African jewel fish (Hemichromis bimaculatus) distinguish individual faces based on their unique iridophore patterns.

Previous research has shown that African jewel fish (Hemichromis bimaculatus) recognize pair-bonded mates during their exchanges of egg-guarding duties. The current research examined the perceptual cues for face recognition by comparing two face models displaying anatomically realistic arrangements of blue iridophores derived from discriminant function analysis of distinct sibling groups. Four groups each consisting of 9 subadults were examined using a narrow compartment restraining lateral movement where face models were presented at eye level for eight trials. Because respiratory movement of the operculum can mechanically displace the eye thereby shifting the retinal image, jewel fish reduce their respiration rate during increased attention. When two experimental groups were presented with the same face models on four trials following initial model presentations, both groups exhibited stable respiration rates indicative of model habituation. When the habituated face models were switched to novel face models on the fifth trial, the rates of respiration decreased as measured by reliable increases in the elapsed times of opercular beats. Switching the models back to the habituated models on the sixth trial caused reliable decreases in the elapsed times of opercular beats, resembling the earlier trials for the habituated models. Switching the face models again to the formerly novel models on the seventh trial produced respiration rates that resembled those of the habituated models. The two control groups viewing the same models for all eight trials exhibited no substantial change in respiration rates. Together, these findings indicate that jewel fish can learn to recognize novel faces displaying unique arrangements of iridorphores after one trial of exposure.

The Saliency of Snake Scales and Leopard Rosettes to Infants: Its Relevance to Graphical Patterns Portrayed in Prehistoric Art.

Geometrically arranged spots and crosshatched incised lines are frequently portrayed in prehistoric cave and mobiliary art. Two experiments examined the saliency of snake scales and leopard rosettes to infants that are perceptually analogous to these patterns. Experiment 1 examined the investigative behavior of 23 infants at three daycare facilities. Four plastic jars (15×14.5cm) with snake scales, leopard rosettes, geometric plaid, and plain patterns printed on yellowish-orange paper inside were placed individually on the floor on separate days during playtime. Fourteen 7-15-month-old infants approached each jar hesitantly and poked it before handling it for five times, the criterion selected for statistical analyses of poking frequency. The jars with snake scales and leopard rosettes yielded reliably higher poking frequencies than the geometric plaid and plain jars. The second experiment examined the gaze and grasping behavior of 15 infants (spanning 5months of age) seated on the laps of their mothers in front of a table. For paired comparisons, the experimenter pushed two of four upright plastic cylinders (13.5×5.5cm) with virtually the same colored patterns simultaneously toward each infant for 6s. Video recordings indicated that infants gazed significantly longer at the cylinders with snake scales and leopard rosettes than the geometric plaid and plain cylinders prior to grasping them. Logistic regression of gaze duration predicting cylinder choice for grasping indicated that seven of 24 paired comparisons were not significant, all of which involved choices of cylinders with snake scales and leopard rosettes that diverted attention before reaching. Evidence that these biological patterns are salient to infants during an early period of brain development might characterize the integration of subcortical and neocortical visual processes known to be involved in snake recognition. In older individuals, memorable encounters with snakes and leopards coupled with the saliency of snake scales and leopard rosettes possibly biased artistic renditions of similar patterns during prehistoric times.

Development of snake-directed antipredator behavior by wild white-faced capuchin monkeys: III. the signaling properties of alarm-call tonality.

In many primates, the acoustic properties of alarm calls can provide information on the level of perceived predatory threat as well as influence the antipredator behavior of nearby conspecifics. The present study examined the harmonics-to-noise ratio (tonality of spectral structure) of alarm calls emitted by white-faced capuchin monkeys (Cebus capucinus) in trees directed at photographic models of a boa constrictor, neotropical rattlesnake, scorpion eater snake, and white snake-shaped control presented on the ground. The average and peak harmonics-to-noise ratios of initial alarm calls by infants, juveniles, and adults and those of nearby second callers were analyzed using PRAAT software. Averaged for age class, the peak harmonics-to-noise ratio of alarm calls directed at the boa constrictor model, characterizing a primary capuchin predator, was reliably higher than the peak harmonics-to-noise ratio of alarm calls directed at the harmless scorpion eater model. This effect was influenced by the higher harmonics-to-noise ratio of infant alarm calls and it disconfirmed our prediction, based on primate vocalization research, that snake perception would increase arousal and alarm-call noisiness. Levels of call tonality did not distinguish the boa and rattlesnake or rattlesnake and scorpion eater models for any age class. Higher alarm-call tonality appeared contagious to nearby perceivers, with focal alarm calling influencing the level of tonality of the first calls of second callers. Together, these findings suggest that the higher peak harmonics-to-noise ratio of capuchin alarm calling directed at snakes is contagious and possibly conveys information about the level of perceived predatory threat.

