Darwin, Haeckel, and the "Mikluskan gas organ theory".

A previously unknown reference to the Russian ethnologist, biologist, and traveler Nikolai N. Miklucho-Maclay (1846-1888) was discovered in correspondence between Charles Darwin (1809-1882) and Ernst Haeckel (1834-1919). This reference has remained unknown to science, even to Miklucho-Maclay's biographers, probably because Darwin used the Russian nickname "Mikluska" when alluding to this young scientist. Here, we briefly outline the story behind the short discussion between Darwin and his German counterpart Haeckel, and highlight its importance for the history of science. Miklucho-Maclay's discovery of a putative swim bladder anlage in sharks, published in 1867, was discussed in four letters between the great biologists. Whereas, Haeckel showed enthusiasm for the finding because it supported (his view on) evolutionary theory, Darwin was less interested, which highlights the conceptual differences between the two authorities. We discuss the scientific treatment of Miklucho-Maclay's observation in the literature and discuss the homology, origin, and destiny of gas organs-swim bladders and lungs-in vertebrate evolution, from an ontogenetic point of view. We show that the conclusions reached by Miklucho-Maclay and Haeckel were rather exaggerated, although they gave rise to fundamental insights, and we illustrate how tree-thinking may lead to differences in the conceptualization of evolutionary change.

Ancient and Nonuniform Loss of Olfactory Receptor Expression Renders the Shark Nose a De Facto Vomeronasal Organ.

Cartilaginous fishes are renowned for a keen sense of smell, a reputation based on behavioral observations and supported by the presence of large and morphologically complex olfactory organs. At the molecular level, genes belonging to the four families coding for most olfactory chemosensory receptors in other vertebrates have been identified in a chimera and a shark, but it was unknown whether they actually code for olfactory receptors in these species. Here, we describe the evolutionary dynamics of these gene families in cartilaginous fishes using genomes of a chimera, a skate, a sawfish, and eight sharks. The number of putative OR, TAAR, and V1R/ORA receptors is very low and stable, whereas the number of putative V2R/OlfC receptors is higher and much more dynamic. In the catshark Scyliorhinus canicula, we show that many V2R/OlfC receptors are expressed in the olfactory epithelium in the sparsely distributed pattern characteristic for olfactory receptors. In contrast, the other three vertebrate olfactory receptor families are either not expressed (OR) or only represented with a single receptor (V1R/ORA and TAAR). The complete overlap of markers of microvillous olfactory sensory neurons with pan-neuronal marker HuC in the olfactory organ suggests the same cell-type specificity of V2R/OlfC expression as for bony fishes, that is, in microvillous neurons. The relatively low number of olfactory receptors in cartilaginous fishes compared with bony fishes could be the result of an ancient and constant selection in favor of a high olfactory sensitivity at the expense of a high discrimination capability.

Exceptionally preserved shark fossils from Mexico elucidate the long-standing enigma of the Cretaceous elasmobranch Ptychodus.

The fossil fish Ptychodus Agassiz, 1834, characterized by a highly distinctive grinding dentition and an estimated gigantic body size (up to around 10 m), has remained one of the most enigmatic extinct elasmobranchs (i.e. sharks, skates and rays) for nearly two centuries. This widespread Cretaceous taxon is common in Albian to Campanian deposits from almost all continents. However, specimens mostly consist of isolated teeth or more or less complete dentitions, whereas cranial and post-cranial skeletal elements are very rare. Here we describe newly discovered material from the early Late Cretaceous of Mexico, including complete articulated specimens with preserved body outline, which reveals crucial information on the anatomy and systematic position of Ptychodus. Our phylogenetic and ecomorphological analyses indicate that ptychodontids were high-speed (tachypelagic) durophagous lamniforms (mackerel sharks), which occupied a specialized predatory niche previously unknown in fossil and extant elasmobranchs. Our results support the view that lamniforms were ecomorphologically highly diverse and represented the dominant group of sharks in Cretaceous marine ecosystems. Ptychodus may have fed predominantly on nektonic hard-shelled prey items such as ammonites and sea turtles rather than on benthic invertebrates, and its extinction during the Campanian, well before the end-Cretaceous crisis, might have been related to competition with emerging blunt-toothed globidensine and prognathodontine mosasaurs.

