Ancestral duplications and highly dynamic opsin gene evolution in percomorph fishes.

Single-gene and whole-genome duplications are important evolutionary mechanisms that contribute to biological diversification by launching new genetic raw material. For example, the evolution of animal vision is tightly linked to the expansion of the opsin gene family encoding light-absorbing visual pigments. In teleost fishes, the most species-rich vertebrate group, opsins are particularly diverse and key to the successful colonization of habitats ranging from the bioluminescence-biased but basically dark deep sea to clear mountain streams. In this study, we report a previously unnoticed duplication of the violet-blue short wavelength-sensitive 2 (SWS2) opsin, which coincides with the radiation of highly diverse percomorph fishes, permitting us to reinterpret the evolution of this gene family. The inspection of close to 100 fish genomes revealed that, triggered by frequent gene conversion between duplicates, the evolutionary history of SWS2 is rather complex and difficult to predict. Coincidentally, we also report potential cases of gene resurrection in vertebrate opsins, whereby pseudogenized genes were found to convert with their functional paralogs. We then identify multiple novel amino acid substitutions that are likely to have contributed to the adaptive differentiation between SWS2 copies. Finally, using the dusky dottyback Pseudochromis fuscus, we show that the newly discovered SWS2A duplicates can contribute to visual adaptation in two ways: by gaining sensitivities to different wavelengths of light and by being differentially expressed between ontogenetic stages. Thus, our study highlights the importance of comparative approaches in gaining a comprehensive view of the dynamics underlying gene family evolution and ultimately, animal diversification.

Comparison of functional and anatomical estimations of visual acuity in two species of coral reef fish.

The high-contrast, complex patterns typical of many reef fish serve several purposes, including providing disruptive camouflage and a basis for vision-based communication. In trying to understand the role of a specific pattern, it is important to first assess the extent to which an observer can resolve the pattern, itself determined, at least in part, by the observer's visual acuity. Here, we studied the visual acuity of two species of reef fish - Pomacentrus amboinensis and Pseudochromis fuscus - using both anatomical and behavioural estimates. The two species share a common habitat but are members of different trophic levels (predator versus herbivore/omnivore) and perform different visual tasks. On the basis of the anatomical study, we estimated visual acuity to lie between 4.1 and 4.6 cycles deg-1 for P. amboinensis and 3.2 and 3.6 cycles deg-1 for P. fuscus Behavioural acuity estimates were considerably lower, ranging between 1.29 and 1.36 cycles deg-1 for P. amboinensis and 1.61 and 1.71 cycles deg-1 for P. fuscus Our results show that two species from the same habitat have only moderately divergent visual capabilities, despite differences in their general life histories. The difference between anatomical and behavioural estimates is an important finding as the majority of our current knowledge on the resolution capabilities of reef fish comes from anatomical measurements. Our findings suggest that anatomical estimates may represent the highest potential acuity of fish but are not indicative of actual performance, and that there is unlikely to be a simple scaling factor to link the two measures across all fish species.

From crypsis to mimicry: changes in colour and the configuration of the visual system during ontogenetic habitat transitions in a coral reef fish.

Animals often change their habitat throughout ontogeny; yet, the triggers for habitat transitions and how these correlate with developmental changes - e.g. physiological, morphological and behavioural - remain largely unknown. Here, we investigated how ontogenetic changes in body coloration and of the visual system relate to habitat transitions in a coral reef fish. Adult dusky dottybacks, Pseudochromis fuscus, are aggressive mimics that change colour to imitate various fishes in their surroundings; however, little is known about the early life stages of this fish. Using a developmental time series in combination with the examination of wild-caught specimens, we revealed that dottybacks change colour twice during development: (i) nearly translucent cryptic pelagic larvae change to a grey camouflage coloration when settling on coral reefs; and (ii) juveniles change to mimic yellow- or brown-coloured fishes when reaching a size capable of consuming juvenile fish prey. Moreover, microspectrophotometric (MSP) and quantitative real-time PCR (qRT-PCR) experiments show developmental changes of the dottyback visual system, including the use of a novel adult-specific visual gene (RH2 opsin). This gene is likely to be co-expressed with other visual pigments to form broad spectral sensitivities that cover the medium-wavelength part of the visible spectrum. Surprisingly, the visual modifications precede changes in habitat and colour, possibly because dottybacks need to first acquire the appropriate visual performance before transitioning into novel life stages.

Opsin evolution in damselfish: convergence, reversal, and parallel evolution across tuning sites.

