season and time of day, the emus were standing still, walking aimlessly or observing the surrounding area.

Lateralisation in agonistic encounters: do mirror tests reflect aggressive behaviour? A study on a West African cichlid.

In this study, population level lateralisation and the suitability of mirror tests as a test of natural aggressive behaviour in male rainbow kribs Pelvicachromis pulcher was investigated. Aggressive behaviour in live agonistic trials correlated positively with behaviours towards a mirror image and no visual lateralisation was detected.

Testing the myth: tolerant dogs and aggressive wolves.

Cooperation is thought to be highly dependent on tolerance. For example, it has been suggested that dog-human cooperation has been enabled by selecting dogs for increased tolerance and reduced aggression during the course of domestication ('emotional reactivity hypothesis'). However, based on observations of social interactions among members of captive packs, a few dog-wolf comparisons found contradictory results. In this study, we compared intraspecies aggression and tolerance of dogs and wolves raised and kept under identical conditions by investigating their agonistic behaviours and cofeeding during pair-wise food competition tests, a situation that has been directly linked to cooperation. We found that in wolves, dominant and subordinate members of the dyads monopolized the food and showed agonistic behaviours to a similar extent, whereas in dogs these behaviours were privileges of the high-ranking individuals. The fact that subordinate dogs rarely challenged their higher-ranking partners suggests a steeper dominance hierarchy in dogs than in wolves. Finally, wolves as well as dogs showed only rare and weak aggression towards each other. Therefore, we suggest that wolves are sufficiently tolerant to enable wolf-wolf cooperation, which in turn might have been the basis for the evolution of dog-human cooperation (canine cooperation hypothesis).

Agonistic reciprocity is associated with reduced male reproductive success within haremic social networks.

While individual variation in social behaviour is ubiquitous and causes social groups to differ in structure, how these structural differences affect fitness remains largely unknown. We used social network analysis of replicate bluebanded goby (Lythrypnus dalli) harems to identify the reproductive correlates of social network structure. In stable groups, we quantified agonistic behaviour, reproduction and steroid hormones, which can both affect and respond to social/reproductive cues. We identified distinct, optimal social structures associated with different reproductive measures. Male hatching success (HS) was negatively associated with agonistic reciprocity, a network structure that describes whether subordinates 'reciprocated' agonism received from dominants. Egg laying was associated with the individual network positions of the male and dominant female. Thus, males face a trade-off between promoting structures that facilitate egg laying versus HS. Whether this reproductive conflict is avoidable remains to be determined. We also identified different social and/or reproductive roles for 11-ketotestosterone, 17ß-oestradiol and cortisol, suggesting that specific neuroendocrine mechanisms may underlie connections between network structure and fitness. This is one of the first investigations of the reproductive and neuroendocrine correlates of social behaviour and network structure in replicate, naturalistic social groups and supports network structure as an important target for natural selection.

Effect of wither application of an analogue of pig appeasing pheromone on encounters between unfamiliar mini-pigs.

BACKGROUND: The practice of mixing unfamiliar pigs on farms is common but results in fighting, welfare problems and performance issues. Pigs have different ways of resolving social conflicts, including aggressive and affiliative behaviours. Synthetic appeasing pheromones have demonstrated many positive effects in animal husbandry and are regularly used by breeders to improve animal welfare and performance. The aim of the study was to investigate the effect of a new method of applying pig appeasing-pheromone (PAP) to the withers in an experimental model of pig mixing to determine whether PAP reduced aggression and fighting, increased prosocial behaviours, and improved behavioural and physiological indicators of welfare. RESULTS: PAP reduced fighting between mini-pigs (df = 1; F = 13.47; P = 0.001; mixed logistic regression). Even if not significant, agonistic behaviours tended to be reduced when the treatment was applied (df = 1; F = 4.14; P = 0.058; mixed logistic regression). Likewise, mini-pigs seemed to be scored as not aggressive at all (df = 1; F = 3.61; P = 0.070; GLMM) and to be less aggressive toward the other pig than when placebo was applied. Concerning the latency of the first contact without aggression, a significant effect was found between the PAP and placebo groups (df = 1; χ2 = 4.74; P = 0.0295; Cox model). Moreover, even if not significant, the treated mini-pigs seemed to spent more time looking at each other (df = 1; F = 3.59; P = 0.071; GLMM) and immobile and/or ground sniffing (df = 1; F = 3.18; P = 0.088; GLMM) than those that received placebo. No significant difference was found between groups for salivary cortisol concentration (df = 1; F = 0.10; P = 0.752; GLMM), but variances between groups were heterogeneous at every time. No significant difference was found between groups regarding alpha-amylase activity (df = 1; F = 0.25; P = 0.621; GLMM), but variances between groups were heterogeneous at T0, T1 and T3. These results indicate that the variability (dispersion) within each group was lower when PAP was applied than when the placebo was applied. CONCLUSIONS: The new method of applying PAP improved welfare of mini-pigs (as models of domestic pigs) by reducing fighting, among other interesting results. PAP seems thus a promising biomimetic tool to enhance animal welfare in pig production systems.

