Acoustic Communication in Butterflyfishes: Anatomical Novelties, Physiology, Evolution, and Behavioral Ecology.

Coral reef fishes live in noisy environments that may challenge their capacity for acoustic communication. Butterflyfishes (Family Chaetodontidae) are prominent and ecologically diverse members of coral reef communities worldwide. The discovery of a novel association of anterior swim bladder horns with the lateral line canal system in the genus Chaetodon (the laterophysic connection) revealed a putative adaptation for enhancement of sound reception by the lateral line system and/or the ear. Behavioral studies show that acoustic communication is an important component of butterflyfish social behavior. All bannerfish (Forcipiger, Heniochus, and Hemitaurichthys) and Chaetodon species studied thus far produce several sound types at frequencies of <1 to >1000 Hz. Ancestral character state analyses predict the existence of both shared (head bob) and divergent (tail slap) acoustic behaviors in these two clades. Experimental auditory physiology shows that butterflyfishes are primarily sensitive to stimuli associated with hydrodynamic particle accelerations of ≤500 Hz. In addition, the gas-filled swim bladder horns in Chaetodon are stimulated by sound pressure, which enhances and extends their auditory sensitivity to 1700-2000 Hz. The broadband spectrum of ambient noise present on coral reefs overlaps with the frequency characteristics of their sounds, thus both the close social affiliations common among butterflyfishes and the evolution of the swim bladder horns in Chaetodon facilitate their short-range acoustic communication. Butterflyfishes provide a unique and unexpected opportunity to carry out studies of fish bioacoustics in the lab and the field that integrate the study of sensory anatomy, physiology, evolution, and behavioral ecology.

Diversity and evolution of sound production in the social behavior of Chaetodon butterflyfishes.

Fish produce context-specific sounds during social communication, but it is not known how acoustic behaviors have evolved in relation to specializations of the auditory system. Butterflyfishes (family Chaetodontidae) have a well-defined phylogeny and produce pulsed communication sounds during social interactions on coral reefs. Recent work indicates that two sound production mechanisms exist in the bannerfish clade and that other mechanisms are used in the Chaetodon clade, which is distinguished by an auditory specialization, the laterophysic connection (LC). Here, we determine the kinematic action patterns associated with sound production during social interactions in four Chaetodon subgenera and the non-laterophysic fish Forcipiger flavissimus. Some Chaetodon species share the head bob acoustic behavior with F. flavissimus, which along with other sounds in the 100-1000 Hz spectrum, are probably adequate to stimulate the ear, swim bladder or LC of a receiver fish. In contrast, only Chaetodon species produced the tail slap sound, which involves a 1-30 Hz hydrodynamic pulse that is likely to stimulate the receiver's ear and lateral line at close distances, but not the swim bladder or LC. Reconstructions of ancestral character states appear equivocal for the head bob and divergent for the tail slap acoustic behaviors. Independent contrast analysis shows a correlation between sound duration and stimulus intensity characters. The intensities of the tail slap and body pulse sounds in Chaeotodon species are correlated with body size and can provide honest communication signals. Future studies on fish acoustic communication should investigate low-frequency and infrasound acoustic fields to understand the integrated function of the ear and lateral line, and their evolutionary patterns.

Sound pressure enhances the hearing sensitivity of Chaetodon butterflyfishes on noisy coral reefs.