The visual system plays a role in nearly every aspect of an organism's life history, and there is a direct link between visual pigment phenotypes and opsin genotypes. In previous studies of African cichlid fishes, we found evidence for positive selection among some opsins, with sequence variation greatest for opsins producing the shortest and longest wavelength visual pigments. In this study, we examined opsin evolution in the closely related damselfish family (Pomacentridae), a group of reef fishes that are distributed widely and have a documented fossil record of at least 50 million years (MY). We found increased functional variation in the protein sequences of opsins at the short- and long-wavelength ends of the visual spectrum, in agreement with the African cichlids, despite an order of magnitude difference in the ages of the two radiations. We also reconstructed amino acid substitutions across opsin tuning sites. These reconstructions indicated multiple instances of parallel evolution, at least one definitive case of convergent evolution, and one evolutionary reversal. Our findings show that the amino acids at spectral tuning sites are labile evolutionarily, and that the same codons evolve repeatedly. These findings emphasize that the aquatic light environment can shape opsin sequence evolution. They further show that phylogenetic approaches can provide important insights into the mechanisms by which natural selection "tinkers" with phenotypes.

Cleaner wrasse Labroides dimidiatus perform above chance in a "matching-to-sample" experiment.

Concept learning have been studied widely in non-human animal species within or not an ecological context. Here we tested whether cleaner fish Labroides dimidiatus, which show generalised rule learning in an ecologically relevant context; they generalise that any predator may provide protection from being chased by other fish; can also learn a general concept when presented with abstract cues. We tested for this ability in the matching-to-sample task. In this task, a sample is shown first, and then the subject needs to choose the matching sample over a simultaneously presented different one in order to obtain a food reward. We used the most general form of the task, using each stimulus only once in a total of 200 trials. As a group, the six subjects performed above chance, and four individuals eventually reached learning criteria. However, individual performance was rather unstable, yielding overall only 57% correct choices. These results add to the growing literature that ectotherms show the ability of abstract concept learning, though the lack of stable high performance may indicate quantitative performance differences to endotherms.

Evidence for a consistent use of external cues by marine fish larvae for orientation.

The larval stage is the main dispersive process of most marine teleost species. The degree to which larval behavior controls dispersal has been a subject of debate. Here, we apply a cross-species meta-analysis, focusing on the fundamental question of whether larval fish use external cues for directional movement (i.e., directed movement). Under the assumption that directed movement results in straighter paths (i.e., higher mean vector lengths) compared to undirected, we compare observed patterns to those expected under undirected pattern of Correlated Random Walk (CRW). We find that the bulk of larvae exhibit higher mean vector lengths than those expected under CRW, suggesting the use of external cues for directional movement. We discuss special cases which diverge from our assumptions. Our results highlight the potential contribution of orientation to larval dispersal outcomes. This finding can improve the accuracy of larval dispersal models, and promote a sustainable management of marine resources.

Behavioural evidence for colour vision in an elasmobranch.

Little is known about the sensory abilities of elasmobranchs (sharks, skates and rays) compared with other fishes. Despite their role as apex predators in most marine and some freshwater habitats, interspecific variations in visual function are especially poorly studied. Of particular interest is whether they possess colour vision and, if so, the role(s) that colour may play in elasmobranch visual ecology. The recent discovery of three spectrally distinct cone types in three different species of ray suggests that at least some elasmobranchs have the potential for functional trichromatic colour vision. However, in order to confirm that these species possess colour vision, behavioural experiments are required. Here, we present evidence for the presence of colour vision in the giant shovelnose ray (Glaucostegus typus) through the use of a series of behavioural experiments based on visual discrimination tasks. Our results show that these rays are capable of discriminating coloured reward stimuli from other coloured (unrewarded) distracter stimuli of variable brightness with a success rate significantly different from chance. This study represents the first behavioural evidence for colour vision in any elasmobranch, using a paradigm that incorporates extensive controls for relative stimulus brightness. The ability to discriminate colours may have a strong selective advantage for animals living in an aquatic ecosystem, such as rays, as a means of filtering out surface-wave-induced flicker.

Categorical face perception in fish: How a fish brain warps reality to dissociate "same" from "different".