Animal Welfare Consequences of Organic Boar Fattening and Occurrence of Boar Taint on Five Commercial Farms.

The welfare of male fattening pigs may be improved by refraining from castration, but may be compromised, in turn, by harmful social behaviour in groups of boars. In addition, boar taint may be problematic. This study aimed to evaluate these potential problems in boar fattening under commercial organic conditions. In total, 625 boars were compared with 433 barrows and 83 gilts regarding their social behaviour, lesions and lameness at 80 kg, before and after split marketing. The mixed-model analysis showed that significantly more short agonistic interactions, fights and mounting behaviours were observed in groups of boars. Agonistic interactions were reduced in spring/summer and when boars grew older. Fights and mounts were increased when boars had contact to female pigs in the neighbouring pen. No effect of split marketing, growth rate, homogeneity of groups, group size, feeding space and illumination hours could be detected. Increased interaction frequencies did not result in significantly more skin lesions, lameness, treatments or mortality. Increased space allowance reduced skin lesions. On 9.8% of the dissected boars' penises, wounds were detected; they were absent on two farms with generous litter provision. Boar taint prevalence, as detected by human nose method, was 1.44%. Under the studied organic husbandry conditions, boar fattening appears to be practicable, although penile injuries should be monitored at slaughter.

Part-time grouping of rabbit does in enriched housing: effects on performances, injury occurrence and enrichment use.

Collective housing is perceived as a possible way to improve rabbit welfare, especially among adult females, which are normally individually housed. Part-time group housing seems to allow a better balance between welfare and health than continuous group housing, but practical implementation and consequences for reproductive performance have not been extensively studied. The aim of this work was to compare weight, feed intake, litter size, injury occurrence, body integrity, and spatial location of female rabbits housed part-time in group housing (PGH group, n = 32) or housed individually (IH group, n = 8). Females were grouped by opening connecting hatches between four individual enriched cages (platform, wood stick and gnawing block). Collective housing started 12 days after artificial insemination and was interrupted at the 10th day (instead of the 15th day later as originally planned) due to high injury rates and severe injuries caused by fighting. The proportion of injured females increased from 25% on the first day of grouping to 63% on the 10th day. Female weight gain during the experiment was similar in the two groups. Litter size at weaning was 9% lower in the PGH group than in the IH group (9.2 vs 10.0, P < 0.01). Platform use was recorded in 32% of the observations and was the highest during the 2 weeks before weaning (46% and 47% of total observations, P < 0.05). During the grouping period, there were at least two females in the same housing unit in 59.4% of the cases, at least two females were located on the same level in 31.3% of the cases. In conclusion, connecting individual cages is an ergonomic solution for part-time group housing, but does not prevent fights for the establishment of a social hierarchy. This is detrimental to the health and body integrity of female rabbits.

Perceived Threat of COVID-19 Contagion and Frontline Paramedics' Agonistic Behaviour: Employing a Stressor-Strain-Outcome Perspective.

Historically, infectious diseases have been the leading cause of human psychosomatic strain and death tolls. This research investigated the recent threat of COVID-19 contagion, especially its impact among frontline paramedics treating patients with COVID-19, and their perception of self-infection, which ultimately increases their agonistic behaviour. Based on the stressor-strain-outcome paradigm, a research model was proposed and investigated using survey-based data through a structured questionnaire. The results found that the perceived threat of COVID-19 contagion (emotional and cognitive threat) was positively correlated with physiological anxiety, depression, and emotional exhaustion, which led toward agonistic behaviour. Further, perceived social support was a key moderator that negatively affected the relationships between agonistic behaviour and physiological anxiety, depression, and emotional exhaustion. These findings significantly contributed to the current literature concerning COVID-19 and pandemic-related effects on human behaviour. This study also theorized the concept of human agonistic behaviour, which has key implications for future researchers.

Dear enemy effect in the Mexican volcano mouse Neotomodon alstoni: Implications of sex in the agonistic behaviour among neighbours.