Butterflyfishes are conspicuous members of coral reefs that communicate with acoustic signals during social interactions with mates and other conspecifics. Members of the genus Chaetodon have a laterophysic connection (LC) - a unique association of anterior swim bladder horns and the cranial lateral line - but the action of the LC system on auditory sensitivity is unexplored. Here, we show in baseline auditory evoked potential threshold experiments that Forcipiger flavissimus (which lacks swim bladder horns and LC) is sensitive to sound tones from 100 Hz up to 1000 Hz, and that thresholds for three species of Chaetodon are 10-15 dB lower, with extended hearing ranges up to 1700-2000 Hz. The relatively high thresholds to sound pressure and low pass response near 500 Hz for all four species are consistent with a primary sensitivity to hydrodynamic particle acceleration rather than sound pressure. Deflation of the swim bladder in F. flavissimus had no measurable effect on auditory sensitivity. In contrast, displacement of gas from the swim bladder horns in Chaetodon multicinctus and Chaetodon auriga increased thresholds (decreased sensitivity) by 5-20 dB, with the greatest effect at 600 Hz. The evolution of swim bladder horns associated with the LC system in Chaetodon species has increased hearing sensitivity through sound pressure transduction in the frequency bands used for social acoustic communication. The close affiliative behaviors that are common in Chaetodon species and other butterflyfish facilitate sound perception and acoustic communication at close distances relative to the high background noise levels found in their natural reef environment.

Cytoarchitecture of the telencephalon in the coral reef multiband Butterflyfish ( Chaetodon multicinctus : Perciformes).

Detailed neuroanatomical studies of model species are necessary to facilitate comparative experiments which test hypotheses relevant to brain evolution and function. Butterflyfishes (Chaetodontidae) boast numerous sympatric species that differ in social behavior, aggression and feeding ecology. However, the ability to test hypotheses relevant to brain function in this family is hindered by the lack of detailed neural descriptions. The cytoarchitecture of the telencephalon in the monogamous and territorial multiband butterflyfish, Chaetodon multicinctus, was determined with Nissl-stained serial sections and an immunohistochemical analysis of arginine vasotocin (AVT), serotonin, substance P and tyrosine hydroxylase. The ventral telencephalon was similar to that of other perciform fishes studied, with one major difference. A previously undescribed postcommissural region, the cuneate nucleus, was identified and putatively assigned to the ventral telencephalon. While the function of this nucleus is unknown, preliminary studies indicate that it may be part of a behaviorally relevant subpallial neural circuit that is modulated by AVT. The dorsal telencephalon consisted of 15 subdivisions among central, medial, lateral, dorsal and posterior zones. Several regions of the dorsal telencephalon of C. multicinctus differed from many other perciform fishes examined thus far. The nucleus taenia was in a more caudal position, and the central and lateral zones were enlarged. Within the lateral zone, an unusual third, ventral subdivision and a large-celled division were present. One hypothesis is that the enlarged ventral subdivision of the lateral zone (potential hippocampus homolog) relates to an enhancement of spatial learning or olfactory memory, which are important for this coral reef fish. This study provides the neuroanatomical basis for future comparative and evolutionary studies of brain organization and neuropeptide distributions, physiological studies of neural processing and insight into the complex social behavior of butterflyfishes.

Fast drum strokes: novel and convergent features of sonic muscle ultrastructure, innervation, and motor neuron organization in the Pyramid Butterflyfish (Hemitaurichthys polylepis).

Sound production that is mediated by intrinsic or extrinsic swim bladder musculature has evolved multiple times in teleost fishes. Sonic muscles must contract rapidly and synchronously to compress the gas-filled bladder with sufficient velocity to produce sound. Muscle modifications that may promote rapid contraction include small fiber diameter, elaborate sarcoplasmic reticulum (SR), triads at the A-I boundary, and cores of sarcoplasm. The diversity of innervation patterns indicate that sonic muscles have independently evolved from different trunk muscle precursors. The analysis of sonic motor pathways in distantly related fishes is required to determine the relationships between sonic muscle evolution and function in acoustic signaling. We examined the ultrastructure of sonic and adjacent hypaxial muscle fibers and the distribution of sonic motor neurons in the coral reef Pyramid Butterflyfish (Chaetodontidae: Hemitaurichthys polylepis) that produces sound by contraction of extrinsic sonic muscles near the anterior swim bladder. Relative to adjacent hypaxial fibers, sonic muscle fibers were sparsely arranged among the endomysium, smaller in cross-section, had longer sarcomeres, a more elaborate SR, wider t-tubules, and more radially arranged myofibrils. Both sonic and non-sonic muscle fibers possessed triads at the Z-line, lacked sarcoplasmic cores, and had mitochondria among the myofibrils and concentrated within the peripheral sarcoplasm. Sonic muscles of this derived eutelost possess features convergent with other distant vocal taxa (other euteleosts and non-euteleosts): small fiber diameter, a well-developed SR, and radial myofibrils. In contrast with some sonic fishes, however, Pyramid Butterflyfish sonic muscles lack sarcoplasmic cores and A-I triads. Retrograde nerve label experiments show that sonic muscle is innervated by central and ventrolateral motor neurons associated with spinal nerves 1-3. This restricted distribution of sonic motor neurons in the spinal cord differs from many euteleosts and likely reflects the embryological origin of sonic muscles from hypaxial trunk precursors rather than occipital somites.