Categorical perception (CP) is the phenomenon by which a smoothly varying stimulus property undergoes a nonlinear transformation during processing in the brain. Consequently, the stimuli are perceived as belonging to distinct categories separated by a sharp boundary. Originally thought to be largely innate, the discovery of CP in tasks such as novel image discrimination has piqued the interest of cognitive scientists because it provides compelling evidence that learning can shape a category's perceptual boundaries. CP has been particularly closely studied in human face perception. In nonprimates, there is evidence for CP for sound and color discrimination, but not for image or face discrimination. Here, we investigate the potential for learned CP in a lower vertebrate, the damselfish Pomacentrus amboinensis. Specifically, we tested whether the ability of these fish to discriminate complex facial patterns tracked categorical rather than metric differences in the stimuli. We first trained the fish to discriminate sets of two facial patterns. Next, we morphed between these patterns and determined the just noticeable difference (JND) between a morph and original image. Finally, we tested for CP by analyzing the discrimination ability of the fish for pairs of JND stimuli along the spectrum of morphs between two original images. Discrimination performance was significant for the image pair straddling the boundary between categories, and chance for equivalent stimulus pairs on either side, thus producing the classic "category boundary" effect. Our results reveal how perception can be influenced in a top-down manner even in the absence of a visual cortex.

Red fluorescence in reef fish: a novel signalling mechanism?

BACKGROUND: At depths below 10 m, reefs are dominated by blue-green light because seawater selectively absorbs the longer, 'red' wavelengths beyond 600 nm from the downwelling sunlight. Consequently, the visual pigments of many reef fish are matched to shorter wavelengths, which are transmitted better by water. Combining the typically poor long-wavelength sensitivity of fish eyes with the presumed lack of ambient red light, red light is currently considered irrelevant for reef fish. However, previous studies ignore the fact that several marine organisms, including deep sea fish, produce their own red luminescence and are capable of seeing it. RESULTS: We here report that at least 32 reef fishes from 16 genera and 5 families show pronounced red fluorescence under natural, daytime conditions at depths where downwelling red light is virtually absent. Fluorescence was confirmed by extensive spectrometry in the laboratory. In most cases peak emission was around 600 nm and fluorescence was associated with guanine crystals, which thus far were known for their light reflecting properties only. Our data indicate that red fluorescence may function in a context of intraspecific communication. Fluorescence patterns were typically associated with the eyes or the head, varying substantially even between species of the same genus. Moreover red fluorescence was particularly strong in fins that are involved in intraspecific signalling. Finally, microspectrometry in one fluorescent goby, Eviota pellucida, showed a long-wave sensitivity that overlapped with its own red fluorescence, indicating that this species is capable of seeing its own fluorescence. CONCLUSION: We show that red fluorescence is widespread among marine fishes. Many features indicate that it is used as a private communication mechanism in small, benthic, pair- or group-living fishes. Many of these species show quite cryptic colouration in other parts of the visible spectrum. High inter-specific variation in red fluorescence and its association with structures used in intra-specific signalling further corroborate this view. Our findings challenge the notion that red light is of no importance to marine fish, calling for a reassessment of its role in fish visual ecology in subsurface marine environments.

The prototype effect revisited: Evidence for an abstract feature model of face recognition.

Humans typically have a remarkable memory for faces. Nonetheless, in some cases they can be fooled. Experiments described in this paper provide new evidence for an effect in which observers falsely "recognize" a face that they have never seen before. The face is a chimera (prototype) built from parts extracted from previously viewed faces. It is known that faces of this kind can be confused with truly familiar faces, a result referred to as the prototype effect. However, recent studies have failed to find evidence for a full effect, one in which the prototype is regarded not only as familiar, but as more familiar than faces which have been seen before. This study sought to reinvestigate the effect. In a pair of experiments, evidence is reported for the full effect based on both an old/new discrimination task and a familiarity ranking task. The results are shown to be consistent with a recognition model in which faces are represented as combinations of reusable, abstract features. In a final experiment, novel predictions of the model are verified by comparing the size of the prototype effect for upright and upside-down faces. Despite the fundamentally piecewise nature of the model, an explanation is provided as to how it can also account for the sensitivity of observers to configural and holistic cues. This discussion is backed up with the use of an unsupervised network model. Overall, the paper describes how an abstract feature-based model can reconcile a range of results in the face recognition literature and, in turn, lessen currently perceived differences between the representation of faces and other objects.

Discrimination of human faces by archerfish (Toxotes chatareus).

