What can whiskers tell us about mammalian evolution,behaviour, and ecology?

1. Most mammals have whiskers; however, nearly everything we know about whiskers derives from just a handful of species, including laboratory rats Rattus norvegicus and mice Mus musculus, as well as some species of pinniped and marsupial.
2. We explore the extent to which the knowledge of the whisker system from a handful of species applies to mammals generally. This will help us understand whisker evolution and function, in order to gain more insights into mammalian behaviour and ecology.
3. This review is structured around Tinbergen’s four questions, since this method is an established, comprehensive, and logical approach to studying behaviour. We ask: how do whiskers work, develop, and evolve? And what are they for?
4. While whiskers are all slender, curved, tapered, keratinised hairs that transmit vibrotactile information, we show that there are marked differences between species with respect to whisker arrangement, numbers, length, musculature, development, and growth cycles.
5. The conservation of form and a common muscle architecture in mammals suggests that early mammals had whiskers. Whiskers may have been functional even in therapsids.
6. However, certain extant mammalian species are equipped with especially long and sensitive whiskers, in particular nocturnal, arboreal species, and aquatic species, which live in complex environments and hunt moving prey.
7. Knowledge of whiskers and whisker use can guide us in developing conservation protocols and designing enriched enclosures for captive mammals.
8. We suggest that further comparative studies, embracing a wider variety of mammalian species, are required before one can make large-scale predictions relating to evolution and function of whiskers. More research is needed to develop robust techniques to enhance the welfare and conservation of mammals.

     

Tactile discrimination of textures by Florida manatees (Trichechus manatus latirostris)

Two male Florida manatees (Trichechus manatus latirostris) demonstrated sensitive tactile discrimination in a two‐alternative forced choice task, using a modified staircase method. Stimuli were acrylic plates with vertical gratings of ridges and grooves. The standard stimulus, present on every trial, had 2 mm gratings and the comparison stimuli had wider gratings. The blindfolded subjects were trained to demonstrate discrimination by pressing the target with wider gratings. Discrimination thresholds (75% correct) for the subjects were 2.05 mm and 2.15 mm, corresponding to Weber fractions of 0.025 and 0.075, respectively. These results indicate thresholds on similar stimuli comparable to humans (index finger tasks) and better than harbor seals, Phoca vitulina, and the closely related Antillean manatee, Trichechus manatus manatus. Memory for the tactile task was quite stable for both subjects, over 2 yr in the case of one of the subjects. Video analysis of responses indicated that bristle‐like hairs, perioral bristles, and skin on the oral disk were involved in the discrimination response.     

Ecomorphology of a size-structured tropical freshwater fish community

Among nine species of a tropical community ecomorphological correlates were sought throughout ontogeny. Ontogenetic changes were distinguished by establishing six pre-defined size-classes. Morphometric data associated with feeding were compared by canonical correspondence analysis to dietary data. This analysis revealed seven significant relationships, showing 71% of the morphological variance explained 77% of the variance in diet. Based on funtional ecomorphological relationships established in other studies and results of the canonical correspondence analysis, three food characters were selected: the size of the food particles, the type of food (vegetable versus animal) and the vertical position in the water column. The morphometric data were reduced using principal component analysis into three axis explaining 83% of the variation. The morphological characters with the highest loadings were: mouth gape on the first principal component axis, length of the intestine tract on the second and the orientation of the mouth together with the presence of barbels on the third. These axis were significantly correlated with, respectively, the size, the type and the vertical position of the food. The importance of morphological changes during ontogeny in explaining dietary changes was shown because 75% of the variation in the first, and most important, morphological principal component was accounted for by differences between size-classes. Assuming functional relationships, the potential niches of the species/size-classes were established, distinguishing herbivorous, omnivorous/molluscivorous and carnivorous species which, in turn, were segregated by their potential to feed on larger prey. This revised version was published online in July 2006 with corrections to the Cover Date.     

Ecomorphology of retinal structures in zooplanktivorous haplochromine cichlids (Pisces) from Lake Victoria

Synopsis Before the decline of the species flock of haplochromine cichlids of Lake Victoria due to the Nile perch upsurge, there were many co-existing haploehromine species such as the taxonomically and ecologically well-studied zooplanktivores of the Mwanza Gulf. In spite of the scarcely separated niches of some of these species, no sign of competition for space or food could be demonstrated. As is argued in this paper, optical differentiation could well be an aspect of adaptive radiation of these zooplanktivores, particularly among the highly sympatric species. Our hypothesis is based on the morphological modifications of retinal structures in nine zooplanktivorous species. Interspecific variation was observed in composition, size and density of the photoreceptors and ganglion cells. The analyses included the intraretinal variation and size dependency of some of the structural parameters. The optical functions deduced from retinal structure indicate distinct interspecific differences in sensitivity thresholds and a slight differentiation in visual resolution. These functions correlate poorly with the photic conditions of the species-specific habitats. The optical properties can, on the other hand, be connected with the more subtle differentiation in food items and feeding behaviour among these species. It is our concluding hypothesis, that the optical differentiation among the haplochromine zooplanktivores primarily served resource partitioning by different modes of visual prey detection rather than niche partitioning by habitat.     