Sound production in the longnose butterflyfishes (genus Forcipiger): cranial kinematics, muscle activity and honest signals.

Many teleost fishes produce sounds for social communication with mechanisms that do not involve swim bladder musculature. Such sounds may reflect physical attributes of the sound-production mechanism, be constrained by body size and therefore control signal reliability during agonistic behaviors. We examined kinematics of the cranium, median fins and caudal peduncle during sound production in two territorial chaetodontid butterflyfish sister species: forcepsfish (Forcipiger flavissimus) and longnose butterflyfish (F. longirostris). During intraspecific agonistic encounters, both species emit a single pulse sound that precedes rapid cranial rotation at velocities and accelerations that exceed those of prey strikes by many ram- and suction-feeding fishes. Electromyography showed that onsets of activity for anterior epaxialis, sternohyoideus, A1 and A2 adductor mandibulae muscles and sound emission are coincident but precede cranial elevation. Observations indicate that sound production is driven by epaxial muscle contraction whereas a ventral linkage between the head and pectoral girdle is maintained by simultaneous activity from the adductor mandibulae and sternohyoideus. Thus, the girdle, ribs and rostral swim bladder are pulled anteriorly before the head is released and rotated dorsally. Predictions of the hypothesis that acoustic signals are indicators of body size and kinematic performance were confirmed. Variation in forcepsfish sound duration and sound pressure level is explained partly by cranial elevation velocity and epaxial electromyogram duration. Body size, however, explains most variation in duration and sound pressure level. These observed associations indicate that forcepsfish sounds may be accurate indicators of size and condition that are related to resource holding potential during social encounters.

Arginine vasotocin neuronal phenotypes and their relationship to aggressive behavior in the territorial monogamous multiband butterflyfish, Chaetodon multicinctus.

Intra and interspecific comparisons of arginine vasotocin (AVT) and its mammalian homolog arginine vasopressin (AVP) demonstrate several relationships between these neuropeptides and aggression/dominance behaviors. Prior studies in coral reef butterflyfishes and other fishes indicate that features of AVT neurons in the gigantocellular preoptic area (gPOA) and axon varicosities within the ventral nucleus of the ventral telencephalon should have a positive relationship with aggressive behavior, whereas AVT-ir neuronal features in the parvocellular preoptic area (pPOA) should have a negative relationship. We measured the offensive aggression of wild caught territorial monogamous multiband butterflyfish, Chaetodon multicinctus, in a simple lab paradigm that controlled for social context and variations in social stimuli. Offensive aggression did not follow a clear stereotyped pattern, but rather a complex sequence that includes five action patterns and two approach behaviors. We then used immunohistochemistry to test for associations between AVT immunoreactive features and projections with overall offensive aggression. Our results indicate that gPOA cell number was positively related to aggression while both the size and number of pPOA cells were negatively related to aggression. No association between aggression and the number of axon varicosities in the telencephalic region proposed to be associated with aggression was found. This study provides further support for the relationship between AVT neuronal features and aggression in fishes, and provides preliminary evidence that this relationship may relate to the motivation to produce aggressive behaviors in the immediate future.

