Animals evoking fear in the Cradle of Humankind: snakes, scorpions, and large carnivores.

Theories explain the presence of fears and specific phobias elicited by animals in contemporary WEIRD (Western, educated, industrialized, rich, and democratic) populations by their evolutionary past in Africa. Nevertheless, empirical data about fears of animals in the Cradle of Humankind are still fragmentary. To fill this gap, we examined which local animals are perceived as the most frightening by Somali people, who inhabit a markedly similar environment and the region where humans have evolved. We asked 236 raters to rank 42 stimuli according to their elicited fear. The stimuli were standardized pictures of species representing the local fauna. The results showed that the most frightening animals were snakes, scorpions, the centipede, and large carnivores (cheetahs and hyenas). These were followed up by lizards and spiders. Unlike in Europe, spiders represent less salient stimuli than scorpions for Somali respondents in this study. This conforms to the hypothesis suggesting that fear of spiders was extended or redirected from other chelicerates.

Determinate growth is predominant and likely ancestral in squamate reptiles.

Body growth is typically thought to be indeterminate in ectothermic vertebrates. Indeed, until recently, this growth pattern was considered to be ubiquitous in ectotherms. Our recent observations of a complete growth plate cartilage (GPC) resorption, a reliable indicator of arrested skeletal growth, in many species of lizards clearly reject the ubiquity of indeterminate growth in reptiles and raise the question about the ancestral state of the growth pattern. Using X-ray micro-computed tomography (µCT), here we examined GPCs of long bones in three basally branching clades of squamate reptiles, namely in Gekkota, Scincoidea and Lacertoidea. A complete loss of GPC, indicating skeletal growth arrest, was the predominant finding. Using a dataset of 164 species representing all major clades of lizards and the tuataras, we traced the evolution of determinate growth on the phylogenetic tree of Lepidosauria. The reconstruction of character states suggests that determinate growth is ancestral for the squamate reptiles (Squamata) and remains common in the majority of lizard lineages, while extended (potentially indeterminate) adult growth evolved several times within squamates. Although traditionally associated with endotherms, determinate growth is coupled with ectothermy in this lineage. These findings combined with existing literature suggest that determinate growth predominates in both extant and extinct amniotes.

Comparing developmental stability in unisexual and bisexual rock lizards of the genus Darevskia.

Parthenogenetic species are usually considered to be short-lived due to the accumulation of adverse mutations, lack of genetic variability, and inability to adapt to changing environment. If so, one may expect that the phenotype of clonal organisms may reflect such genetic and/or environmental stress. To test this hypothesis, we compared the developmental stability of bisexual and parthenogenetic lizards of the genus Darevskia. We assessed asymmetries in three meristic traits: ventral, preanal, and supratemporal scales. Our results suggest that the amount of ventral and preanal asymmetries is significantly higher in clones compared with their maternal, but not paternal, progenitor species. However, it is questionable, whether this is a consequence of clonality, as it may be considered a mild form of outbreeding depression as well. Moreover, most ventral asymmetries were found in the bisexual species Darevskia valentini. We suggest that greater differences in asymmetry levels among bisexuals may be, for instance, a consequence of the population size: the smaller the population, the higher the inbreeding and the developmental instability. On the basis of the traits examined in this study, the parthenogens do not seem to be of significantly poorer quality.

A gyroscopic advantage: phylogenetic patterns of compensatory movements in frogs.

Head and eye compensatory movements known as vestibulo-ocular and vestibulo-cervical reflexes are essential to stay orientated in space while moving. We have used a previously developed methodology focused on the detailed mathematical description of head compensatory movements in frogs without the need for any surgical procedures on the examined specimens. Our comparative study comprising 35 species of frogs from different phylogenetic backgrounds revealed species-specific head compensatory abilities ensuring gaze stabilization. Moreover, we found a strong phylogenetic signal highlighting the great ability of compensatory head movements in families of Pyxicephalidae and Rhacophoridae from the Natatanura group. By contrast, families of Dendrobatidae and Microhylidae exhibited only poor or no head compensatory movements. Contrary to our expectation, the results did not corroborate an ecomorphological hypothesis anticipating a close relationship between ecological parameters and the head compensatory movements. We did not find any positive association between more complex (3D structured, arboreal or aquatic) habitats or more saltatory behavior and elevated abilities of head compensatory movements. Moreover, we found compensatory movements in most basal Archeobatrachia, giving an indication of common ancestry of these abilities in frogs that are variously pronounced in particular families. We hypothesize that the uncovered proper gaze stabilization during locomotion provided by the higher head compensatory abilities can improve or even enable visual perception of the prey. We interpret this completely novel finding as a possible gyroscopic advantage in a foraging context. We discuss putative consequences of such advanced neuromotor skills for diversification and ecological success of the Natatanura group.