Forward-facing predators attract attention in humans (Homo sapiens).

Even prey that successfully evade attack incur costs when responding to predators. These nonlethal costs can impact their reproductive success and survival. One strategy that prey can use to minimize these costs is to adjust their antipredator behavior based on the perceived level of risk. We tested whether humans adopt this strategy by presenting participants with photographic arrays of predators (lions) that varied in their level of risk. While their eye movements were recorded, the participants searched for a forward-facing predator (signifying potential predator interest; high-risk target) among an array of inattentive predators that were facing away (low-risk distractors) or searched for a predator that was facing away from them among an array of forward-facing predators. As a control, participants also searched through similar arrays that displayed a potential prey species (impalas) rather than predators. Participants detected forward-facing predators more quickly than predators facing away from them. Unexpectedly, they were also quicker at detecting forward-facing prey versus prey facing away from them, but slower to detect these forward-facing prey compared with forward-facing predators. They were slower to detect predators and prey facing away from them because they spent more time looking at the forward-facing distractors and looked at more of those distractors. The results indicate that human attention is drawn toward dangerous predators with forward-facing orientations, and this could allow humans to quickly assess predator intentions. (PsycINFO Database Record (c) 2018 APA, all rights reserved).

Evolutionary constraints on equid domestication: Comparison of flight initiation distances of wild horses (Equus caballus ferus) and plains zebras (Equus quagga).

Habituation to humans was an essential component of horse (Equus caballus ferus) domestication, with the nondomestication of zebras (Equus quagga) possibly reflecting an adaptive constraint on habituation. We present the human hunting hypothesis, arguing that ancestral humans hunted African animals, including zebras, long enough to promote a persistent wariness of humans, whereas a briefer period of hunting horses in Central Asia influenced by glacial cycles was unlikely to produce an equally persistent wariness. An alternative habituation to humans hypothesis, prompted by field observations, posits that zebras can habituate well to nonthreatening humans given sufficient exposure. If so, other factors must account for zebra nondomestication. To examine these hypotheses, we compared the flight initiation distances (FIDs) of wild horses in the United States and plains zebras in Africa to a human approaching on foot (N = 87). We compared the flight behavior of both species at sites with low and high exposure to humans (mean humans/acre = .004 and .209, respectively). Analyses revealed a significant interaction (p = .0001) between equid species and level of human exposure. The mean FIDs of horses (146 m) and zebras (105 m) with low human exposure did not differ appreciably (p = .412), but these distances were substantially longer (p < .0001) than those of horses (17 m) and zebras (37 m) with high human exposure that did differ significantly (p < .0001). The finding that plains zebras habituate less completely to humans than horses do might reflect an adaptive response to historical hunting and partly explain their resistance to domestication.

Isolation rearing reveals latent antisnake behavior in California ground squirrels (Otospermophilus becheeyi) searching for predatory threats.

This study of California ground squirrels (Otospermophilus beecheyi) investigated the long-term effects of isolation rearing on alarm-call recognition. Six wild-caught squirrels, trapped as yearlings, and six laboratory-reared squirrels were maintained in solitary cages for approximately 3 years prior to the study. Visual searching and olfactory searching were measured as squirrels emerged from their burrow-like nest box into a laboratory room after hearing repetitive playbacks of alarm calls or control sounds consisting of pulses of white-noise or ambient laboratory sounds. Before exiting completely after hearing alarm calls, both groups exhibited similar levels of visual searching that was reliably higher than after hearing the other sounds. After exiting completely, the laboratory-reared squirrels exhibited a reliably greater amount of olfactory investigation than the wild-caught squirrels. Five laboratory-reared squirrels turned around after exiting and inspected their dark nest-box opening, three of which tail flagged repeatedly and one threw substrate into the opening. Since pups recognize snakes and engage in this behavior, this latent expression of antisnake behavior illustrates its robust organizational properties in the appropriate burrow-like context irrespective of the presumed retardation of neural development known to occur in other species of rodent subjected to similar isolation rearing.