Two opposite phenomena have been found in territorial animals, the "dear enemy'' and the "nasty neighbour'', which refer to individuals that show less aggression toward neighbours than toward strangers and vice versa. However, the need to maintain territory should differ for males and females because sexual reproduction is the result of the different adaptive strategies of the sexes. In this study, we explore territorial behaviour in the context of dear-enemy and nasty neighbour effects in the Mexican volcano mouse (Neotomodon alstoni). Patterns of spatial relationships between individuals and the degrees of agonistic behaviour among neighbours were analysed for a period of one year. Results exhibit a greater spatial proximity between male pairs during the non-reproductive period than during the reproductive period, and greater spatial proximity between pairs of females during the reproductive period than during the non-reproductive period. The analysis of agonistic behaviour showed that there is less tolerance for distant neighbours than for nearest neighbours. However, there is a greater frequency of aggression between male pairs than between females, while females appear to exhibit non-aggressive avoidance among individuals. The results support the theory that Mexican volcano mouse exhibits the "dear enemy phenomenon".

The Effect of the Type of Non-Caged Housing System, Genotype and Age on the Behaviour of Laying Hens.

This study investigated the welfare of laying hens in different non-caged housing systems, namely a deep-litter barn system (BS), a free-range system (FRS) and an organic system (OS). The study was conducted on 270 hens of a native breed Green-legged Partridge (Z-11) and 270 Hy-Line Brown hybrids. Visual scans were performed to record behaviour of hens. Hens were housed in groups of 30 and observed over the course of one day at 20, 36 and 56 weeks of age. Dustbathing, scratching, wing stretching, wing flapping and preening were recorded as comfort behaviours. Pecking, fighting, threatening and chasing were recorded as agonistic behaviours. The percentage of run use was higher in native hens than in commercial hens (p < 0.05). The proportion of hens exhibiting comfort behaviours housed in the FRS and OS was similar but over twice as high as in the BS (p < 0.05). In the FRS and OS, the percentage of hens displaying comfort behaviours increased with age (p < 0.05). In all the production systems, the percentage of birds displaying comfort behaviours was higher in native breed hens than in commercial breeds (p < 0.05). In the BS, the higher proportion of hens displaying an agonistic behaviour was seen more in commercial breed than in the native breed hens (p < 0.05). The percentage of birds displaying an agonistic behaviour declined with hen age, both in commercial and native breed hens.

Aggression and discrimination among closely versus distantly related species of Drosophila.

Fighting between different species is widespread in the animal kingdom, yet this phenomenon has been relatively understudied in the field of aggression research. Particularly lacking are studies that test the effect of genetic distance, or relatedness, on aggressive behaviour between species. Here we characterized male-male aggression within and between species of fruit flies across the Drosophila phylogeny. We show that male Drosophila discriminate between conspecifics and heterospecifics and show a bias for the target of aggression that depends on the genetic relatedness of opponent males. Specifically, males of closely related species treated conspecifics and heterospecifics equally, whereas males of distantly related species were overwhelmingly aggressive towards conspecifics. To our knowledge, this is the first study to quantify aggression between Drosophila species and to establish a behavioural bias for aggression against conspecifics versus heterospecifics. Our results suggest that future study of heterospecific aggression behaviour in Drosophila is warranted to investigate the degree to which these trends in aggression among species extend to broader behavioural, ecological and evolutionary contexts.

The serotonin or dopamine by cyclic adenosine monophosphate-protein kinase A pathway involved in the agonistic behaviour of Chinese mitten crab, Eriocheir sinensis.