Strong support for Rensch's rule in an American clade of lizards (Teiidae and Gymnophtalmidae) and a paradox of the largest tejus.

Rensch's rule predicts an allometric relationship between male and female body size stating that the sexual size dimorphism (SSD) increases with body size in male-larger taxa and decreases in female-larger taxa in groups of related species. It means that the relationship between the male and female body size is hyperallometric, i.e., the allometric exponent of this relationship exceeds the unity. We explored the relationship between the male and female body size in a New World clade of lizards consisting of sister families Teiidae and Gymnophthalmidae, which exhibit a great variation in both their adult body sizes and SSD. All our estimates of the reduced major axis regression slopes ranged from 1.067 to 1.229 and clearly followed a pattern consistent with the Rensch's rule. Despite a clear general trend, giant species from the subfamily Tupinambinae show paradoxically only poor SSD. The cases of extreme male-larger SSD were found in species of moderate body size belonging to the genera Ameiva and Cnemidophorus. The abovementioned deviations from the hyperallometric relationship between the male and female body size are surprising and require further examination.

The first description of dermal armour in snakes.

Osteoderms, also called dermal armour, often play a role in predator defence. The presence of osteoderms is highly irregularly distributed across the squamate phylogeny and they have not been found in snakes. In this study, we searched for candidate snake species that would benefit from such armour to protect their body, focusing primarily on fossorial species with defensive tail displays. We examined the tail morphology of 27 snake species from different families using micro-computed tomography (µCT) and micro- radiography. We discovered dermal armour in four species of sand boas (Erycidae) that also feature enlarged and highly modified caudal vertebrae. This is the first description of dermal armour in snakes. Ancestral state reconstructions revealed that osteoderms likely evolved once or multiple times in Erycidae. We have not found osteoderms in any other examined snake species. Nevertheless, similar structures are known from unrelated squamate clades, such as gerrhosaurids and geckos. This supports the idea of underlying deep developmental homology. We propose the hypothesis that osteoderms protect sand boas like the "brigandine armour" of medieval warriors. We interpret it as another component of the sand boas' rich defence strategy.

Are vipers prototypic fear-evoking snakes? A cross-cultural comparison of Somalis and Czechs.

Snakes are known as highly fear-evoking animals, eliciting preferential attention and fast detection in humans. We examined the human fear response to snakes in the context of both current and evolutionary experiences, conducting our research in the cradle of humankind, the Horn of Africa. This region is characterized by the frequent occurrence of various snake species, including deadly venomous viperids (adders) and elapids (cobras and mambas). We conducted experiments in Somaliland and compared the results with data from Czech respondents to address the still unresolved questions: To which extent is human fear of snakes affected by evolutionary or current experience and local culture? Can people of both nationalities recognize venomous snakes as a category, or are they only afraid of certain species that are most dangerous in a given area? Are respondents of both nationalities equally afraid of deadly snakes from both families (Viperidae, Elapidae)? We employed a well-established picture-sorting approach, consisting of 48 snake species belonging to four distinct groups. Our results revealed significant agreement among Somali as well as Czech respondents. We found a highly significant effect of the stimulus on perceived fear in both populations. Vipers appeared to be the most salient stimuli in both populations, as they occupied the highest positions according to the reported level of subjectively perceived fear. The position of vipers strongly contrasts with the fear ranking of deadly venomous elapids, which were in lower positions. Fear scores of vipers were significantly higher in both populations, and their best predictor was the body width of the snake. The evolutionary, cultural, and cognitive aspects of this phenomenon are discussed.