Electrosensitive spatial vectors in elasmobranch fishes: implications for source localization.

The electrosense of sharks and rays is used to detect weak dipole-like bioelectric fields of prey, mates and predators, and several models propose a use for the detection of streaming ocean currents and swimming-induced fields for geomagnetic orientation. We assessed pore distributions, canal vectors, complementarity and possible evolutionary divergent functions for ampullary clusters in two sharks, the scalloped hammerhead (Sphyrna lewini) and the sandbar shark (Carcharhinus plumbeus), and the brown stingray (Dasyatis lata). Canal projections were determined from measured coordinates of each electrosensory pore and corresponding ampulla relative to the body axis. These species share three ampullary groups: the buccal (BUC), mandibular (MAN) and superficial ophthalmic (SO), which is subdivided into anterior (SOa) and posterior (SOp) in sharks. The stingray also has a hyoid (HYO) cluster. The SOp in both sharks contains the longest (most sensitive) canals with main projections in the posterior-lateral quadrants of the horizontal plane. In contrast, stingray SO canals are few and short with the posterior-lateral projections subsumed by the HYO. There was strong projection coincidence by BUC and SOp canals in the posterior lateral quadrant of the hammerhead shark, and laterally among the stingray BUC and HYO. The shark SOa and stingray SO and BUC contain short canals located anterior to the mouth for detection of prey at close distance. The MAN canals of all species project in anterior or posterior directions behind the mouth and likely coordinate prey capture. Vertical elevation was greatest in the BUC of the sandbar shark, restricted by the hammerhead cephalofoil and extremely limited in the dorsoventrally flattened stingray. These results are consistent with the functional subunit hypothesis that predicts specialized ampullary functions for processing of weak dipole and geomagnetic induced fields, and provides an anatomical basis for future experiments on central processing of different forms of relevant electric stimuli.

Gonadotropin-releasing hormone (GnRH) modulates auditory processing in the fish brain.

Gonadotropin-releasing hormone 1 (GnRH1) neurons control reproductive activity, but GnRH2 and GnRH3 neurons have widespread projections and function as neuromodulators in the vertebrate brain. While these extra-hypothalamic GnRH forms function as olfactory and visual neuromodulators, their potential effect on processing of auditory information is unknown. To test the hypothesis that GnRH modulates the processing of auditory information in the brain, we used immunohistochemistry to determine seasonal variations in these neuropeptide systems, and in vivo single-neuron recordings to identify neuromodulation in the midbrain torus semicircularis of the soniferous damselfish Abudefduf abdominalis. Our results show abundant GnRH-immunoreactive (-ir) axons in auditory processing regions of the midbrain and hindbrain. The number of extra-hypothalamic GnRH somata and the density of GnRH-ir axons within the auditory torus semicircularis also varied across the year, suggesting seasonal changes in GnRH influence of auditory processing. Exogenous application of GnRH (sGnRH and cGnRHII) caused a primarily inhibitory effect on auditory-evoked single neuron responses in the torus semicircularis. In the majority of neurons, GnRH caused a long-lasting decrease in spike rate in response to both tone bursts and playbacks of complex natural sounds. GnRH also decreased response latency and increased auditory thresholds in a frequency and stimulus type-dependent manner. To our knowledge, these results show for the first time in any vertebrate that GnRH can influence context-specific auditory processing in vivo in the brain, and may function to modulate seasonal auditory-mediated social behaviors.

Arginine vasotocin neuronal phenotypes, telencephalic fiber varicosities, and social behavior in butterflyfishes (Chaetodontidae): potential similarities to birds and mammals.