Smelly shark, smelly ray: what is infecting you?

AIMS: Although elasmobranchs are consumed worldwide, bacteriological assessments for this group are still sorely lacking. In this context, this study assessed bacteria of sharks and rays from one of the most important landing ports along the Rio de Janeiro coast. METHODS AND RESULTS: Bacteria were isolated from the cloacal swabs of the sampled elasmobranchs. They were cultured, and Vibrio, Aeromonas, and Enterobacterales were isolated and identified. The isolated bacteria were then biochemically identified and antimicrobial susceptibility assays were performed. Antigenic characterizations were performed for Salmonella spp. and Polymerase Chain Reaction (PCR) assays were performed to identify Escherichia coli pathotypes. Several bacteria of interest in the One Health context were detected. The most prevalent Enterobacterales were Morganella morganii and Citrobacter freundii, while Vibrio harveyi and Vibrio fluvialis were the most prevalent among Vibrio spp. and Aeromonas allosacharophila and Aeromonas veronii bv. veronii were the most frequent among Aeromonas spp. Several bacteria also displayed antimicrobial resistance, indicative of Public Health concerns. A total of 10% of Vibrio strains were resistant to trimethoprim-sulfamethoxazole and 40% displayed intermediate resistance to cefoxitin. Salmonella enterica strains displayed intermediate resistance to ciprofloxacin, nalidixic acid and streptomycin. All V. cholerae strains were identified as non-O1/non-O139. The detected E. coli strains did not exhibit pathogenicity genes. This is the first study to perform serology assessments for S. enterica subsp. enterica isolated from elasmobranchs, identifying the zoonotic Typhimurium serovar. Salmonella serology evaluations are, therefore, paramount to identify the importance of elasmobranchs in the epidemiological salmonellosis chain. CONCLUSIONS: The detection of several pathogenic and antibiotic-resistant bacteria may pose significant Public Health risks in Brazil, due to high elasmobranch consumption rates, indicating the urgent need for further bacteriological assessments in this group.

Comparative Proteome Analysis of Four Stages of Spermatogenesis in the Small-Spotted Catshark (Scyliorhinus canicula), Using High-Resolution NanoLC-ESI-MS/MS.

Spermatogenesis is a highly specialized process of cell proliferation and differentiation leading to the production of spermatozoa from spermatogonial stem cells. Due to its testicular anatomy, Scyliorhinus canicula is an interesting model to explore stage-based changes in proteins during spermatogenesis. The proteomes of four testicular zones corresponding to the germinative niche and to spermatocysts (cysts) with spermatogonia (zone A), cysts with spermatocytes (zone B), cysts with young spermatids (zone C), and cysts with late spermatids (zone D) have been analyzed by nanoLC-ESI-MS/MS. Gene ontology and KEGG annotations were also performed. A total of 3346 multiple protein groups were identified. Zone-specific protein analyses highlighted RNA-processing, chromosome-related processes, cilium organization, and cilium activity in zones A, D, C, and D, respectively. Analyses of proteins with zone-dependent abundance revealed processes related to cellular stress, ubiquitin-dependent degradation by the proteasome, post-transcriptional regulation, and regulation of cellular homeostasis. Our results also suggest that the roles of some proteins, such as ceruloplasmin, optineurin, the pregnancy zone protein, PA28ß or the Culling-RING ligase 5 complex, as well as some uncharacterized proteins, during spermatogenesis could be further explored. Finally, the study of this shark species allows one to integrate these data in an evolutionary context of the regulation of spermatogenesis. Mass spectrometry data are freely accessible via iProX-integrated Proteome resources (https://www.iprox.cn/) for reuse purposes.