Dangerous animals capture and maintain attention in humans.

Predation is a major source of natural selection on primates and may have shaped attentional processes that allow primates to rapidly detect dangerous animals. Because ancestral humans were subjected to predation, a process that continues at very low frequencies, we examined the visual processes by which men and women detect dangerous animals (snakes and lions). We recorded the eye movements of participants as they detected images of a dangerous animal (target) among arrays of nondangerous animals (distractors) as well as detected images of a nondangerous animal (target) among arrays of dangerous animals (distractors). We found that participants were quicker to locate targets when the targets were dangerous animals compared with nondangerous animals, even when spatial frequency and luminance were controlled. The participants were slower to locate nondangerous targets because they spent more time looking at dangerous distractors, a process known as delayed disengagement, and looked at a larger number of dangerous distractors. These results indicate that dangerous animals capture and maintain attention in humans, suggesting that historical predation has shaped some facets of visual orienting and its underlying neural architecture in modern humans.

Wild Asian elephants distinguish aggressive tiger and leopard growls according to perceived danger.

Prey species exhibit antipredator behaviours such as alertness, aggression and flight, among others, in response to predators. The nature of this response is variable, with animals reacting more strongly in situations of increased vulnerability. Our research described here is the first formal study to investigate night-time antipredator behaviour in any species of elephants, Asian or African. We examined the provocative effects of elephant-triggered tiger and leopard growls while elephants attempted to crop-raid. Tigers opportunistically prey on elephant calves, whereas leopards pose no threat; therefore, we predicted that the elephant response would be reflective of this difference. Elephants reacted similarly cautiously to the simulated presence of felids of both species by eventually moving away, but differed markedly in their more immediate behavioural responses. Elephants retreated silently to tiger-growl playbacks, whereas they responded with aggressive vocalizations, such as trumpets and grunts, to leopard-growl playbacks. Elephants also lingered in the area and displayed alert or investigative behaviours in response to leopard growls when compared with tiger growls. We anticipate that the methods outlined here will promote further study of elephant antipredator behaviour in a naturalistic context, with applications for conservation efforts as well.

A comparison of rural and urban Indian children's visual detection of threatening and nonthreatening animals.

Recent studies indicate that young children preferentially attend to snakes, spiders, and lions compared with nondangerous species, but these results have yet to be replicated in populations that actually experience dangerous animals in nature. This multi-site study investigated the visual-detection biases of southern Indian children towards two potentially dangerous taxa, snakes and lions, that constituted major threats during human evolution. Three- to 8-year-old children from two distinct populations were presented with visual-search tasks containing one target image embedded in matrices of eight distractor images. Children living in Bangalore city, an urban setting in which exposure to dangerous animals would only occur occasionally during family outings to zoos and forest areas, were compared with children living in and around National Parks where exposure to dangerous species is frequent. In the first two experiments, children from both locations detected snake and lion images more rapidly than nonthreatening lizard and antelope images, respectively. Neither urban nor rural children displayed a bias for detecting horses versus cows, the latter constituting a familiar animal with strong religious significance. For all three experiments, the reaction times of urban and rural children were very similar, indicating that periodic exposure to dangerous animals early in life, coupled with adult cautioning, did not facilitate better snake and lion detection. This consistency of urban and rural children with different exposure to dangerous animals suggests that detection of some dangerous species may reflect both experience in nature and visual biases shaped by natural selection.

Development of snake-directed antipredator behavior by wild white-faced capuchin monkeys: I. Snake-species discrimination.