The neuropeptide arginine vasopressin (AVP) influences many social behaviors through its action in the forebrain of mammals. However, the function of the homologous arginine vasotocin (AVT) in the forebrain of fishes, specifically the telencephalon remains unresolved. We tested whether the density of AVT-immunoreactive (-ir) fiber varicosities, somata size or number of AVT-ir neuronal phenotypes within the forebrain were predictive of social behavior in reproductive males of seven species of butterflyfishes (family Chaetodontidae) in four phylogenetic clades. Similar to other fishes, the aggressive (often territorial) species in most cases had larger AVT-ir cells within the gigantocellular preoptic cell group. Linear discriminant function analyses demonstrated that the density of AVT-ir varicosities within homologous telencephalic nuclei to those important for social behavior in mammals and birds were predictive of aggressive behavior, social affiliations, and mating system. Of note, the density of AVT-ir varicosities within the ventral nucleus of the ventral telencephalon, thought to be homologous to the septum of other vertebrates, was the strongest predictor of aggressive behavior, social affiliation, and mating system. These results are consistent with the postulate that AVT within the telencephalon of fishes plays an important role in social behavior and may function in a similar manner to that of AVT/AVP in birds and mammals despite having cell populations solely within the preoptic area.

Pulse sound generation, anterior swim bladder buckling and associated muscle activity in the pyramid butterflyfish, Hemitaurichthys polylepis.

Acoustic behaviors are widespread among diverse fish taxa but mechanisms of sound production are known from relatively few species, vary widely and convergent mechanisms are poorly known. We examined the sound production mechanism in the pyramid butterflyfish, Hemitaurichthys polylepis, a member of the socially and ecologically diverse reef fish family Chaetodontidae. In the field, fish produce pulse trains at dusk during social interactions that are probably related to mate attraction and courtship. In laboratory experiments, sound production was synchronized to high-speed video to determine body movement associated with sound generation. In addition, electromyography (EMG) recordings tested the activity of six candidate muscles. Fish produced individual pulses with a mean peak frequency of 97 Hz in rapid succession. EMG experiments show that anterior hypaxial muscles contract at high bilaterally synchronous rates (up to 120 Hz) in near perfect association with rapid inward buckling visible outside the body over the anterior swim bladder. Muscle activity often showed EMG doublets that occurred within the time of a single sound pulse but was not sustained. Buckling and sound pulse rates correlated strongly (R(2)≈1.00) and sound pulse rate measured over two successive pulses (maximum of 38 pulses s(-1)) was lower than muscle firing rate. These results show that the extrinsic swim bladder muscles of pyramid butterflyfish involve single contractions that produce pulses in a manner similar to distantly related teleosts, but involve a novel doublet motor-neuron firing pattern. Thus, the sound production mechanism in pyramid butterflyfish is likely convergent with several percomorph taxa and divergent from the related chaetodontid genus Forcipiger.

Encoding properties of auditory neurons in the brain of a soniferous damselfish: response to simple tones and complex conspecific signals.

The fish auditory system encodes important acoustic stimuli used in social communication, but few studies have examined response properties of central auditory neurons to natural signals. We determined the features and responses of single hindbrain and midbrain auditory neurons to tone bursts and playbacks of conspecific sounds in the soniferous damselfish, Abudefduf abdominalis. Most auditory neurons were either silent or had slow irregular resting discharge rates <20 spikes s(-1). Average best frequency for neurons to tone stimuli was approximately 130 Hz but ranged from 80 to 400 Hz with strong phase-locking. This low-frequency sensitivity matches the frequency band of natural sounds. Auditory neurons were also modulated by playbacks of conspecific sounds with thresholds similar to 100 Hz tones, but these thresholds were lower than that of tones at other test frequencies. Thresholds of neurons to natural sounds were lower in the midbrain than the hindbrain. This is the first study to compare response properties of auditory neurons to both simple tones and complex stimuli in the brain of a recently derived soniferous perciform that lacks accessory auditory structures. These data demonstrate that the auditory fish brain is most sensitive to the frequency and temporal components of natural pulsed sounds that provide important signals for conspecific communication.

