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To evaluate scale morphological variability (shape, size, topological macro- and microstructures, ornamentation patterns) of cyprinid fishes, nine species of the genus Garra were selected and their scales from five different body regions were studied by using light and scanning electron microscopy. The scales of the examined species were thin with a central or antero-centrally positioned focus, with no cteni in the posterior part. In addition to these typical characteristics, some morphological variation was observed in the overall shape (irregular round, true oval, round-triangular, irregular pentagonal, hexagonal, irregular hexagonal, pentagonal, ovoid), and the focus shape. These variations were mainly dependent on the fish lengths and the flank region. Morphological analysis clustered the examined species into two distinct groups. Group I consisted of G. amirhosseini and G. gymnothorax, while Group II has three subgroups, which include G. persica and G. mondica (subgroup I), G. meymehensis, G. rossica, G. nudiventris, G. hormuzensis (subgroup II), and G. rufa (subgroup III). The grouping of the studied Garra species based on the current scale morphological characters and the molecular data was only consistent for G. rossica and G. nudiventris. In addition, in the phylogenetic tree, G. persica, G. mondica, G. amirhosseini, and G. hormuzensis formed a distinct clade. However, these species did not represent close relationships in the dendrogram obtained from the scale morphology. A possible explanation why the grouping of the studied Garra species based on their scale morphological characters does not match their phylogenetic relationships is that most of the scale morphological traits vary depending on the fish size and the location of the scales on the flank. Therefore, except for some traits, that is, central or antero-centrally positioned focus, having no cteni, a specific sectioned form so called "tetra-sectioned" type, the other examined variables are not useful enough to be used in the taxonomic study of the examined cyprinid fishes. Therefore, scale morphological characters should be used carefully for taxonomic purposes. RESEARCH HIGHLIGHTS: Shape, size, topological macro- and microstructures, and ornamentation patterns of nine species of the genus Garra were studied by using light and scanning electron microscopy. The scales were thin with a central or antero-centrally positioned focus, with no cteni in the posterior part. Morphological variation was observed in the overall shape (irregular round, true oval, round-triangular, irregular pentagonal, hexagonal, irregular hexagonal, pentagonal, and ovoid), and the focus shape. The grouping of Garra species based on the current scale morphological characters and the molecular data was only consistent for G. rossica and G. nudiventris., (© 2024 Wiley Periodicals LLC.)

This study investigated the morphological characteristics of scales in six Cyprinion species, using light and scanning electron microscopy focusing on key features such as scale type, key scales, lateral line scales, radius/radii, rostral margin, focus, circuli, lepidonts, tubercles, and scale indices. The research analyzed the scales using ultramicroscopy and light microscopy imaging, categorizing them based on size classes and body regions. The morphological variations in scale characteristics were examined across different species, regions, and size classes. Notable findings included the tetra-sectioned form of scales, representing a unique characteristic of the Cyprinion genus. Morphological changes in scale features were observed with fish growth, particularly in the overall shape, focus shape, and size. Quantitative analysis revealed variations in average relative scale length and width among different species, regions, and size classes. The study utilized canonical discriminant analysis for multivariate assessment, classifying the species into distinct groups based on morphometric indices. The findings contribute to the understanding of scale morphology in Cyprinion species and exploring morphological variation between the examined species., Competing Interests: Conflict of Interest: The authors declare that they have no known competing financial interests or personal relationships that could have appeared to influence the work reported in this paper., (© The Author(s) 2024. Published by Oxford University Press on behalf of the Microscopy Society of America. All rights reserved. For commercial re-use, please contact reprints@oup.com for reprints and translation rights for reprints. All other permissions can be obtained through our RightsLink service via the Permissions link on the article page on our site—for further information please contact journals.permissions@oup.com.)