Young animals are known to direct alarm calls at a wider range of species than adults. Our field study examined age-related differences in the snake-directed antipredator behavior of infant, juvenile, and adult white-faced capuchin monkeys (Cebus capucinus) in terms of alarm calling, looking behavior, and aggressive behavior. In the first experiment, we exposed infant and juvenile white-faced capuchins to realistic-looking inflatable models of their two snake predators, the boa constrictior (Boa constrictor) and neotropical rattlesnake (Crotalus durissus) and a white airplane as a novel control. In the second experiment, infants, juveniles, and adults were presented photographic models of a coiled boa constrictor, rattlesnake, indigo snake (Drymarchon corais), a noncapuchin predator, and a white snake-like model. We found that antipredator behavior changed during the immature stage. Infants as young as 4 months old were able to recognize snakes and display antipredator behavior, but engaged in less snake-model discrimination than juveniles. All age classes exhibited a lower response to the white snake-like model, indicating that the absence of color and snake-scale patterns affected snake recognition. Infants also showed a higher level of vigilance after snake-model detection as exhibited by a higher proportion of time spent looking and head cocking at the models. Aggressive antipredator behavior was found in all age classes, but was more prevalent in juveniles and adults than infants. This study adds to the knowledge of development of antipredator behavior in primates by showing that, although alarm calling behavior and predator recognition appear at a very young age in capuchins, snake-species discrimination does not become apparent until the juvenile stage.

Rapid detection of visually provocative animals by preschool children and adults.

The ability to detect dangerous animals rapidly in complex landscapes has been historically important during human evolution. Previous research has shown that snake images are more readily detected than images of benign animals. To provide a stringent test of superior snake detection in preschool children and adults, Experiment 1 consisted of two parts using a touch-screen visual search task. Reaction times to detect different target snakes embedded in matrices of lizards were compared with reaction times to detect target lizards embedded in matrices of snakes. Experiment 2 compared the visual salience of lions with that of similarly colored antelopes. This experiment tested the prediction that historically dangerous felid predators would also engender rapid detection. Results from the two experiments revealed that both preschool children and adults located snakes and lions more quickly than their nonthreatening counterparts. Experiment 3 examined the ability of children and adults to distinguish between similar appearing cows and horses. Preschool children and adult men exhibited no reliable differences in detecting the two animal types. Adult women located horses reliably faster than cows, suggesting that visual biases for some animals can be acquired after childhood.

Development of snake-directed antipredator behavior by wild white-faced capuchin monkeys: II. Influence of the social environment.

Young animals are known to direct alarm calls at a wider range of animals than adults. If social cues are safer and/or more reliable to use than asocial cues for learning about predators, then it is expected that the development of this behavior will be affected by the social environment. Our study examined the influence of the social environment on antipredator behavior in infant, juvenile, and adult wild white-faced capuchin monkeys (Cebus capucinus) at Lomas Barbudal Biological Reserve in Costa Rica during presentations of different species of model snakes and novel models. We examined (a) the alarm calling behavior of the focal animal when alone versus in the vicinity of conspecific alarm callers and (b) the latency of conspecifics to alarm call once the focal animal alarm called. Focal animals alarm called more when alone than after hearing a conspecific alarm call. No reliable differences were found in the latencies of conspecifics to alarm call based on age or model type. Conspecifics were more likely to alarm call when focal individuals alarm called at snake models than when they alarm called at novel models. Results indicate (a) that alarm calling may serve to attract others to the predator's location and (b) that learning about specific predators may begin with a generalized response to a wide variety of species, including some nonthreatening ones, that is winnowed down via Pavlovian conditioned inhibition into a response directed toward specific dangerous species. This study reveals that conspecifics play a role in the development of antipredator behavior in white-faced capuchins.

Rock squirrel (Spermophilus variegatus) blood sera affects proteolytic and hemolytic activities of rattlesnake venoms.

Rock squirrels (Spermophilus variegatus) from two sites in south central New Mexico, where prairie (Crotalus viridis viridis) and western diamondback (Crotalus atrox) rattlesnakes are common predators, were assayed for inhibition of rattlesnake venom digestive and hemostatic activities. At statistically significant levels rock squirrel blood sera reduced the metalloprotease and hemolytic activity of venoms from C. v. viridis and C. atrox more than venom from an allopatric snake species, the northern Pacific rattlesnake (Crotalus oreganus). In contrast, general proteolytic activity of venom from C. oreganus was inhibited more by S. variegatus serum defenses than activity of venom from sympatric snakes. For all three venoms, incubation with squirrel sera increased the level of fibrinolysis over venom-only treatments. These results suggest that rock squirrels (S. variegatus) can defend against metalloproteases and other proteases after envenomation from at least two of five rattlesnake predators they might encounter. However, there were statistically significant differences between general proteolytic activity and fibrinolytic activity of C. v. viridis and C. atrox venom, suggesting that rock squirrels might be differentially vulnerable to these two predators. The hypothesis that prey resistance influences snake venom evolution in a predator-prey arms race is given further support by the previously cryptic variation in venoms detected when assayed against prey defenses.