Head anatomy of a lantern shark wet-collection specimen (Chondrichthyes: Etmopteridae).

In this study, we apply a two-step (untreated and soft tissue stained) diffusible iodine-based contrast-enhanced micro-computed tomography array to a wet-collection Lantern Shark specimen of Etmopterus lucifer. The focus of our scanning approach is the head anatomy. The unstained CT data allow the imaging of mineralized (skeletal) tissue, while results for soft tissue were achieved after staining for 120 h in a 1% ethanolic iodine solution. Three-dimensional visualization after the segmentation of hard as well as soft tissue reveals new details of tissue organization and allows us to draw conclusions on the significance of organs in their function. Outstanding are the ampullae of Lorenzini for electroreception, which appear as the dominant sense along with the olfactory system. Corresponding brain areas of these sensory organs are significantly enlarged as well and likely reflect adaptations to the lantern sharks' deep-sea habitat. While electroreception supports the capture of living prey, the enlarged olfactory system can guide the scavenging of these opportunistic feeders. Compared to other approaches based on the manual dissection of similar species, CT scanning is superior in some but not all aspects. For example, fenestrae of the cranial nerves within the chondrocranium cannot be identified reflecting the limitations of the method, however, CT scanning is less invasive, and the staining is mostly reversible and can be rinsed out.

Myxidium tunisiensis n. sp. (Myxosporea: Myxidiidae) infecting the rough skate Raja radula Delaroche, 1908 (Rajiformes: Rajidae) from North East Tunisia.

A new marine myxosporean, Myxidium tunisiensis n. sp. (Myxosporea: Myxidiidae), is described infecting the gallbladder of the rough skate Raja radula Delaroche, 1908 (Chondrichthyes: Rajidae) harvested from the Bay of Bizerte, Tunisia. Observed stages of the parasite were floating free in the bile. Plasmodia, disporic or polysporic, contain disporic pansporoblasts measuring 23.0-27.2 µm in maximum diameter and 21.0-25.5 µm in minimum diameter. Mature myxospores, which are elongated, typically fusiform measured 18.6 ± 1.7 (17.2-22.0) µm in length, 8.6 ± 1.2 (7.2-11.2) µm in width and 8.2 ± 0.9 (6.4-10.1) in thickness. Myxospore valves are symmetrical with 9-10 longitudinal striations. Two polar capsules, pyriform, were equal in size, measuring 5.8 ± 0.4 (5.2-6.7) µm in length and 3.1 ± 0.2 (3.0-3.2) µm in width, with 5-6 polar filament turns. The infection was prevalent throughout the year with the highest prevalence in autumn (81.8%) and the lowest in winter (32.3%). The seasonal prevalence of M. tunisiensis n. sp. in the Bay of Bizerte appeared to fluctuate according to seawater temperature. The morphological and morphometrical differences with all Myxidium spp. described so far, complemented by molecular data, prove clearly that the present species is a new member of the genus Myxidium. The SSU rDNA phylogenetic analyses revealed that Myxidium tunisiensis n. sp. clustered within the oligochaete-gallbladder clade in a basal position to the marine subclade of Myxidium baueri and Myxidium coryphaenoideum. While the new species branching pattern is stable in the ML and the BI analyses, it changed in the MP analysis. This is the first molecular study of a Myxidium species infecting an elasmobranch host inhabiting the Mediterranean Sea.

Birth and growth of the shortnose guitarfish Zapteryx brevirostris (Müller & Henle, 1841) (Chondrichthyes, Rhinobatidae) in captivity.