The bigger the threat, the longer the gaze? A cross-cultural study of Somalis and Czechs.

High fear reaction, preferential attention, or fast detection are only a few of the specific responses which snakes evoke in humans. Previous research has shown that these responses are shared amongst several distinct cultures suggesting the evolutionary origin of the response. However, populations from sub-Saharan Africa have been largely missing in experimental research focused on this issue. In this paper, we focus on the effect of snake threat display on human spontaneous attention. We performed an eye-tracking experiment with participants from Somaliland and the Czechia and investigated whether human attention is swayed towards snakes in a threatening posture. Seventy-one Somalis and 71 Czechs were tested; the samples were matched for gender and comparable in age structure and education level. We also investigated the effect of snake morphotype as snakes differ in their threat display. We found that snakes in a threatening posture were indeed gazed upon more than snakes in a relaxed (non-threatening) posture. Further, we found a large effect of snake morphotype as this was especially prominent in cobras, less in vipers, and mostly non-significant in other morphotypes. Finally, despite highly different cultural and environmental backgrounds, the overall pattern of reaction towards snakes was similar in Somalis and Czechs supporting the evolutionary origin of the phenomenon. We concluded that human attention is preferentially directed towards snakes, especially cobras and vipers, in threatening postures.

Do Spiders Ride on the Fear of Scorpions? A Cross-Cultural Eye Tracking Study.

Deep fear of spiders is common in many countries, yet its origin remains unexplained. In this study, we tested a hypothesis based on recent studies suggesting that fear of spiders might stem from a generalized fear of chelicerates or fear of scorpions. To this end, we conducted an eye tracking experiment using a spontaneous gaze preference paradigm, with spiders and scorpions (previously neglected but crucial stimuli) as threatening stimuli and grasshoppers as control stimuli. In total, 67 participants from Somaliland and 67 participants from the Czech Republic were recruited and presented with a sequence of paired images. Both Somali and Czech people looked longer (total duration of the gaze) and more often (number of fixations) on the threatening stimuli (spiders and scorpions) when presented with a control (grasshopper). When both threatening stimuli were presented together, Somali participants focused significantly more on the scorpion, whereas in Czech participants, the effect was less pronounced, and in Czech women it was not significant. This supports the hypothesis that fear of spiders originated as a generalized fear of scorpions. Moreover, the importance of spiders as fear-eliciting stimuli may be enhanced in the absence of scorpions in the environment.

Specificity of spiders among fear- and disgust-eliciting arthropods: Spiders are special, but phobics not so much.

To investigate a specificity of spiders as a prototypical fear- and disgust-eliciting stimuli, we conducted an online experiment. The respondents rated images of 25 spiders, 12 non-spider chelicerates, and 10 other arthropods on a fear and disgust 7-point scale. The evaluation of 968 Central European respondents confirmed the specificity of spiders among fear- and disgust-eliciting arthropods and supported the notion of spiders as a cognitive category. We delineated this category as covering extant spider species as well as some other chelicerates bearing a physical resemblance to spiders, mainly whip spiders and camel spiders. We suggested calling this category the spider-like cognitive category. We discussed evolutionary roots of the spider-like category and concluded that its roots should be sought in fear, with disgust being secondary of the two emotions. We suggested other chelicerates, e.g., scorpions, might have been important in formation and fixation of the spider-like category. Further, we investigated an effect of respondent's sensitivity to a specific fear of spiders on evaluation of the stimuli. We found that suspected phobic respondents were in their rating nearly identical to those with only high fear of spiders and similar to those with only moderate fear of spiders. We concluded that results based on healthy respondents with elevated fear should also be considered relevant for arachnophobia research.

Emotions triggered by live arthropods shed light on spider phobia.