Two rival theories of how humans recognize faces exist: (i) recognition is innate, relying on specialized neocortical circuitry, and (ii) recognition is a learned expertise, relying on general object recognition pathways. Here, we explore whether animals without a neocortex, can learn to recognize human faces. Human facial recognition has previously been demonstrated for birds, however they are now known to possess neocortex-like structures. Also, with much of the work done in domesticated pigeons, one cannot rule out the possibility that they have developed adaptations for human face recognition. Fish do not appear to possess neocortex-like cells, and given their lack of direct exposure to humans, are unlikely to have evolved any specialized capabilities for human facial recognition. Using a two-alternative forced-choice procedure, we show that archerfish (Toxotes chatareus) can learn to discriminate a large number of human face images (Experiment 1, 44 faces), even after controlling for colour, head-shape and brightness (Experiment 2, 18 faces). This study not only demonstrates that archerfish have impressive pattern discrimination abilities, but also provides evidence that a vertebrate lacking a neocortex and without an evolutionary prerogative to discriminate human faces, can nonetheless do so to a high degree of accuracy.

Same/Different Abstract Concept Learning by Archerfish (Toxotes chatareus).

While several phylogenetically diverse species have proved capable of learning abstract concepts, previous attempts to teach fish have been unsuccessful. In this report, the ability of archerfish (Toxotes chatareus) to learn the concepts of sameness and difference using a simultaneous two-item discrimination task was tested. Six archerfish were trained to either select a pair of same or different stimuli which were presented simultaneously. Training consisted of a 2-phase approach. Training phase 1: the symbols in the same and different pair did not change, thereby allowing the fish to solve the test through direct association. The fish were trained consecutively with four different sets of stimuli to familiarize them with the general procedure before moving on to the next training phase. Training phase 2: six different symbols were used to form the same or different pairs. After acquisition, same/different concept learning was tested by presenting fish with six novel stimuli (transfer test). Five fish successfully completed the first training phase. Only one individual passed the second training phase, however, transfer performance was consistent with chance. This individual was given further training using 60 training exemplars but the individual was unable to reach the training criterion. We hypothesize that archerfish are able to solve a limited version of the same/different test by learning the response to each possible stimulus configuration or by developing a series of relatively simple choice contingencies. We conclude that the simultaneous two-item discrimination task we describe cannot be successfully used to test the concepts of same and different in archerfish. In addition, despite considerable effort training archerfish using several tests and training methods, there is still no evidence that fish can learn an abstract concept-based test.

Facing the environment: onset and development of UV markings in young fish.

Most colour patterns in animals represent an elegant compromise between conspicuousness to ensure effective communication with preferred receivers and camouflage to avoid attracting the attention of unwanted predators. Many species, including several coral reef fishes, overcome this conflict by using ultraviolet (UV) colouration and signalling, as these colours are visible only over short distances and are often invisible to their predators. Despite a great interest in their behavioural significance and ecological influence on survival, little is known about when these colours first develop on the bodies of free-living animals. Here we show for the first time that the UV facial patterns of a coral reef fish do not develop in captivity but only when juveniles experience the socio-behavioural conditions of their natural environment. Using field and laboratory experiments, we determined that the onset and early development of these UV facial markings did not occur at metamorphosis. Instead, juveniles developed the UV markings during their first two weeks on the reef. Exposure to different reef environments revealed significant plasticity in the development of these markings. The direct or indirect (through intraspecific interactions) exposure to predators is a likely candidate trigger for the plastic development of these UV markings in the wild.

Do human activities influence survival and orientation abilities of larval fishes in the ocean?

The larval stages of most marine fishes spend days to weeks in the pelagic environment, where they must find food and avoid predators in order to survive. Some fish only spend part of their life history in the pelagic environment before returning to their adult habitat, for example, a coral reef. The sensory systems of larval fish develop rapidly during the first few days of their lives, and here we concentrate on the various sensory cues the fish have available to them for survival in the pelagic environment. We focus on the larvae of reef fishes because most is known about them. We also review the major threats caused by human activities that have been identified to have worldwide impact and evaluate how these threats may impact larval-fish survival and orientation abilities. Many human activities negatively affect larval-fish sensory systems or the cues the fish need to detect. Ultimately, this could lead to decreased numbers of larvae surviving to settlement, and, therefore, to decreased abundance of adult fishes. Although we focus on species wherein the larvae and adults occupy different habitats (pelagic and demersal, respectively), it is important to acknowledge that the potential anthropogenic effects we identify may also apply to larvae of species like tuna and herring, where both larvae and adults are pelagic.