Pseudoreplication conventions are testable hypotheses.

Hurlbert's conceptions of pseudoreplication, such as the loss of independent replicates with repeated sampling over time and the lack of appropriate spatial interspersion of experimental units to achieve statistical independence, are really theoretical hypotheses that warrant empirical confirmation or disconfirmation. Schank and Koehnle have provided a valuable service to researchers in ecology, conservation biology, and animal behavior using logical argument and repeated Monte Carlo simulations to challenge Hurlbert's theoretical assumptions about statistical independence. Their simulations showed that averaging samples to produce a single datum destroys the information within experimental units and is mathematically equivalent to the treatment level in a nested analysis of variance. Such results counter Hurlbert's argument that pooling is wrong. Rigid thinking about sampling, as engendered by Hurlbert's pseudoreplication arguments, does not promote the exploration of new methodologies and there is something disturbingly inconsistent with the scientific method when journal editors and reviewers adopt Hurlbert's pseudoreplication arguments unquestionably.

Relaxed selection in the wild.

Natural populations often experience the weakening or removal of a source of selection that had been important in the maintenance of one or more traits. Here we refer to these situations as 'relaxed selection,' and review recent studies that explore the effects of such changes on traits in their ecological contexts. In a few systems, such as the loss of armor in stickleback, the genetic, developmental and ecological bases of trait evolution are being discovered. These results yield insights into whether and how fast a trait is reduced or lost under relaxed selection. We provide a prospectus and a framework for understanding relaxed selection and trait loss in natural populations. We also examine its implications for applied issues, such as antibiotic resistance and the success of invasive species.

The re-emergence of felid camouflage with the decay of predator recognition in deer under relaxed selection.

When a previously common predator disappears owing to local extinction, the strong source of natural selection on prey to visually recognize that predator becomes relaxed. At present, we do not know the extent to which recognition of a specific predator is generalized to similar looking predators or how a specific predator-recognition cue, such as coat pattern, degrades under prolonged relaxed selection. Using predator models, we show that deer exhibit a more rapid and stronger antipredator response to their current predator, the puma, than to a leopard displaying primitive rosettes similar to a locally extinct predator, an early jaguar. Presentation of a novel tiger with a striped coat engendered an intermediate speed of predator recognition and strength of antipredator behaviour. Responses to the leopard model slightly exceeded responses to a non-threatening deer model, suggesting that thousands of years of relaxed selection have led to the loss of recognition of the spotted coat as a jaguar-recognition cue, and that the spotted coat has regained its ability to camouflage the felid form. Our results shed light on the evolutionary arms race between adoption of camouflage to facilitate hunting and the ability of prey to quickly recognize predators by their formerly camouflaging patterns.

California ground squirrel (Spermophilus beecheyi) defenses against rattlesnake venom digestive and hemostatic toxins.

Previous studies have shown that some mammals are able to neutralize venom from snake predators. California ground squirrels (Spermophilus beecheyi) show variation among populations in their ability to bind venom and minimize damage from northern Pacific rattlesnakes (Crotalus oreganus), but the venom toxins targeted by resistance have not been investigated. Four California ground squirrel populations, selected for differences in local density or type of rattlesnake predators, were assayed for their ability to neutralize digestive and hemostatic effects of venom from three rattlesnake species. In Douglas ground squirrels (S. b. douglasii), we found that animals from a location where snakes are common showed greater inhibition of venom metalloprotease and hemolytic activity than animals from a location where snakes are rare. Effects on general proteolysis were not different. Douglas ground squirrels also reduced the metalloprotease activity of venom from sympatric northern Pacific rattlesnakes (C. o. oreganus) more than the activity of venom from allopatric western diamondback rattlesnakes (C. atrox), but enhanced the fibrinolysis of sympatric venom almost 1.8 times above baseline levels. Two Beechey ground squirrel (S. b. beecheyi) populations had similar inhibition of venoms from northern and southern Pacific rattlesnakes (C. o. helleri), despite differences between the populations in the locally prevalent predator. However, the venom toxins inhibited by Beechey squirrels varied among venom from Pacific rattlesnake subspecies, and between these venoms and venom from allopatric western diamondback rattlesnakes. Blood plasma from Beechey squirrels showed highest inhibition of metalloprotease activity of northern Pacific rattlesnake venom, general proteolytic activity and hemolysis of southern Pacific rattlesnake venom, and hemolysis by allopatric western diamondback venom. These results reveal previously cryptic variation in venom activity against resistant prey that suggests reciprocal adaptation at the molecular level.