Spiders are mostly harmless, yet they often trigger high levels of both fear and disgust, and arachnophobia (the phobia of spiders) ranks among the most common specific animal phobias. To investigate this apparent paradox, we turned to the only close relatives of spiders that pose a real danger to humans: scorpions. We adopted a unique methodology in order to assess authentic emotions elicited by arthropods. Over 300 respondents were asked to rate live specimens of 62 arthropod species (including spiders, scorpions, cockroaches, and other insects) based on perceived fear, disgust, and beauty. We found that species' scores on all three scales depended on the higher taxon as well as on body size. Spiders, scorpions, and other arachnids scored the highest in fear and disgust, while beetles and crabs scored the highest in beauty. Moreover, all chelicerates were perceived as one cohesive group, distinct from other arthropods, such as insects or crabs. Based on these results, we hypothesize that the fear of spiders might be triggered by a generalized fear of chelicerates, with scorpions being the original stimulus that signals danger.

Ontogeny of sexual size dimorphism in monitor lizards: males grow for a longer period, but not at a faster rate.

Monitor lizards belong to the largest and the most sexually dimorphic lizards in terms of size, making this group an ideal model for studies analyzing ontogenetic causes of sexual dimorphism. Understanding of these ontogenetic factors is essential to the current discussion concerning patterns of sexual dimorphism in animals. We examined the ontogenetic trajectories of body weight and snout-vent length to analyze the emergence of sexual size dimorphism. Experimental animals were 22 males and 13 females of mangrove-dwelling monitors (Varanus indicus) hatched at the Prague Zoo. They were regularly weighed and measured up to the age of 33-40 months, and subsequently sexed by ultrasonographic imaging. The logistic growth equation was used to describe and analyze the observed growth patterns. Our results confirm considerable sexual size dimorphism in the mangrove monitor. The mean asymptotic body weight of males was nearly three times higher than that of females. As the body size of male and female hatchlings is almost equal, and the growth rate parameter (K) of the logistic growth equation as well as the absolute growth rate up to the age of 12 months do not differ between the sexes, size differences between fully grown males and females should be attributed to timing of the postnatal growth. Males continue to grow several months after they reach the age when the growth of females is already reduced. Therefore, the sexual size dimorphism emerges and sharply increases at this period.

Ultraviolet reflectance and pattern properties in leopard geckos (Eublepharis macularius).

Complex visual signaling through various combinations of colors and patterns has been well documented in a number of diurnal reptiles. However, there are many nocturnal species with highly sensitive vision, being able to discriminate colors in night conditions, as was shown in geckos. Because of their sensitivity to chromatic signals, including UV (ultraviolet), they may have potential hidden features in their coloration, which may play role in intraspecific communication (e.g. mate choice) or interspecific signals (e.g. antipredatory function). We explored this hypothesis in nocturnal Leopard geckos (Eublepharis macularius), a species using visual signals in both antipredation defense and courtship, having ontogenetic color change accompanied by a shift in behavior. We used UV photography and visual modeling in order to compare various aspects of their coloration (luminance, contrast, color proportions) between sexes, age groups and populations. We found that Leopard geckos have considerable UV reflectance in white patches on their tails (and on the head in juveniles). Though, no prominent differences were detected in their coloration between various groups. We hypothesize that the limitation of UV reflectance to the head and tail, which are both actively displayed during defense, especially in juveniles, might potentially boost the effect of antipredation signaling.

Universality of indeterminate growth in lizards rejected: the micro-CT reveals contrasting timing of growth cartilage persistence in iguanas, agamas, and chameleons.

Squamate reptiles are considered to exhibit indeterminate growth. Nevertheless, current literature disputes the available definitions of this growth type, presents new theoretical models, and questions its universality in cold-blooded vertebrates. We have followed up on our previous research employing micro-CT to explore growth plate cartilage (GPC) in the epiphysis of long bones, which is responsible for longitudinal skeletal growth by the endochondral ossification process. We focused on numerous and highly diversified group of the Iguania clade comprising Acrodonta (agamas and chameleons) and Pleurodonta ("iguanas"). We recorded the absence of GPC in most of the examined adult Pleurodonta specimens and interpret it as an irreversible arrest of skeletal growth. This finding clearly rejects the universality of indeterminate growth in lizards. On the other hand, we found apparent GPC preservation in most of the adult specimens belonging to Acrodonta. This suggests a preserved ability to continue body growth throughout most of their life. We discuss the uncovered disparity between Acrodonta and Pleurodonta and emphasize the importance of GPC degradation timing.