Concept learning and the use of three common psychophysical paradigms in the archerfish (Toxotes chatareus).

Archerfish are well known for their specialized hunting technique of spitting water at prey located above the water line. This unique ability has made them a popular focus of study as researchers try to understand the mechanisms involved in targeting and spitting. In more recent years, archerfish have also become an increasingly popular model for studying visual discrimination and learning in general. Until now, only the alternative forced-choice (AFC) task has been used with archerfish, however, they may be capable of learning other classical discrimination tasks. As well as providing alternative, and potentially more efficient, means for testing their visual capabilities, these other tasks may also provide deeper insight into the extent to which an organism with no cortex can grasp the concepts underlying these tasks. In this paper, we consider both the matched-to-sample (MTS) and the odd-one-out (OOO) tasks as they require the subject to learn relatively sophisticated concepts rather than a straight, stimulus-reward relationship, of the kind underlying AFC tasks. A variety of line drawings displayed on a monitor were used as stimuli. We first determined if archerfish could complete the MTS and OOO test and then evaluated their ability to be retrained to new stimuli using a 4-AFC test. We found that archerfish were unable to learn the MTS and had only a limited capacity for learning the OOO task. We conclude that the MTS and OOO are impractical as paradigms for behavioral experiments with archerfish. However, the archerfish could rapidly learn to complete an AFC test and select the conditioned stimulus with a high degree of accuracy when faced with four stimuli, making this a powerful test for behavioral studies testing visual discrimination. In addition, the fish were able to learn the concept of oddity under particular training circumstances. This paper adds to the growing evidence that animals without a cortex are capable of learning some higher order concepts.

Spectral and spatial selectivity of luminance vision in reef fish.

Luminance vision has high spatial resolution and is used for form vision and texture discrimination. In humans, birds and bees luminance channel is spectrally selective-it depends on the signals of the long-wavelength sensitive photoreceptors (bees) or on the sum of long- and middle-wavelength sensitive cones (humans), but not on the signal of the short-wavelength sensitive (blue) photoreceptors. The reasons of such selectivity are not fully understood. The aim of this study is to reveal the inputs of cone signals to high resolution luminance vision in reef fish. Sixteen freshly caught damselfish, Pomacentrus amboinensis, were trained to discriminate stimuli differing either in their color or in their fine patterns (stripes vs. cheques). Three colors ("bright green", "dark green" and "blue") were used to create two sets of color and two sets of pattern stimuli. The "bright green" and "dark green" were similar in their chromatic properties for fish, but differed in their lightness; the "dark green" differed from "blue" in the signal for the blue cone, but yielded similar signals in the long-wavelength and middle-wavelength cones. Fish easily learned to discriminate "bright green" from "dark green" and "dark green" from "blue" stimuli. Fish also could discriminate the fine patterns created from "dark green" and "bright green". However, fish failed to discriminate fine patterns created from "blue" and "dark green" colors, i.e., the colors that provided contrast for the blue-sensitive photoreceptor, but not for the long-wavelength sensitive one. High resolution luminance vision in damselfish, Pomacentrus amboinensis, does not have input from the blue-sensitive cone, which may indicate that the spectral selectivity of luminance channel is a general feature of visual processing in both aquatic and terrestrial animals.

Temporal constraints on predation risk assessment in a changing world.

Habitat degradation takes various forms and likely represents the most significant threat to our global biodiversity. Recently, we have seen considerable attention paid to increasing global CO2 emissions which lead to ocean acidification (OA). Other stressors, such as changing levels of ultraviolet radiation (UVR), also impact biodiversity but have received much less attention in the recent past. Here we examine fundamental questions about temporal aspects of risk assessment by coral reef damselfish and provide critical insights into how OA and UVR influence this assessment. Chemical cues released during a predator attack provide a rich source of information that other prey animals use to mediate their risk of predation and are the basis of the majority of trait-mediated indirect interactions in aquatic communities. However, we have surprisingly limited information about temporal aspects of risk assessment because we lack knowledge about how long chemical cues persist after they are released into the environment. Here, we showed that under ambient CO2 conditions (~385 µatm), alarm cues of ambon damselfish (Pomacentrus amboinensis) did not degrade within 30 min in the absence of ultraviolet radiation (UVR), but were degraded within 15 min when the CO2 was increased to ~905 µatm. In experiments that used filters to eliminate UVR, we found minimal degradation of alarm cues within 30 min, whereas under ambient UVR conditions, alarm cues were completely degraded within 15 min. Moreover, in the presence of both UVR and elevated CO2, alarm cues were broken down within 5 min. Our results highlight that alarm cues degrade surprisingly quickly under natural conditions and that anthropogenic changes have the potential to dramatically change rates of cue degradation in the wild. This has considerable implications for risk assessment and consequently the importance of trait-mediated indirect interactions in coral-reef communities.