Medical imaging techniques such as digital radiography and ultrasonography are non-invasive and provide precise results for examining internal organs and structures within fish. Their effectiveness can be further enhanced by using body parts like scales as markers for the organs beneath them. This study utilized the number of scales as landmarks in digital radiography and ultrasonography to non-invasively evaluate the muscles, bones, and images of internal and reproductive organs of common carp (Cyprinus carpio). Digital radiography was performed in the dorsoventral and lateral views of the fish, whereas ultrasonography was conducted in longitudinal and transverse views on sequence scale numbers with brightness and colour Doppler-modes. Digital radiography of the common carp revealed the whole-body morphology, including the bony parts from the head, pectoral fins, dorsal fins, pelvic fins, anal fins, and vertebrae to the tail that appeared radiopaque. Internal organs were also observed, with the swim bladder and heart appeared radiolucent, while the intestines, liver, testes, and ovaries appeared radiopaque. Ultrasonography in brightness mode displayed the digestive organs, reproductive organs, and muscle thickness. Additionally, colour Doppler mode demonstrated blood flow within the heart's ventricle., (© 2024 Wiley‐VCH GmbH. Published by John Wiley & Sons Ltd.)

Many actinopterygian fish groups, including fossil and extant polypteriforms and lepisosteiforms, fossil halecomorphs, and some basal teleosts, have stout bony scales covered by layers of ganoin-an enamel layer ornamented with minute tubercles. Ganoid scales preserve well as disarticulated remains and notably constitute most of the fossil record for polypteriform in both South America and Africa. Based on two variables (tubercle size and distance between tubercles), some authors reported that the ganoin tubercle ornamentation in these scales is constant within a species and differs between species and allows distinguishing species or at least groups of species. However, despite its promising potential for assessing polypteriform paleodiversity, this tool has remained unused, probably because the variables are not well defined, and intraspecific variation does not seem to have been considered. To address this gap, we aimed to test the intraspecific and intra-individual variation in the ornamentation of ganoid scales in the type species Polypterus bichir. We propose three different parameters to describe the tubercle ornamentation: the distance between contiguous tubercles centers, their density, and their relative spatial organization. With these parameters, we investigate the variation in ganoin ornamentation among four specimens and across different regions of the body. Our results show that the distribution of the tubercles is highly variable within a same species, regardless of the body region, and sometimes even between different sectors of a same scale. Moreover, the variation observed in P. bichir overlaps with the distribution described in the literature for several extant and fossil species. Thus far, the ornamentation of ganoid scales is not a reliable diagnostical feature for polypterids., (© 2024 Fisheries Society of the British Isles.)

Arboreality has evolved in all major vertebrate lineages and is often associated with morphological adaptations and increased diversification concomitant with accessing novel niche space. In squamate reptiles, foot, claw, and tail morphology are well-studied adaptations shown to be associated with transitions to arboreality. Here, we examined a less well understood trait-the keeled scale-in relation to microhabitat, climate, and diversification dynamics across a diverse lizard radiation, Agamidae. We found that the ancestral agamid had keeled dorsal but not ventral scales; further, dorsal and ventral keels are evolutionarily decoupled. Ventral keeled scales evolved repeatedly in association with arboreality and may be advantageous in reducing wear or by promoting interlocking when climbing. We did not find an association between keeled scales and diversification, suggesting keels do not allow finer-scale microhabitat partitioning observed in other arboreal-associated traits. We additionally found a relationship between keeled ventral scales and precipitation in terrestrial species where we posit that the keels may function to reduce scale degradation. Our results suggest that keeled ventral scales facilitated transitions to arboreality across agamid lizards, and highlight a need for future studies that explore their biomechanical function in relation to microhabitat and climate.

Fossil feathers have transformed our understanding of integumentary evolution in vertebrates. The evolution of feathers is associated with novel skin ultrastructures, but the fossil record of these changes is poor and thus the critical transition from scaled to feathered skin is poorly understood. Here we shed light on this issue using preserved skin in the non-avian feathered dinosaur Psittacosaurus. Skin in the non-feathered, scaled torso is three-dimensionally replicated in silica and preserves epidermal layers, corneocytes and melanosomes. The morphology of the preserved stratum corneum is consistent with an original composition rich in corneous beta proteins, rather than (alpha-) keratins as in the feathered skin of birds. The stratum corneum is relatively thin in the ventral torso compared to extant quadrupedal reptiles, reflecting a reduced demand for mechanical protection in an elevated bipedal stance. The distribution of the melanosomes in the fossil skin is consistent with melanin-based colouration in extant crocodilians. Collectively, the fossil evidence supports partitioning of skin development in Psittacosaurus: a reptile-type condition in non-feathered regions and an avian-like condition in feathered regions. Retention of reptile-type skin in non-feathered regions would have ensured essential skin functions during the early, experimental stages of feather evolution., (© 2024. The Author(s).)