Endemic to the south-west Atlantic Ocean, the shortnose guitarfish (Zapteryx brevirostris) is a small species, classified as endangered by the IUCN. Although reproduction in captivity has been successful for some species, a range of factors can limit the success of captive breeding programmes for elasmobranchs. In Brazil, the Ubatuba Aquarium was the first public aquarium to reproduce small-sized elasmobranchs. Since 2018, at least five parturition events have been recorded for Z. brevirostris at the institution. From a total of 13 live neonates that rearing was attempted, the mean ± standard error of weight, total length (TL) and disc width at birth were 17.47 ± 1.6 g, 13.25 ± 0.7 cm and 6.53 ± 0.2 cm, respectively. The mean weight as well as mean TL were higher for females at all births, with 26.15 g and 15.07 cm for females in comparison with 17.09 g and 13.94 cm for males. Considering the increasing risk of extinction that the species is facing, age and growth studies are fundamental for the success of conservation, improving the knowledge of Z. brevirostris life cycle so that a more efficient and sustainable management can be carried out. This study provides important data, as well as directions for captive breeding of the species.

Nonlethally assessing elasmobranch ontogenetic shifts in energetics.

Body condition is an important proxy for the overall health and energetic status of fishes. The classically used Fulton's condition factor requires length and mass measurements, but mass can be difficult to obtain in large species. Girth measurements can replace mass for wild pelagic sharks. However, girth-calculated condition has not been validated against Fulton's condition factor intraspecifically, across ontogeny or reproduction, or in a controlled setting. We used the epaulette shark (Hemiscyllium ocellatum), because they are amenable to captive reproduction, to track fine-scale body condition changes across life stages, oviparous reproduction and between condition indices. We measured four girths, total length and mass of 16 captive epaulette sharks across 1 year and tracked female reproduction daily. We also collected length and mass data from an additional 72 wild-caught sharks and 155 sharks from five previous studies and two public aquaria to examine the relationship between length and mass for this species. Even though data were derived from a variety of sources, a predictable length-mass relationship (R2 = 0.990) was achievable, indicating that combining data from a variety of sources could help overcome knowledge gaps regarding basic life history characteristics. We also found that condition factor decreased during early life stages, then increased again into adulthood, with predictable changes across the female reproductive cycle. Finally, we determined that both Fulton's and girth condition analyses were comparable. Outcomes from this study uniquely provide body condition changes across the complete life history, including fine-scale female reproductive stages, and validate the use of girths as a nonlethal whole-organism energetic assessment for fishes.

Vision in sharks and rays: Opsin diversity and colour vision.

The visual sense of elasmobranch fishes is poorly studied compared to their bony cousins, the teleosts. Nevertheless, the elasmobranch eye features numerous specialisations that have no doubt facilitated the diversification and evolutionary success of this fascinating taxon. In this review, I highlight recent discoveries on the nature and phylogenetic distribution of visual pigments in sharks and rays. Whereas most rays appear to be cone dichromats, all sharks studied to date are cone monochromats and, as a group, have likely abandoned colour vision on multiple occasions. This situation in sharks mirrors that seen in other large marine predators, the pinnipeds and cetaceans, which leads us to reassess the costs and benefits of multiple cone pigments and wavelength discrimination in the marine environment.

Review of the parasitic nematodes of marine fishes from off the American continent.

The ichthyofauna of the Atlantic and Pacific coasts off the American continent is very rich. Consequently, a high biodiversity of nematodes parasitizing these vertebrates is also expected. Currently, data on nematode parasites of marine fish off the Americas are fragmented. A review of all adult nematode species reported parasitizing marine fish from off the American continent is herein presented, as well as comments on their patterns of diversity, life cycles and advances in the taxonomic and phylogenetic knowledge. A total of 209 valid species, 19 species inquirendae and 6 dubious records have been recorded, the majority from the fish taxa Eupercaria and Perciformes. The families Sciaenidae, Serranidae and Lutjanidae, as well as the tropical and temperate Atlantic waters, exhibited the highest records of parasitic nematodes. The Cucullanidae, Philometridae and Cystidicolidae were the most speciose families of nematodes, which may be related to technological advances and relatively recent efforts of taxonomists, resulting in description of new taxa and the resolution of taxonomic problems. Numerous taxonomic questions still need resolution and, even though genetic data have been important for this process, the database is very scarce. This is the first review on all currently known nematode species parasitizing marine fish off the Americas and may serve as an important basis of reference for future approaches on these organisms.