Response of the hammerhead shark olfactory epithelium to amino acid stimuli.

Sharks and rays are highly sensitive to chemical stimuli in their natural environment but several hypotheses predict that hammerhead sharks, with their expanded head and enlarged olfactory epithelium, have particularly acute olfactory systems. We used the electro-olfactogram (EOG) technique to compare the relative response of the scalloped hammerhead shark (Sphyrna lewini) olfactory epithelium to 20 proteinogenic amino acids and determine the sensitivity for 6 amino acids. At micromolar concentrations, cysteine evoked the greatest EOG response which was approximately twice as large as that of alanine. The weakest response was obtained for proline followed by aspartic acid and isoleucine. The olfactory epithelium showed adaptation to sequential stimulation, and recovery was related to the inter-stimulus time period. Estimated EOG response thresholds were in the sub-nanomolar range for both alanine (9.2 x 10(-11) M) and cysteine (8.4 x 10(-10) M) and in the micromolar range for proline and serine. These thresholds from 10(-10) to 10(-6) M for the scalloped hammerhead shark are comparable or lower than those reported for other teleost and elasmobranch species. Future work should focus on binary and more complex compounds to test for competition and cross-adaptation for different classes of peripheral receptors, and their responses to molecules found in biologically relevant stimuli.

Central projections of octavolateralis nerves in the brain of a soniferous damselfish (Abudefduf abdominalis).

Sounds and hydrodynamic stimuli are important cues detected by the octavolateralis system in fishes. The central organization of auditory, mechanosensory, and vestibular projections is known for only a few phylogenetically diverse fishes, and less is known about projections in derived perciforms that use sounds for acoustic communication. We used neuronal labeling to provide a detailed analysis of octavolateralis endorgan projections in a soniferous perciform that does not have accessory morphological structures to enhance hearing. Octavolateralis nerves terminate ipsilaterally within seven medullary octaval nuclei: caudal (CON) and medial (MON) octavolateralis, anterior (AON), descending (DON), magnocellular (MgON), tangential (TON), and posterior (PON) octaval nuclei, and the eminentia granularis (EG). Anterior and posterior lateral line nerves project to the CON and MON, with dense projections to the EG. Semicircular canal nerves project primarily to ventral regions including the TON, ventral DON, intermediate DON (DONi), and MgON. Otolithic, semicircular canal, and anterior lateral line nerves all project to the MgON, which may serve a sensorimotor integration function. The DONi receives primarily segregated projections from all otolithic and semicircular canal nerves, whereas the ventral DON and TON receive principally utricular and semicircular canal afferents. The AON receives dense lateral and ventral projections from the saccule and utricle, and medial and dorsal projections from the lagena. These projection patterns are similar to those reported for non-sonic perciforms, and indicate the absence of neuroanatomical modifications in first-order octavolateralis nuclei in species that use acoustic communication. Thus patterns of central projections may be conserved among vocal and non-vocal perciforms.

Functional morphology of the feeding apparatus, feeding constraints, and suction performance in the nurse shark Ginglymostoma cirratum.

The nurse shark, Ginglymostoma cirratum, is an obligate suction feeder that preys on benthic invertebrates and fish. Its cranial morphology exhibits a suite of structural and functional modifications that facilitate this mode of prey capture. During suction-feeding, subambient pressure is generated by the ventral expansion of the hyoid apparatus and the floor of its buccopharyngeal cavity. As in suction-feeding bony fishes, the nurse shark exhibits expansive, compressive, and recovery kinematic phases that produce posterior-directed water flow through the buccopharyngeal cavity. However, there is generally neither a preparatory phase nor cranial elevation. Suction is generated by the rapid depression of the buccopharyngeal floor by the coracoarcualis, coracohyoideus, and coracobranchiales muscles. Because the hyoid arch of G. cirratum is loosely connected to the mandible, contraction of the rectus cervicis muscle group can greatly depress the floor of the buccopharyngeal cavity below the depressed mandible, resulting in large volumetric expansion. Suction pressures in the nurse shark vary greatly, but include the greatest subambient pressures reported for an aquatic-feeding vertebrate. Maximum suction pressure does not appear to be related to shark size, but is correlated with the rate of buccopharyngeal expansion. As in suction-feeding bony fishes, suction in the nurse shark is only effective within approximately 3 cm in front of the mouth. The foraging behavior of this shark is most likely constrained to ambushing or stalking due to the exponential decay of effective suction in front of the mouth. Prey capture may be facilitated by foraging within reef confines and close to the substrate, which can enhance the effective suction distance, or by foraging at night when it can more closely approach prey.