Ultraviolet-B wavelengths regulate changes in UV absorption of cleaner fish Labroides dimidiatus mucus.

High-energy wavelengths in the ultraviolet-B (UVB, 280-315 nm) and the UVA (315-400-nm) portion of the spectrum are harmful to terrestrial and aquatic organisms. Interestingly, UVA is also involved in the repair of UV induced damage. Organisms living in shallow coral reef environments possess UV absorbing compounds, such as mycosporine-like amino acids, to protect them from UV radiation. While it has been demonstrated that exposure to UV (280-400 nm) affects the UV absorbance of fish mucus, whether the effects of UV exposure vary between UVB and UVA wavelengths is not known. Therefore, we investigated whether the UVB, UVA, or photosynthetically active radiation (PAR, 400-700 nm) portions of the spectrum affected the UV absorbance of epithelial mucus and Fulton's body condition index of the cleaner fish Labroides dimidiatus. We also compared field-measured UV absorbance with laboratory based high-performance liquid chromatography measurements of mycosporine-like amino acid concentrations. After 1 week, we found that the UV absorbance of epithelial mucus was higher in the UVB+UVA+PAR treatment compared with the UVA+PAR and PAR only treatments; after 2 and 3 weeks, however, differences between treatments were not detected. After 3 weeks, Fulton's body condition index was lower for fish in the UVB+UVA+PAR compared with PAR and UVA+PAR treatments; furthermore, all experimentally treated fish had a lower Fulton's body condition index than did freshly caught fish. Finally, we found a decrease with depth in the UV absorbance of mucus of wild-caught fish. This study suggests that the increase in UV absorbance of fish mucus in response to increased overall UV levels is a function of the UVB portion of the spectrum. This has important implications for the ability of cleaner fish and other fishes to adjust their mucus UV protection in response to variations in environmental UV exposure.

Exogenous material in the inner ear of the adult Port Jackson shark, Heterodontus portusjacksoni (Elasmbranchii).

Previous studies have suggested that the inner ear of some benthic species of elasmobranchs contain only exogenous material within their otoconial organs, a unique feature within vertebrates. However, these examinations have not accounted for the possibility of otoconial degeneration or used modern experimental methods to identify the materials present. Both of these issues are addressed in this study using inner ear samples from the adult Port Jackson shark, Heterodontus portusjacksoni. A comparison of the otoconial mass in fixed specimens over short and medium time scales reveals that over those timescales the degeneration of calcium carbonate-based otoconia does not occur and confirms that calcium carbonate-based otoconia are not found within the otoconial organs of H. portusjacksoni. Additionally, microanalysis of the chemical composition and ultrastructure of the otoconial mass using energy dispersive X-ray microanalysis and scanning electron microscopy confirms that the entire otoconial mass is comprised of exogenous silicon dioxide particles, bound within a carbon matrix. This exogenous material is suggested to play an equivalent role to the otoconia found in other species of elasmobranchs, and allows both hearing and vestibular control to occur in benthic sharks that spend their lives foraging within a sandy substrate.

A comparison of the external morphology of the membranous inner ear in elasmobranchs.

Studies on the elasmobranch inner ear have focused predominantly on a small group of sharks, particularly, carcharhinids. As a result, subsequent studies in other species have subdivided species into two main groups: those typical and those atypical of carcharhinid sharks. This study proposes a different set of inner-ear morphology groupings to those previously suggested. The inner ears from 17 species of elasmobranchs (representing both sharks and rays) are examined in this study and based on morphometric data some groups include both rays and sharks. Four groups are now proposed based predominantly on the shape and dimensions of the membranous otoconial organs, and characteristics of the semicircular canals. Evident morphological differences between the ear types belonging to the new groups include the membranes of the semicircular canals being bound to the otoconial organs in some species, while only being connected via the canal ducts in others, as well as clear variation present in saccular organ size. Previous studies examining variation in the inner ear have attributed differences to either phylogeny or functional significance. Results from this study suggest that neither phylogeny nor feeding strategy solely accounts for the morphological diversity present in the external morphology of the elasmobranch inner ear.