Morphological abnormalities in seven American round ray specimens: A review of America's batomorph anomalies.

Although morphological abnormalities in several rays and skate species around the American continents have frequently mentioned, their numbers are unknown. The present work record morphological abnormalities in four Urotrygonidae species. Two anophthalmic specimens were detected (Urotrygon microphthalmum and Urobatis halleri). Two individuals lacked caudal fins (Urobatis maculatus and Urotrygon chilensis). Two round rays showed incomplete fusion of the pectoral fin to the head (U. microphthalmum and U. chilensis). Vertebral compression and fusion were found in a 6-year-old female Urotrygon rogersi. In addition, 118 abnormal batomorph specimens were gathered from the available bibliography, spanning the last six decades (1959-2021). Amblyraja doellojuradoi was the species with the highest number of abnormalities (18). The most common anomaly was an incomplete fusion of the pectoral fin with the head. Since 2010, at least 30 anomalous batomorphs have been recorded every 5 years. Sixty-nine abnormal specimens occurred in the Northern Hemisphere (1.00-60.00 N). The Cortezian (Pacific) and Southeastern Brazil (Atlantic) marine ecoregions stood out with the highest number of these specimens. Mexico recorded 58 anomalous specimens, followed by Brazil (n = 36). Biological, abiotic and anthropogenic factors are probably the leading causes. However, additional studies are necessary to elucidate these speculations.

Ontogenetic development of the holocephalan dentition: Morphological transitions of dentine in the absence of teeth.

Among the cartilaginous fishes (Chondrichthyes), the Holocephali are unique in that teeth are absent both in ontogeny and adult regenerative growth. Instead, the holocephalan dentition of ever-growing nonshedding dental plates is composed of dentine, trabecular in arrangement, forming spaces into which a novel hypermineralized dentine (whitlockin) is deposited. These tissue features form a variety of specific morphologies as the defining characters of dental plates in the three families of extant holocephalans. We demonstrate how this morphology changes through ontogenetic development with continuity between morphologies, through successive growth stages of the dentition represented by the dental plate. For example, rod-shaped whitlockin appears early, later transformed into the tritoral pad, including a regular arrangement of vascular canals and whitlockin forming with increasing mineralization (95%-98%). While the tritoral pads develop lingually, stacks of individual ovoids of whitlockin replace the rods in the more labial parts of the plate, again shaped by the forming trabecular dentine. The ability to make dentine into new, distinctive patterns is retained in the evolution of the Holocephali, despite the lack of teeth forming in development of the dentition. We propose that developmentally, odontogenic stem cells, retained through evolution, control the trabecular dentine formation within the dental plate, and transition to form whitlockin, throughout lifetime growth. Our model of cellular activity proposes a tight membrane of odontoblasts, having transformed to whitloblasts, that can control active influx of minerals to the rapidly mineralizing dentine, forming whitlockin. After the reduced whitloblast cells transition back to odontoblasts, they continue to monitor the levels of minerals (calcium, phosphate and magnesium) and at a slower rate of growth in the peritubate 'softer' dentine. This model explains the unique features of transitions within the holocephalan dental plate morphology.

Mercury in neonatal and juvenile blacktip sharks (Carcharhinus limbatus). Part I: exposure assessment.