A comparative study of growth: different body weight trajectories in three species of the genus Eublepharis and their hybrids.

An extensive research effort is devoted to the evolution of life-histories and processes underlying the variation in adult body weight; however, in this regard, some animal taxa remain neglected. Here we report rates and timing of growth recorded in two wild-derived populations of a model lizard species, Eublepharis macularius (M, W), other two related species, i.e., E. angramainyu (A) and E. sp. (D), and their between-species hybrids. We detected clear differences among the examined species/populations, which can be interpreted in the terms of "fast - slow" continuum of life-history strategies. The mean asymptotic body size was the highest in A and further decreased in the following order: M, W, and D. In contrast, the growth rate showed an opposite pattern. Counter-intuitively, the largest species exhibited the slowest growth rates. The final body size was determined mainly by the inflexion point. This parameter reflecting the duration of exponential growth increased with mean asymptotic body size and easily overcompensated the effect of decreasing growth rates in larger species. Compared to the parental species, the F1 and backcross hybrids exhibited intermediate values of growth parameters. Thus, except for the case of the F2 hybrid of MxA, we failed to detect deleterious effects of hybridization in these animals with temperature sex determination.

New records of one of the least known snakes, Telescopus pulcher (Squamata: Colubridae) from the Horn of Africa.

Telescopus pulcher is an enigmatic colubrid snake only known from the holotype and paratype specimens described from 'Migiurtinia' in Puntland (Somalia) in 1935. Herein we recorded the third and fourth-ever known specimens of this species from the Toon village, Woqooyi Galbeed Region, and 15 km southeast of Sheikh, Saahil Region, Somaliland. The species is endemic to Somaliland and adjoining parts of Ethiopia and Puntland. Data on morphology and natural history, as well as the first photographs of live specimens are provided. We also provide a detailed description of the paratype. The coloration of the species resembles that of the vipers of the genus Echis and we hypothesize that T. pulcher mimics these common and sympatric vipers in the Horn of Africa.

Development of behavioural profile in the Northern common boa (Boa imperator): Repeatable independent traits or personality?

Recent studies of animal personality have focused on its proximate causation and ecological and evolutionary significance in particular, but the question of its development was largely overlooked. The attributes of personality are defined as between-individual differences in behaviour, which are consistent over time (differential consistency) and contexts (contextual generality) and both can be affected by development. We assessed several candidates for personality variables measured in various tests with different contexts over several life-stages (juveniles, older juveniles, subadults and adults) in the Northern common boa. Variables describing foraging/feeding decision and some of the defensive behaviours expressed as individual average values are highly repeatable and consistent. We found two main personality axes-one associated with foraging/feeding and the speed of decision, the other reflecting agonistic behaviour. Intensity of behaviour in the feeding context changes during development, but the level of agonistic behaviour remains the same. The juveniles and adults have a similar personality structure, but there is a period of structural change of behaviour during the second year of life (subadults). These results require a new theoretical model to explain the selection pressures resulting in this developmental pattern of personality. We also studied the proximate factors and their relationship to behavioural characteristics. Physiological parameters (heart and breath rate stress response) measured in adults clustered with variables concerning the agonistic behavioural profile, while no relationship between the juvenile/adult body size and personality concerning feeding/foraging and the agonistic behavioural profile was found. Our study suggests that it is important for studies of personality development to focus on both the structural and differential consistency, because even though behaviour is differentially consistent, the structure can change.

Morphological characteristics of blood cells in monitor lizards: is erythrocyte size linked to actual body size?

Blood cell morphology and count are not uniform across species. Recently, between-species comparisons revealed that the size of red blood cells is associated with body size in some lizard taxa, and this finding was interpreted in the context of the metabolic theory. In the present study, we examined the numbers and the size of blood cells in 2 species of monitor lizards, the mangrove-dwelling monitor (Varanus indicus) and the savannah monitor (V. exanthematicus), and we compared these traits in individuals of different body size. The results revealed that during the course of ontogeny, the size of red blood cells increases with body mass. Because the mass-specific metabolic rate decreases with body size and the cell volume-to-surface ratio decreases with the cell size, changes in the erythrocyte size might be the result of oxygen transport adjustment.