The study of morphological characteristics and growth information in fish scales is a crucial component of modern fishery biological research, while it has been less studied in fossil materials. This paper presents a detailed morphological description and growth analysis of a fossil ctenoid scale obtained from the Upper Cretaceous Campanian lacustrine deposits in northeastern China. The morphological features of this fossil scale are well-preserved and consistent with the structures found in ctenoid scales of extant fish species and display prominent ring ornamentation radiating outward from the central focus, with grooves intersecting the rings. A comparative analysis of the morphological characteristics between the fossil ctenoid scale and those well-studied extant fish Mugilidae allows us to explore the applicability of modern fishery biological research methods to the field of fossil scales. The scale length, scale width, the vertical distance from the focus to the apex of the scale, and the total number of radii have been measured. The age of the fish that possessed this ctenoid scale has been estimated by carefully counting the annuli, suggesting an age equal to or more than seven years. The distribution of growth rings on the scale potentially reflects the warm paleoclimatic condition and fish-friendly paleoenvironment prevalent during that period. This paper, moreover, serves as a notable application of fishery biological methods in the examination of fossil materials., Competing Interests: The authors have declared that no competing interests exist., (Copyright: © 2024 Liu et al. This is an open access article distributed under the terms of the Creative Commons Attribution License, which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited.)

Corneous skin appendages are not only common and diverse in crown-group amniotes but also present in some modern amphibians. This raises the still unresolved question of whether the ability to form corneous skin appendages is an apomorphy of a common ancestor of amphibians and amniotes or evolved independently in both groups. So far, there is no palaeontological contribution to the issue owing to the lack of keratin soft tissue preservation in Palaeozoic anamniotes. New data are provided by a recently discovered ichnofossil specimen from the early Permian of Poland that shows monospecific tetrapod footprints associated with a partial scaly body impression. The traces can be unambiguously attributed to diadectids and are interpreted as the globally first evidence of horned scales in tetrapods close to the origin of amniotes. Taking hitherto little-noticed scaly skin impressions of lepospondyl stem amniotes from the early Permian of Germany into account, the possibility has to be considered that the evolutionary origin of epidermal scales deeply roots among anamniotes.

A fundamental gap in the study of the origin of limbed vertebrates lies in understanding the morphological and functional diversity of their closest relatives. Whereas analyses of the elpistostegalians Panderichthys rhombolepis, Tiktaalik roseae and Elpistostege watsoni have revealed a sequence of changes in locomotor, feeding and respiratory structures during the transition 1-9 , an isolated bone, a putative humerus, has controversially hinted at a wider range in form and function than now recognized 10-14 . Here we report the discovery of a new elpistostegalian from the Late Devonian period of the Canadian Arctic that shows surprising disparity in the group. The specimen includes partial upper and lower jaws, pharyngeal elements, a pectoral fin and scalation. This new genus is phylogenetically proximate to T. roseae and E. watsoni but evinces notable differences from both taxa and, indeed, other described tetrapodomorphs. Lacking processes, joint orientations and muscle scars indicative of appendage-based support on a hard substrate 13 , its pectoral fin shows specializations for swimming that are unlike those known from other sarcopterygians. This unexpected morphological and functional diversity represents a previously hidden ecological expansion, a secondary return to open water, near the origin of limbed vertebrates., (© 2022. The Author(s).)

During metamorphosis, the wings of a butterfly sprout hundreds of thousands of scales with intricate microstructures and nano-structures that determine the wings' optical appearance, wetting characteristics, thermodynamic properties, and aerodynamic behavior. Although the functional characteristics of scales are well known and prove desirable in various applications, the dynamic processes and temporal coordination required to sculpt the scales' many structural features remain poorly understood. Current knowledge of scale growth is primarily gained from ex vivo studies of fixed scale cells at discrete time points; to fully understand scale formation, it is critical to characterize the time-dependent morphological changes throughout their development. Here, we report the continuous, in vivo, label-free imaging of growing scale cells of Vanessa cardui using speckle-correlation reflection phase microscopy. By capturing time-resolved volumetric tissue data together with nanoscale surface height information, we establish a morphological timeline of wing scale formation and gain quantitative insights into the underlying processes involved in scale cell patterning and growth. We identify early differences in the patterning of cover and ground scales on the young wing and quantify geometrical parameters of growing scale features, which suggest that surface growth is critical to structure formation. Our quantitative, time-resolved in vivo imaging of butterfly scale development provides the foundation for decoding the processes and biomechanical principles involved in the formation of functional structures in biological materials., Competing Interests: The authors declare no competing interest., (Copyright © 2021 the Author(s). Published by PNAS.)