Agonistic behaviour is common in an encounter between two crustaceans. It often causes limb disability and consumes a lot of energy, which is harmful for the growth and survival of commercially important crustaceans. In the present study, we mainly focused on the agonistic behaviour of the Chinese mitten crab, Eriocheir sinensis, which is an important species of the aquaculture industry in China. We recorded agnostic behaviour with a high-definition camera and preliminarily evaluated the role of serotonin (5-HT) or dopamine (DA)-cyclic adenosine monophosphate (cAMP)-protein kinase A (PKA) pathway and eyestalk in the behaviour. The results showed that agonistic behaviour in E. sinensis consisted of three stages: approach, contact and fight. We found that the number of fights and cumulative time of fight were significantly higher in the male vs. male group than in the female vs. female and female vs. male groups (P < 0.05). After 1 h of agonistic behaviour, 5-HT concentration showed a significant increase and DA concentration showed a significant decrease when compared with the control group (no encounter; P < 0.05). 5-HT1B and 5-HT2B mRNA levels showed a significant increase in the eyestalk (P < 0.05). 5-HT7 mRNA levels showed significant downregulation in the thoracic ganglia and DA1A mRNA levels showed upregulation in the intestine (P < 0.05). DA2 mRNA levels showed a significant decrease in the eyestalk (P < 0.05). These changes were accompanied by a significant increase in cAMP level and significant decrease in PKA level in the haemolymph (P < 0.05). In addition, a significant decrease in glucose levels was detected after the agonistic behaviour. Crustacean hyperglycemic hormone (CHH) mRNA levels showed significant upregulation in the eyestalk and significant downregulation in the intestine (P < 0.05). The number of fights and cumulative time of fight in the left eyestalk ablation (L-X vs. L-X) group were more and longer than those in the intact eyestalk (C vs. C), right eyestalk ablation (R-X vs. R-X) and bilateral eyestalk ablation (D-X vs. D-X) groups. In short, E. sinensis shows special agonistic behaviour modulated by 5-HT or DA-cAMP-PKA pathway and eyestalk, especially the left eyestalk.

Intrasexually selected weapons.

We propose a practical concept that distinguishes the particular kind of weaponry that has evolved to be used in combat between individuals of the same species and sex, which we term intrasexually selected weapons (ISWs). We present a treatise of ISWs in nature, aiming to understand their distinction and evolution from other secondary sex traits, including from 'sexually selected weapons', and from sexually dimorphic and monomorphic weaponry. We focus on the subset of secondary sex traits that are the result of same-sex combat, defined here as ISWs, provide not previously reported evolutionary patterns, and offer hypotheses to answer questions such as: why have only some species evolved weapons to fight for the opposite sex or breeding resources? We examined traits that seem to have evolved as ISWs in the entire animal phylogeny, restricting the classification of ISW to traits that are only present or enlarged in adults of one of the sexes, and are used as weapons during intrasexual fights. Because of the absence of behavioural data and, in many cases, lack of sexually discriminated series from juveniles to adults, we exclude the fossil record from this review. We merge morphological, ontogenetic, and behavioural information, and for the first time thoroughly review the tree of life to identify separate evolution of ISWs. We found that ISWs are only found in bilateral animals, appearing independently in nematodes, various groups of arthropods, and vertebrates. Our review sets a reference point to explore other taxa that we identify with potential ISWs for which behavioural or morphological studies are warranted. We establish that most ISWs come in pairs, are located in or near the head, are endo- or exoskeletal modifications, are overdeveloped structures compared with those found in females, are modified feeding structures and/or locomotor appendages, are most common in terrestrial taxa, are frequently used to guard females, territories, or both, and are also used in signalling displays to deter rivals and/or attract females. We also found that most taxa lack ISWs, that females of only a few species possess better-developed weapons than males, that the cases of independent evolution of ISWs are not evenly distributed across the phylogeny, and that animals possessing the most developed ISWs have non-hunting habits (e.g. herbivores) or are faunivores that prey on very small prey relative to their body size (e.g. insectivores). Bringing together perspectives from studies on a variety of taxa, we conceptualize that there are five ways in which a sexually dimorphic trait, apart from the primary sex traits, can be fixed: sexual selection, fecundity selection, parental role division, differential niche occupation between the sexes, and interference competition. We discuss these trends and the factors involved in the evolution of intrasexually selected weaponry in nature.

Fight or flight? Effects of vaginal oestrus on cortisol, testosterone, and behaviour in guinea pig female-female interaction.

It is accepted that social stress in relation to confrontation and competition can elicit behavioural and hormonal changes in social mammals. These effects have, however, been less frequently studied among female-female interactions. In the present study female-female confrontation experiments were carried out to monitor socio-positive and agonistic behaviour by controlling for the oestrus cycles of 12 individuals. Additionally, plasma cortisol (CORT) and testosterone (T) levels were determined before and after the experiments. During non-oestrus conditions a significant increase in CORT levels from pre- to post confrontation was registered and females spent more time to sit side by side. During vaginal oestrus the confrontation experiments revealed avoiding of a conspecific female by showing increased flight behaviour. However, during that period no changes in CORT levels were found. But, a non-significant increase in T was measured from pre- to post confrontation in both cycle phases, while no differences in the display of aggressive behaviours were found. These findings indicate considerable influences of different oestrus cycle phases on social stress-induced CORT secretion and the modulation of socio-positive and agonistic behaviour in female guinea pigs.

Influence of seasonality and stimulus of oestrous does in bucks' aggressiveness.