Ecomorphology of plesiosaur flipper geometry

The Plesiosauria is an extinct group of marine reptiles once common in mesozoic seas. Previous work on plesiosaur hunting styles has suggested that short‐necked, large‐headed animals were pursuit predators, whereas long‐necked, small‐headed animals were ambush predators. This study presents new data on the aspect ratios (ARs) of plesiosaur flippers, and interprets these data via comparison with AR in birds, bats and aircraft. Performance trade‐offs implicit in AR variation are well‐understood in the context of aircraft design, and these trade‐offs have direct ecomorphological analogues in birds and bats. Knowledge of these trade‐offs allows interpretation of variation in plesiosaur AR. By analogy, short‐necked taxa were specialized for manoeuvrability and pursuit, whereas long‐necked taxa were generally specialized for efficiency and cruising. These interpretations agree with previous assessments of maximum swimming speed.     

Ecomorphology of the digestive tract of the brazilian electric ray Narcine brasiliensis (Olfers, 1831) (Torpediniformes: Narcinidae)

Elasmobranchii shows particular characteristics that make them susceptible to the impacts caused by fishing pressure, especially bottom trawling, which mostly affect rays, as they have demersal habits. Based on food availability and feeding ability, Polychaeta and Sipunculiforme are the bases of Brazilian electric ray Narcine brasiliensis diet. Morphologically, the digestion starts in the oesophagus with acid mucines action. The existence of acid and neutral mucines in the posterior intestine prevents lesions in the mucosa and favours the passage of faeces and shell residues. Thus, this study shows the interaction between their habitat, the organisms and their morphophysiology.     

Ecomorphology of Locomotion in Labrid Fishes

The Labridae is an ecologically diverse group of mostly reef associated marine fishes that swim primarily by oscillating their pectoral fins. To generate locomotor thrust, labrids employ the paired pectoral fins in motions that range from a fore-aft rowing stroke to a dorso-ventral flapping stroke. Species that emphasize one or the other behavior are expected to benefit from alternative fin shapes that maximize performance of their primary swimming behavior. We document the diversity of pectoral fin shape in 143 species of labrids from the Great Barrier Reef and the Caribbean. Pectoral fin aspect ratio ranged among species from 1.12 to 4.48 and showed a distribution with two peaks at about 2.0 and 3.0. Higher aspect ratio fins typically had a relatively long leading edge and were narrower distally. Body mass only explained 3% of the variation in fin aspect ratio in spite of four orders of magnitude range and an expectation that the advantages of high aspect ratio fins and flapping motion are greatest at large body sizes. Aspect ratio was correlated with the angle of attachment of the fin on the body (r = 0.65), indicating that the orientation of the pectoral girdle is rotated in high aspect ratio species to enable them to move their fin in a flapping motion. Field measures of routine swimming speed were made in 43 species from the Great Barrier Reef. Multiple regression revealed that fin aspect ratio explained 52% of the variation in size-corrected swimming speed, but the angle of attachment of the pectoral fin only explained an additional 2%. Labrid locomotor diversity appears to be related to a trade-off between efficiency of fast swimming and maneuverability in slow swimming species. Slow swimmers typically swim closer to the reef while fast swimmers dominate the water column and shallow, high-flow habitats. Planktivory was the most common trophic associate with high aspect ratio fins and fast swimming, apparently evolving six times.     

Magnitude Estimation of Contact Force When Objects with Different Shapes Are Applied Passively to the Fingerpad

Stimuli with spherically curved surfaces were presented passively to the fingerpads of human subjects. There were 28 stimuli, consisting of all combinations of 4 different curvatures and 7 different contact forces; these were presented in random order. Subjects scaled their perceived magnitude of the contact force using magnitude estimation. Perceived force increased markedly with an increase in experimentally applied contact force. An increase in curvature resulted in a slight increase in perceived contact force. Thus, when humans are passively presented with objects changing in both shape and contact force, they are able to extract information about the force. Because of the passive nature of the task, all such information must be conveyed to the brain by the cutaneous mechanoreceptors.     