The males of more than 80% of the Lycaenidae species belonging to the tribe Polyommatini exhibit structural coloration on their dorsal wing surfaces. These colors have a role in reinforcement in prezygotic reproductive isolation. The species-specific colors are produced by the cellular self-assembly of chitin/air nanocomposites. The spectral position of the reflectance maximum of such photonic nanoarchitectures depends on the nanoscale geometric dimensions of the elements building up the nanostructure. Previous work showed that the coloration of male Polyommatus icarus butterflies in the Western and Eastern Palearctic exhibits a characteristic spectral difference (20 nm). We investigated the coloration and the de novo developed DNA microsatellites of 80 P. icarus specimens from Europe from four sampling locations, spanning a distance of 1621 km. Remarkably good concordance was found between the spectral properties of the blue sexual signaling color (coincident within 5 nm) and the population genetic structure as revealed by 10 microsatellites for the P. icarus species., (© 2021. The Author(s).)

Scales are skin appendages in fishes that evolutionarily predate feathers in birds and hair in mammals. Zebrafish scales are dermal in origin and develop during metamorphosis. Understanding regulation of scale development in zebrafish offers an exciting possibility of unraveling how the mechanisms of skin appendage formation evolved in lower vertebrates and whether these mechanisms remained conserved in birds and mammals. Here we have investigated the expression and function of twist 2/dermo1 gene - known for its function in feather and hair formation - in scale development and regeneration. We show that of the four zebrafish twist paralogues, twist2/dermo1 and twist3 are expressed in the scale forming cells during scale development. Their expression is also upregulated during scale regeneration. Our knockout analysis reveals that twist2/dermo1 gene functions in the maintenance of the scale shape and organization during development as well as regeneration. We further show that the expression of twist2/dermo1 and twist3 is regulated by Wnt signaling. Our results demonstrate that the function of twist2/dermo1 in skin appendage formation, presumably under regulation of Wnt signaling, originated during evolution of basal vertebrates., (Copyright © 2021 Elsevier B.V. All rights reserved.)

We investigated the geographic diversification of Plestiodon finitimus , which occurs in the central to northern parts of the Japanese Islands, based on a time-calibrated mitochondrial DNA (mtDNA) phylogeny and external morphological characters. The mtDNA phylogeny suggests that P. finitimus diverged from its sister species Plestiodon japonicus in western Japan 2.82-4.63 million years ago (MYA), which can be explained by geographic isolation due to the spread of sedimentary basins in the Pliocene. The primary intraspecific divergence was that between P. finitimus lineages in central and northeastern Japan 1.58-2.76 MYA, which could have been caused by the upliftings of major mountain ranges. In the northeastern lineage, mtDNA and morphological characters suggest a geographic differentiation between sub-lineages of the northwestern Tohoku District (α) and other areas (β). Although the sub-lineage β occurs in a disjunct geographic range, consisting of Hokkaido and the central to south of Tohoku, these areas are bridged by populations with intermediate characteristics along the Pacific side of northern Tohoku. Overall, the geographic variation in P. finitimus in northern Japan can be explained by an initial allopatric divergence of the sub-lineages α and β at 0.71-1.39 MYA, a recent northward expansion of the sub-lineage β, and subsequent secondary introgressive hybridization between the sub-lineages.