Elasmobranchs are exposed to mercury (Hg) through a variety of pathways in the environment. This study assessed maternal offloading and diet-based Hg exposure for neonatal and juvenile blacktip sharks (Carcharhinus limbatus) from Charlotte Harbor located along southwest Florida's coast, a recognized Hg hotspot. Neonates (n = 57) had highest total Hg (THg) concentrations in the kidney (0.56 ± 0.26 mg kg-1; n = 38) and muscle (0.53 ± 0.17 mg kg-1; n = 57), followed by liver (0.31 ± 0.11 mg kg-1; n = 38), and blood (0.05 ± 0.033 mg kg-1; n = 57). Juveniles (n = 13) exhibited a different distribution with highest THg in the liver (0.868 ± 0.54 mg kg-1; n = 6), followed by the muscle (0.84 ± 0.28 mg kg-1; n = 13), kidney (0.55 ± 0.22 mg kg-1; n = 6), and blood (0.11 ± 0.04 mg kg-1; n = 11). The distribution of THg among tissues and liver-to-muscle ratios indicated that Hg originated primarily from maternal offloading in neonates, whereas juveniles continued to accumulate Hg through dietary exposure post-parturition. Additionally, comparisons between results of the present study and previous Florida blacktip shark surveys suggested that Hg levels have not declined in southwest Florida estuaries for over two decades.

The elasmobranchs of Malpelo Flora and Fauna Sanctuary, Colombia.

Management and conservation actions in marine-protected areas require baselines for monitoring threatened marine fauna such as elasmobranchs. This article provides evidence of the occurrence of 34 species of elasmobranchs (21 sharks and 13 batoids) in the Malpelo Flora and Fauna Sanctuary, Colombia, including five new records of sharks and three of rays. From 1987 to 2021, new records were obtained by underwater visual census using SCUBA, manned submersibles and deep-ocean cameras to depths of up to 2211 m. Of the recorded species, 21 are considered as threatened taxa (64%) by the IUCN, making the Malpelo Flora and Fauna Sanctuary an essential conservation area for this highly threatened group of species.

Additional Pressure to a Threatened Species: High Persistent Organic Pollutant Concentrations in the Tropical Estuarine Batoid Gymnura altavela.

Persistent organic pollutants (POPs) were determined in Gymnura altavela liver samples from one of the most impacted estuarine environments in the Atlantic Ocean (Guanabara Bay, Brazil). PCBs were the predominant compounds (91.2%) averaging 6773 ± 4659 ng.g-1 lipid weight (lw). DDT, Mirex, HCH and HCB concentrations were 633 ± 715, 6.2 ± 9.3, 3.4 ± 1.4 and 0.5 ± 0.5 ng.g-1 lw, respectively. Recent DDT input in the estuary was minimum, as verified by the p,p'-DDE/∑DDT ratio (0.67). The prevalence of industrial compounds was highlighted by the ΣDDT/ΣPCB ratio (0.08). A negative correlation detected between disc width and PCBs, Mirex and HCH concentrations may be a consequence of maternal offloading. High POP concentrations in G. altavela liver samples from Guanabara Bay suggests that PCBs, in particular, may pressure the conservation of this threatened with extinction tropical estuarine batoid species.

First evidence of a functional spiracle in stem chondrichthyan acanthodians, with the oldest known elastic cartilage.

Spiracles are a general character of gnathostomes (jawed fishes), being present in antiarch placoderms, commonly regarded as the most basal gnathostome group. The presence of spiracular tubes in acanthodians has been deduced from grooves on the neurocranium of the derived acanthodiform Acanthodes bronni from the Permian of Germany, but until now these tubes were presumed to lack an external opening, rendering them non-functional. Here we describe the external spiracular elements in specimens of the Middle Devonian acanthodiforms Cheiracanthus murchisoni, Cheiracanthus latus and Mesacanthus pusillus from northern Scotland, and the internal structure of these elements in C. murchisoni, demonstrating that the spiracle in acanthodiforms differed from all known extant and extinct fishes in having paired cartilage-pseudobranch structures. This arrangement represents a transitional state between the presumed basal gnathostome condition with an unconstricted first gill slit (as yet not identified in any fossil) and the derived condition with a spiracle and a single pseudobranch derived from the posterior hemibranch of the mandibular arch. We identify the main tissue forming the pseudobranch as elastic cartilage, a tissue previously unrecorded in fossils.