Orbital frontal cortex ablations of rock squirrels (Spermophilus variegatus) disinhibit innate antisnake behavior.

The antisnake behavior of rock squirrels (Spermophilus variegatus) was examined to determine the role of the orbital frontal cortex in regulating physiological arousal and behavioral excitability during encounters with a rattlesnake predator. Rock squirrels with orbital frontal cortex ablations and sham-surgery control squirrels were presented with a caged rattlesnake pre- and postsurgery. Orbital frontal cortex ablations had no substantial effect on the expression of gross motor behavior in dealing with the rattlesnake, but they augmented the speed of snake recognition and clearly disinhibited sympathetic arousal as manifested by increased tail piloerection and tail-flagging activity, which is a specific antisnake behavior. Natural selection from snakes has probably shaped the neural organization of the orbital frontal cortex to afford adaptive behavioral flexibility during snake encounters.

California ground squirrel (Spermophilus beecheyi) defenses against rattlesnake venom digestive and hemostatic toxins.

Previous studies have shown that some mammals are able to neutralize venom from snake predators. California ground squirrels (Spermophilus beecheyi) show variation among populations in their ability to bind venom and minimize damage from northern Pacific rattlesnakes (Crotalus oreganus), but the venom toxins targeted by resistance have not been investigated. Four California ground squirrel populations, selected for differences in local density or type of rattlesnake predators, were assayed for their ability to neutralize digestive and hemostatic effects of venom from three rattlesnake species. In Douglas ground squirrels (S. b. douglasii), we found that animals from a location where snakes are common showed greater inhibition of venom metalloprotease and hemolytic activity than animals from a location where snakes are rare. Effects on general proteolysis were not different. Douglas ground squirrels also reduced the metalloprotease activity of venom from sympatric northern Pacific rattlesnakes (Crotalus oreganus oreganus) more than the activity of venom from allopatric western diamondback rattlesnakes (C. atrox), but enhanced fibrinolysis of sympatric venom almost 1.8 times above baseline levels. Two Beechey ground squirrel (S. b. beecheyi) populations had similar inhibition of venoms from northern and southern Pacific rattlesnakes (C. o. helleri), despite differences between the populations in the locally prevalent predator. However, the venom toxins inhibited by Beechey squirrels did vary among venom from Pacific rattlesnake subspecies, and between these venoms and venom from allopatric western diamondback rattlesnakes. Blood plasma from Beechey squirrels showed highest inhibition of metalloprotease activity of northern Pacific rattlesnake venom, general proteolytic activity and hemolysis of southern Pacific rattlesnake venom, and hemolysis by allopatric western diamondback venom. These results reveal previously cryptic variation in venom activity against resistant prey that suggests reciprocal adaptation at the molecular level.

Recognition of partially concealed leopards by wild bonnet macaques (Macaca radiata). The role of the spotted coat.

Wild bonnet macaques (Macaca radiata) have been shown to recognize models of leopards (Panthera pardus), based on their configuration and spotted yellow coat. This study examined whether bonnet macaques could recognize the spotted and dark melanic morph when partially concealed by vegetation. Seven troops were studied at two sites in southern India, the Mudumalai Wildlife Sanctuary and the Kalakad-Mundanthurai Tiger Reserve. The forequarters and hindquarters of the two leopard morphs were presented from behind thick vegetation to individuals at feeding stations 25 m away. Flight reaction times and frequency of flight were obtained from video for only those individuals who oriented towards the models prior to hearing alarm calls. Bonnet macaques exhibited faster reaction times and greater frequency of flight after looking at the spotted morph's forequarter than after looking at either its spotted hindquarter or the dark morph's forequarter. The hindquarter of the dark morph was ignored completely. Artificial neural network modeling examined the perceptual aspects of leopard face recognition and the role of spots as camouflage. When spots were integrated into the pattern recognition process via network training, these spots contributed to leopard face recognition. When networks were not trained with spots, spots did not act as camouflage by disrupting facial features.