Ecomorphology and habitat utilization of Cottus species

We investigated relationships between morphological attributes and mesohabitat use in populations of Cottus carolinae. Field studies revealed significant differences between Ozark and prairie stream mesohabitats where populations of C. carolinae occurred. Features of the mesohabitat that were characteristic of each stream type correlated with intra- and inter-specific differences in morphological attributes of Cottus. The morphology of Cottus occurring in the deeper, slower, and siltier prairie stream exhibited larger head width and depth, body width and depth, and caudal peduncle width while those occurring in a shallower, faster flowing Ozark stream had longer pelvic fins, larger eyes, and deeper caudal peduncles. Each population had a morphology tailored to optimize its performance in a specific habitat. Results of laboratory based performance studies using a variable flow chamber revealed that critical current velocities (CCV) of live and preserved specimens were weakly associated with distinct combinations of morphological traits. Specimens with higher CCV measurements typically had longer pelvic fins, larger eyes, and deeper caudal fins (Ozark morphology). Those with lower CCVs had larger head width and depth, body width and depth, and caudal peduncle width (prairie morphology) in studies using cobble as the substrate. Differences in flow regimes between prairie and Ozark streams may generate specific niches for specific morphotypes, and current velocity may exert selective pressure on morphology.     

Ecomorphology and foraging behaviour of Pacific boobies

Wing size and shape, expressed as wing loading and aspect ratio respectively, together with bill morphology are parameters that can reveal differences related to the foraging ecology of seabirds. Six species of booby (Sulidae) that inhabit the Pacific are the focus of this study: four mainly pelagic species, Masked Booby Sula dactylatra, Nazca Booby Sula granti, Red‐footed Booby Sula sula and Brown Booby Sula leucogaster, and two coastal species, Blue‐footed Booby Sula nebouxii and Peruvian Booby Sula variegata. Pelagic boobies showed segregation among species in body mass and relative bill size, and they differed in wing morphology (wing loading and aspect ratio) from the coastal boobies. The coastal Peruvian and Blue‐footed Boobies are largely allopatric but overlap in northern Peru. In their area of sympatry, they showed evidence of character displacement in body size and in wing and bill morphology, which suggests that competition plays an important role in sympatry. This study improves our understanding of ecological interactions among Pacific boobies and of how selective pressures have shaped their ecomorphology and foraging behaviours.     

Reconstitution reveals the functional core of mammalian eIF3

Eukaryotic translation initiation factor (eIF)3 is the largest eIF ( approximately 650 kDa), consisting of 10-13 different polypeptide subunits in mammalian cells. To understand the role of each subunit, we successfully reconstituted a human eIF3 complex consisting of 11 subunits that promoted the recruitment of the 40S ribosomal subunit to mRNA. Strikingly, the eIF3g and eIF3i subunits, which are evolutionarily conserved between human and the yeast Saccharomyces cerevisiae are dispensable for active mammalian eIF3 complex formation. Extensive deletion analyses suggest that three evolutionarily conserved subunits (eIF3a, eIF3b, and eIF3c) and three non-conserved subunits (eIF3e, eIF3f, and eIF3h) comprise the functional core of mammalian eIF3.     

Sensory biology of Phalangida harvestmen (Arachnida,Opiliones): a review,with new morphological data on 18 species

Phalangida includes three of the four suborders of Opiliones (Arachnida): Eupnoi, Dyspnoi and Laniatores. We review the literature on the sensory structures and capabilities of Phalangida, provide new morphological data for 18 species and discuss the 11 sensory structures that have been described in the group. Based on the published data encompassing both behaviour and morphology, three conclusions are apparent: (1) species of Phalangida appear to have limited abilities to detect stimuli at a distance; (2) close range olfaction probably helps to find foods with strong odours, but (3) they appear to be highly dependent on contact chemoreception to detect live prey, predators and mates. We also highlight the fact that legs I in the three suborders and pedipalps in Dyspnoi and Eupnoi are very important sensory appendages, thus legs II should not be called the ‘sensory appendages’ of harvestmen. In conclusion, we highlight the fact that the sensory capabilities, diet, prey capturing and handling ability, and foraging behaviour of species of Phalangida seem to be different from those of most other arachnids. Finally, we suggest future directions for studies in the field of the sensory system of the group.     