The normal and lateral line scales from above the head and four flank regions were investigated in two goatfishes by using light microscopy to evaluate the microscopic characteristics of scale surface ornamentation from different body parts of these fishes and to add new morphological data for their discrimination. The body scales were removed and digital images captured with a Dino-Lite digital camera connected to a Leica compound microscope. The presence of both transverse and longitudinal radii was a distinctive characteristic for the scales of studied goatfishes. The most distinctive features of the key scales discriminating the two goatfishes were the scale shape, the presence of irregular grooves in scale surface, the presence of transverse radii in the lateral fields, and the availability of lepidont at the anterior field. The lateral line scale ornamentations were almost similar in two species; however, the lateral line canal was branching in Parupeneus heptacanthus. Scale extension index indicated that in both species, the dimension of normal key scales tends to the right, while the dimension of lateral line scales tends to the left. This study accentuated that scales from different body regions may exhibit variation in surface ornamentations, that scale morphology could successfully be used to discriminate the examined goatfishes, and that the use of fish scale microscopy is an easy and relatively rapid method and does not require fish sacrifice. Since fish identification is essential for the conservation and management of species, the use of scale morphology to this purpose appears particularly promising., (© 2020 Wiley Periodicals LLC.)

The skin of the foot provides the interface between the bird and the substrate. The foot morphology involves the bone shape and the integument that is in contact with the substrate. The podotheca is a layer of keratinized epidermis forming scales that extends from the tarsometatarsus to the toe extremities. It varies in size, shape, amount of overlap and interacts with the degree of fusion of the toes (syndactyly). A study of toe shape and the podotheca provides insights on the adaptations of perching birds. Our analysis is based on micro-CT scans and scanning electron microscopy images of 21 species from 17 families, and includes examples with different orientations of the toes: zygodactyl (toes II and III forward), anisodactyl (toes II, III, and IV forward), and heterodactyl (toes III and IV forward). We show that in these three groups, the skin forms part of a perching adaptation that involves syndactyly to different degrees. However, syndactyly does not occur in Psittacidae that use their toes also for food manipulation. The syndactyly increases the sole surface and may reinforce adherence with the substrate. Scale shape and toe orientation are involved in functional adaptations to perch. Thus, both bone and skin features combine to form a pincer-like foot., (© 2020 Wiley Periodicals LLC.)

This study describes Characidium nambiquara, a new species from the upper rio Guaporé, rio Madeira basin, Brazil. The new species differs from most congeners by the presence of isthmus and area between the contralateral pectoral-fin bases completely naked. From congeners with some degree of scaleless ventral surface of the body C. nambiquara differs by having 10 circumpeduncular scales. The new species is also distinguished from congeners by the spotted colour pattern on body of the large-sized specimens and by having black dashes on all fins and conspicuous midlateral longitudinal dark stripe or conspicuous vertical bars absent. Characidium nambiquara further differs from most congeners by the presence of 34-36 pored scales on the lateral line, 3 horizontal scale rows above the lateral line and 3 horizontal scale rows from the lateral line to the midventral scale series. Remarks on intraspecific colour variation within the genus, not related to sexual dimorphism, are also provided., (© 2020 Fisheries Society of the British Isles.)

Chaetonotidae is the most diverse and widely distributed family of the order Chaetonotida (Gastrotricha) and includes both marine and freshwater species. Although the family is regarded as a sister taxon to the exclusively marine Xenotrichulidae, the type of environment, marine or freshwater, where Chaetonotidae originated is still not known. Here, we reconstructed the phylogeny of the family based on molecular sequence data and mapped both morphological and ecological characters to determine the ancestral environment of the first members of the family. Our results revealed that the freshwater genus Bifidochaetus is the earliest branching lineage in the paraphyletic Chaetonotidae (encompassing Dasydytidae and Neogosseidae). Moreover, we reconstructed Lepidochaetus-Cephalionotus clade as a monophyletic sister group to the remaining chaetonotids, which supports Kisielewski's morphological based hypothesis concerning undifferentiated type of body scales as a most primary character in Chaetonotidae. We also found that reversals to marine habitats occurred independently in different Chaetonotidae lineages, thus marine species in the genera Heterolepidoderma, Halichaetonotus, Aspidiophorus and subgenera Chaetonotus (Schizochaetonotus) or Chaetonotus (Marinochaetus) should be assumed as having secondarily invaded the marine environment. Character mapping revealed a series of synapomorphies that define the clade that includes Chaetonotidae (with Dasydytidae and Neogosseidae), the most important of which may be those linked to reproduction., (© The Willi Hennig Society 2020.)