Aggressiveness is directly related to testosterone concentration, which varies with seasons and in response to female stimulation. The aim was to determine if the frequency and pattern of agonistic interactions between bucks varies seasonally and in response to oestrous female stimulation. In the first study we characterized the pattern of agonistic interactions during feeding throughout a whole year in groups of Iberian ibex bucks; and in the second study, we determined the influence of oestrous does on the frequencies of agonistic behaviours between Gabon bucks in different seasons. In Iberian ibex bucks, the frequency of agonistic behaviours was maximum in summer and testosterone concentration in late autumn. In Gabon bucks, total agonistic interactions increased in bucks stimulated by females in late summer and autumn. In late autumn there were more interactions without physical contact in stimulated than in isolated bucks. Aggressiveness in bucks varied along the year, but the seasonal pattern was related to the context in which behaviours were recorded. Stimulation with oestrous females induced an increase of aggressiveness during the breeding season. At least in the conditions of these studies, testosterone concentrations seemed not to be the main influence on the frequency in which aggressive behaviour was displayed.

[Effects of socializing piglets in lactation on behaviour, including tail-biting, in growing and finishing pigs]. / Effekte einer frühen Sozialisierung von Ferkeln auf das Verhalten, unter anderem auf das Schwanzbeißen.

OBJECTIVE: The aim of this study was to determine whether early socialising of piglets influences the later behaviour and the risk of tail biting in growing and finishing pigs. MATERIAL AND METHODS: The behaviour of 183 animals (divided in three successive runs) was recorded from birth until the end of the fattening period. Furthermore, the condition of the teats of the sows as well as the integument and tails of the growing and finishing pigs were evaluated. To socialize the piglets, four litters of the experimental group (V) were provided with additional space (walkway) by opening "piglet-doors" (day 10 postpartum of the youngest litter). The piglets of the control group (K) were reared in conventional farrowing crates located in the same compartment of the stable. Post-weaning, the piglets were assigned to three groups: the experimental group (V/V, two litters of group V), the control group (K/K, two litters of group K) and the mixed group (V/K, one litter of group V and one litter of group K). RESULTS: After opening the "piglet-doors", piglets of group V displayed significantly more playing behaviour than piglets of group K. Additionally, the agonistic behaviour increased in group V. Post-weaning, at allocation and mixing, animals of group V/V showed significantly less agonistic behaviour than pigs of group K/K. Tail-biting behaviour occurred in all three groups, but only in runs two and three. On the 100th day of the fattening period, 58.7% of the pigs of group V/V, 51.7% of group V/K and 43.3% of group K/K still had intact tails. In pigs of group K/K, the tails were significantly shorter compared to the other two groups. CONCLUSION AND CLINICAL RELEVANCE: Early socialization enhances piglet welfare in farrowing pens by encouraging playing behaviour. Less agonistic behaviour at allocation could furthermore reduce stress at mixing and allocation. The aim to reduce the incidence of tail-biting could not be achieved. However, socializing piglets in lactation could contribute, in accordance with other measures, to an influence on biting behaviour.

Tiarajudens eccentricus and Anomocephalus africanus, two bizarre anomodonts (Synapsida, Therapsida) with dental occlusion from the Permian of Gondwana.

Anomodontia was a highly successful tetrapod clade during the Permian and the Triassic. New morphological information regarding two bizarre basal anomodonts is provided and their palaeoecological significance is explored. The osteology of the recently discovered Tiarajudens eccentricus Cisneros et al. 2011, from the Brazilian Permian, is described in detail. The taxon exhibits unusual postcranial features, including the presence of gastralia. Additional preparation and computed tomography scans of the holotype of Anomocephalus africanus Modesto et al. 1999 discovered in the Karoo Basin of South Africa allow a reappraisal of this genus. Anomocephalus is similar to Tiarajudens with regard to several traits, including a battery of large, transversally expanded, palatal teeth. Molariform teeth are present in the mandible of the African taxon, providing additional insight into the function of the earliest tooth-occlusion mechanism known in therapsids. At least two waves of tooth replacement can be recognized in the palate of Anomocephalus. The outsized, blade-like caniniforms of the herbivorous Tiarajudens allow several non-exclusive ecological interpretations, among which we favour intraspecific display or combat. This behaviour was an alternative to the head-butting practised by the contemporary dinocephalians. Combat specializations that are considered typical of Cenozoic herbivores likely evolved during the Middle Permian, at the time the first communities with diverse, abundant tetrapod herbivores were being assembled.