Ecomorphology of the giant bear-dogs Amphicyon and Ischyrocyon

Giant bear-dogs of the genera Amphicyon and Ischyrocyon (Carnivora, Amphicyonidae, Amphicyoninae) were the largest carnivorans in North America during middle and late Miocene (17.5–8.8 Mya) with a dental and skeletal morphology that combined features found in living Ursidae, Canidae, and Felidae. This study tests previously proposed models of diet and hunting behaviour of these extinct carnivorans. Relative grinding area (RGA) of lower molars and wear pattern on upper molars suggest that bear-dogs were carnivorous. Amphicyon and Ischyrocyon possessed skeletal features of both ambush (short distal limb segments) and pursuit (caudally bent olecranon process of ulna) living predators. Therefore, bear-dogs probably pursued their prey (mediportal ungulates) for a longer distance but at a slower speed than do living ambush predators. Upon catching up to its prey a bear-dog probably seized it with powerfully muscled forelimbs and killed it by tearing into its ribcage or neck with canines set in a narrow rostrum.     

Intralocality Variation in Feeding Biomechanics and Prey Use in Archosargus probatocephalus (Teleostei,Sparidae), with Implications for the Ecomorphology of Fishes

Archosargus probatocephalusin a Florida estuary was investigated to explore intraspecific variation in prey utilization and jaw biomechanics. Volumetric contribution of major prey types and seven biomechanical features of the oral jaws that characterize prey-capture and processing performance were contrasted between two locations within the estuary. At Mosquito Lagoon, where A. probatocephalusinhabited mostly oyster beds, mangroves and salt marshes, fish consumed mostly thick-shelled bivalves, gastropods, crabs, and tubiculous polychaetes and amphipods. In contrast, conspecifics at Indian River Lagoon that inhabited mostly seagrass beds and algal turf consumed predominantly algae, seagrass, epiphytic invertebrates and small bivalves and gastropods. Difference in magnitude of durophagy between locations was associated with differences in oral-jaw biomechanics. Analyses of covariance indicated that A. probatocephalusat Mosquito Lagoon had more massive jaw muscles and bones, than conspecifics at Indian River Lagoon. Variations in lever ratios for jaw-opening and jaw-closing between locations were not significant. It is hypothesized that intralocality differences in food habits have induced the development of feeding morphologies that enhance the ability of A. probatocephalusto successfully exploit locally dominant prey resources within the estuary. Plasticity of the feeding mechanism of A. probatocephalusmay buffer the species from the adverse effects of settling on heterogeneous habitats that contain variable prey resources such as those found within estuaries.     

Network scaling reveals consistent fractal pattern in hierarchical mammalian societies

Recent studies have demonstrated that human societies are hierarchically structured with a consistent scaling ratio across successive layers of the social network; each layer of the network is between three and four times the size of the preceding (smaller) grouping level. Here we show that similar relationships hold for four mammalian taxa living in multi-level social systems. For elephant (Loxodonta africana), gelada (Theropithecus gelada) and hamadryas (Papio hamadryas hamadryas) baboon, successive layers of social organization have a scaling ratio of almost exactly 3, indicating that such branching ratios may be a consistent feature of all hierarchically structured societies. Interestingly, the scaling ratio for orca (Orcinus orca) was 3.8, which might mean that aquatic environments place different constraints on the organization of social hierarchies. However, circumstantial evidence from a range of other species suggests that scaling ratios close to 3 may apply widely, even in species where hierarchical social structures have not traditionally been identified. These results identify the origin of the hierarchical, fractal-like organization of mammalian social systems as a fundamental question.     

Melanopsin-dependent photo-perturbation reveals desynchronization underlying the singularity of mammalian circadian clocks

Singularity behaviour in circadian clocks--the loss of robust circadian rhythms following exposure to a stimulus such as a pulse of bright light--is one of the fundamental but mysterious properties of clocks. To quantitatively perturb and accurately measure the dynamics of cellular clocks, we synthetically produced photo-responsiveness within mammalian cells by exogenously introducing the photoreceptor melanopsin and continuously monitoring the effect of photo-perturbation on the state of cellular clocks. Here we report that a critical light pulse drives cellular clocks into singularity behaviour. Our theoretical analysis consistently predicts and subsequent single-cell level observation directly proves that desynchronization of individual cellular clocks underlies singularity behaviour. Our theoretical framework also explains why singularity behaviours have been experimentally observed in various organisms, and it suggests that desynchronization is a plausible mechanism for the observable singularity of circadian clocks. Importantly, these in vitro and in silico findings are further supported by in vivo observations that desynchronization underlies the multicell-level amplitude decrease in the rat suprachiasmatic nucleus induced by critical light pulses.