Bryconops cyrtogaster, a poorly known species endemic from the Oyapock River at the border between French Guyana and Brazil, is redescribed herein based on examination of available type material, as well as newly collected material. Additionally, a new rheophilic species from the rio Jari rapids, lower Amazon basin, Brazil, is described. The two species belong to the subgenus Creatochanes and are unique among the congeneres for possessing a posteriorly positioned humeral blotch at the level of the sixth and seventh lateral line scales. They differ from each other by meristic and morphometric characters. The list of endemic species in the rio Jari basin is revised., (© 2020 The Fisheries Society of the British Isles.)

Puffers show good drag reduction performance during migration. It is worth noting that spines which are different from ordinary fish scales are densely distributed on the puffer skin. Here, the special morphological structure of puffer spines was observed using microscopy techniques, accurate contour models were established based on image processing techniques and curve fitting, then feature sizes were obtained. Based on the results, the nonsmooth surface was established by orthogonal test to simulate the flow field. In addition, the influence of spinal structure on boundary layer flow field and the drag reduction property of nonsmooth surface were further analyzed. The nonsmooth surface formed by spinal structure elements can effectively reduce the wall shear stress and Reynolds stress, and there was a special "climbing vortex" phenomenon, so as to reduce the surface viscous friction resistance and achieve drag reduction. Compared with the smooth surface, the drag reduction rate of the nonsmooth surface was 12.94% when the inflow velocity was 5 m/s, which revealed and verified the drag reduction performance of the spines of puffer skin. The results lay a foundation for further research and optimization of drag reduction ability of nonsmooth surface of bionic spines. HIGHLIGHTS: The contour of the spinous process was accurately reflected by the Fourier function. The spines of puffer skin have good drag reduction effect. There was a special "climbing vortex" phenomenon to explain the drag reduction property., (© 2020 Wiley Periodicals, Inc.)

The covering by scales of the wings of Lepidoptera contributes to multiple functions that are critical for their survival and reproduction. In order to gain a better understanding about their distribution, we have exhaustively studied 4 specimens of Colias crocea (Geoffroy, 1785). We have quantified the sources of variability affecting scale density. The results indicate that the scale covering of butterfly wings may be remarkably heterogeneous, and that the importance of the sources of variability differs between forewings and hindwings. Thus, in forewing the greatest variability occurs between sectors, while in the hindwings it occurs between sides, with a higher density of scales on the underside, considerably higher (almost 19%) than on the upperside. It seems likely that this difference has an adaptive value, as the hindwing underside is more exposed (in resting position) to predators. These results are in contrast with the generally accepted notion that scale covering is uniform and homogeneous. Moreover, the cover scale density is independent of the size of the specimen and therefore an average density of scales can be attributed to this species. According to our measurements C. crocea has 312 scales/mm 2 and the total number of scales per individual is about 520,000 on average., (Copyright © 2020 Elsevier Ltd. All rights reserved.)

The quest for new light-weight materials with superior mechanical properties is a goal of materials scientists and engineers worldwide. A promising route in this pursuit is drawing inspiration from nature to design and develop materials with enhanced properties. By emulating the graded mineral content and hierarchical structure of fish scales of the Arapaima gigas from the nano to macro scales, we were able to develop bioinspired laminated composites with improved impact resistance. Activated by the addition of nano-particles of Al 2 O 3 and nano-layers of TiN to a thermoplastic fiber substrate, new energy dissipation mechanisms operating at the nanoscale enhanced the energy absorption and stiffness of the bioinspired material. Remarkably, the newly developed materials are easily transferred to the industry with minimum associated manufacturing costs.

Recently, it has been shown that animals such as jumping spiders, birds, and butterflies have evolved ultra-black coloration comparable to the blackest synthetic materials. Of these, certain papilionid butterflies have reflectances approaching 0.2%, resulting from a polydisperse honeycomb structure. It is unknown if other ultra-black butterflies use this mechanism. Here, we examine a phylogenetically diverse set of butterflies and demonstrate that other butterflies employ simpler nanostructures that achieve ultra-black coloration in scales thinner than synthetic alternatives. Using scanning electron microscopy, we find considerable interspecific variation in the geometry of the holes in the structures, and verify with finite-difference time-domain modeling that expanded trabeculae and ridges, found across ultra-black butterflies, reduce reflectance up to 16-fold. Our results demonstrate that butterflies produce ultra-black by creating a sparse material with high surface area to increase absorption and minimize surface reflection. We hypothesize that butterflies use ultra-black to increase the contrast of color signals.