Sharks, rays, and chimaeras of the Revillagigedo National Park: An update of new and confirmed records.

The Revillagigedo Archipelago National Park is the largest no-take Marine Protected Area in North America and it is essential for the conservation of globally endangered marine megafauna, including cetaceans, sea birds and cartilaginous fishes. The present paper reports the occurrence of 28 chondrichthyan species, including 19 sharks, seven batoids and two chimaeras in the Revillagigedo Archipelago, comprising eight new records and a brief revision of confirmed species to date.

Myosin heavy chain isoforms in the myocardium of the atrioventricular junction of Scyliorhinus canicula (Chondrichthyes, Carcharhiniformes).

The atrioventricular junction of the fish heart, namely the segment interposed between the single atrium and the single ventricle, has been studied anatomically and histologically in several chondrichthyan and teleost species. Nonetheless, knowledge about myosin heavy chain (MyHC) in the atrioventricular myocardium remains scarce. The present report is the first one to provide data on the MyHC isoform distribution in the myocardium of the atrioventricular junction in chondrichthyans, specifically in the lesser spotted dogfish, Scyliorhinus canicula, a shark species whose heart reflects the primitive cardiac anatomical design in gnathostomes. Hearts from five dogfish were examined using histochemical and immunohistochemical techniques. The anti-MyHC A4.1025 antibody was used to detect differences in the occurrence of MyHC isoforms in the dogfish, as the fast-twitch isoforms MYH2 and MYH6 have a higher affinity for this antibody than the slow-twitch isoforms MYH7 and MYH7B. The histochemical findings show that myocardium of the atrioventricular junction connects the trabeculated myocardium of the atrium with the trabeculated layer of the ventricular myocardium. The immunohistochemical results indicate that the distribution of MyHC isoforms in the atrioventricular junction is not homogeneous. The atrial portion of the atrioventricular myocardium shows a positive reactivity against the A4.1025 antibody similar to that of the atrial myocardium. In contrast, the ventricular portion of the atrioventricular junction is not labelled, as is the case with the ventricular myocardium. This dual condition suggests that the myocardium of the atrioventricular junction has two contraction patterns: the myocardium of the atrial portion contracts in line with the atrial myocardium, whereas that of the ventricular portion follows the contraction pattern of the ventricular myocardium. Thus, the transition of the contraction wave from the atrium to the ventricle may be established in the atrioventricular segment because of its heterogeneous MyHC isoform distribution. The findings support the hypothesis that a distinct MyHC isoform distribution in the atrioventricular myocardium enables a synchronous contraction of inflow and outflow cardiac segments in vertebrates lacking a specialized cardiac conduction system.

The early elasmobranch Phoebodus: phylogenetic relationships, ecomorphology and a new time-scale for shark evolution.

Anatomical knowledge of early chondrichthyans and estimates of their phylogeny are improving, but many taxa are still known only from microremains. The nearly cosmopolitan and regionally abundant Devonian genus Phoebodus has long been known solely from isolated teeth and fin spines. Here, we report the first skeletal remains of Phoebodus from the Famennian (Late Devonian) of the Maïder region of Morocco, revealing an anguilliform body, specialized braincase, hyoid arch, elongate jaws and rostrum, complementing its characteristic dentition and ctenacanth fin spines preceding both dorsal fins. Several of these features corroborate a likely close relationship with the Carboniferous species Thrinacodus gracia, and phylogenetic analysis places both taxa securely as members of the elasmobranch stem lineage. Identified as such, phoebodont teeth provide a plausible marker for range extension of the elasmobranchs into the Middle Devonian, thus providing a new minimum date for the origin of the chondrichthyan crown-group. Among pre-Carboniferous jawed vertebrates, the anguilliform body shape of Phoebodus is unprecedented, and its specialized anatomy is, in several respects, most easily compared with the modern frilled shark Chlamydoselachus. These results add greatly to the morphological, and by implication ecological, disparity of the earliest elasmobranchs.