Development of DNA microsatellite markers in the multiband butterflyfish (Chaetodon multicinctus).

Twelve polymorphic microsatellite loci were developed in the multiband (pebbled) butterflyfish Chaetodon multicinctus. The loci were scored in 45 individuals from Hawaii. There were five to 21 alleles per locus with observed heterozygosity ranging from 0.419 to 0.883. Four of the primer sets also reliably amplified polymorphic loci in Chaetodon quadrimaculatus. We expect these markers to be useful for studies of genetic population structure and kinship, for example to determine whether new recruits settling onto reefs are related.

Sound production and spectral hearing sensitivity in the Hawaiian sergeant damselfish, Abudefduf abdominalis.

Sounds provide important signals for inter- and intraspecific communication in fishes, but few studies examine fish acoustic behavior in the context of coevolution of sound production and hearing ability within a species. This study characterizes the acoustic behavior in a reproductive population of the Hawaiian sergeant fish, Abudefduf abdominalis, and compares acoustic features to hearing ability, measured by the auditory evoked potential (AEP) technique. Sergeant fish produce sounds at close distances to the intended receiver (

Sex, seasonal, and stress-related variations in elasmobranch corticosterone concentrations.

Serum corticosterone was previously studied in numerous elasmobranch fishes (sharks, skates and rays), but the role of this steroid, widespread throughout many taxa, has yet to be defined. The goal of this study was to test whether corticosterone varied in response to acute and chronic capture stress, and across the reproductive cycle in the bonnethead shark, Sphyrna tiburo, and Atlantic stingray, Dasyatis sabina. Serum corticosterone in S. tiburo increased following capture and again 24 h post-capture, possibly caused by interference with 1alpha-hydroxycorticosterone, the primary stress hormone in elasmobranchs. Higher serum concentrations in males compared to females were observed in both species. Variations in corticosterone also occurred during the reproductive cycle in both species. Consistent with other taxa, elevations in male bonnethead sharks and stingrays coincided with peak testicular development and mating. Elevations in female bonnethead sharks occurred from the time of mating through sperm storage into early gestation. In contrast, corticosterone levels in female stingrays were low during their protracted mating season, but elevated through late gestation and parturition. These results indicate that corticosterone has a limited role, if any, in acute and chronic stress associated with capture in S. tiburo, but likely has physiological functions associated with its glucocorticoid properties across the reproductive cycle of both species.

From morphology to neural information: the electric sense of the skate.

Morphology typically enhances the fidelity of sensory systems. Sharks, skates, and rays have a well-developed electrosense that presents strikingly unique morphologies. Here, we model the dynamics of the peripheral electrosensory system of the skate, a dorsally flattened batoid, moving near an electric dipole source (e.g., a prey organism). We compute the coincident electric signals that develop across an array of the skate's electrosensors, using electrodynamics married to precise morphological measurements of sensor location, infrastructure, and vector projection. Our results demonstrate that skate morphology enhances electrosensory information. Not only could the skate locate prey using a simple population vector algorithm, but its morphology also specifically leads to quick shifts in firing rates that are well-suited to the demonstrated bandwidth of the electrosensory system. Finally, we propose electrophysiology trials to test the modeling scheme.