We describe the morphology of alar androconia and the female abdominal scent gland of Heliconius erato phyllis, Heliconius ethilla narcaea, and Heliconius besckei. Androconial scales of Heliconius, which are arranged in overlapping wing bands, release pheromones during courtship, probably through vibratory movements of male wings over the female to induce her to mate. An antiaphrodisiac is produced by glands located in the valves of the male and is transferred during copulation to the yellow dorsal abdominal sac present in the virgin female, causing this sac to emit a scent that reduces the attractiveness of the female for courtship with other males. Stereomicroscopy, SEM, and TEM analyses were conducted to describe the morphology of the internal and external scales and the external abdominal scent sac. The findings revealed different sizes of external androconial scales and an internal group of porous structural vesicles that are probably related to the preservation of internal space, reception and storage of secretions, and elimination of volatiles when the male is actively involved in courtship. Translucent projections on the female abdominal scent sac create open reservoirs for the reception, storage, and emission of antiaphrodisiac volatiles along with stink clubs. Male valve denticles vary in form and probably attach securely to the female sac during mating, thus ensuring secretion transfer. These features are discussed in the context of a comparative analysis., (© 2020 Wiley Periodicals, Inc.)

Using morphometric, meristic and qualitative characters, we confirmed a specific status of a clade discovered based on molecular data only. The newly described species Phoxinus krkae has a very local distribution in the upper Krka River but is easily morphologically distinguished from geographically neighbouring species with much wider ranges: Phoxinus lumaireul, Phoxinus marsilii and Phoxinus csikii. Phoxinus krkae is well differentiated in statistical analyses and differs from other species of Phoxinus in northern Adriatic Sea drainages and right-bank tributaries of the middle and lower River Danube by a combination of characters, none of them unique: caudal peduncle length 1.9-2.2 times minimum depth of caudal peduncle; upper lip not projecting beyond lower lip; rostral fold absent; total number of scales in lateral series 74-84; lateral line incomplete and interrupted, commonly 17-49 (< 60% of total number of scales in lateral series) with no or few canal segments on posterior body and caudal peduncle; scales on belly extending forward to over middle of distance between pelvic-fin base and pectoral-fin origin; patches of breast scales commonly connected by scattered scales or forming 1-3 regular rows of scales; total vertebrae commonly 38-39 (22 + 17, 22 + 16 or 21 + 17). The study reveals unrecognised morphological diversity within the group and provides the groundwork for future taxonomic study in Phoxinus in the Adriatic region and the Danube Basin., (© 2019 The Fisheries Society of the British Isles.)

The simultaneous use of the scale and otolith morphometry was assessed as a potential tool for the identification of Persian brown trout Salmo trutta stocks of the Lar Lake and five rivers from Lar Basin, Iran. Fourier coefficients (FC) and circularity, rectangularity, roundness, ellipticity and form factor shape indices (SI) were calculated for otolith and scale. Several SIs were significantly different among sites for both structures. Permutational multivariate analysis of variance revealed significant differences between several pairwise comparisons for otolith and scale (FCs and indices separately). Discriminant analysis showed otolith FCs (cross-classification rates: 25-86%) and SI (20-45%) appear to be a relatively acceptable tool to discriminate between several locations. Comparatively, the scale morphometry showed lower discriminatory power (FC = 3-65%; SI = 15-34%), with the exception of SI for Elarm River (60%), Kamardasht River (56%) and Lar Lake (75%). Cross-classification rates improved up to 100% when discriminate analysis incorporating all variables for otolith and scale was performed. The results showed a potential segregation between some water bodies, suggesting that the otolith and scale morphometry could be a useful tool to delimit S. trutta populations in relatively close freshwater environments., (© 2019 The Fisheries Society of the British Isles.)

The outermost surface of insect cuticle is a high-performance interface that provides wear protection, hydration, camouflage and sensing. The complex and inhomogeneous structure of insect cuticle imposes stringent requirements on approaches to elucidate its molecular structure and surface chemistry. Therefore, a molecular understanding and possible mimicry of the surface of insect cuticle has been a challenge. Conventional optical and electron microscopies as well as biochemical techniques provide information about morphology and chemistry but lack surface specificity. We here show that a near edge X-ray absorption fine structure microscope at the National Synchrotron Light Source can probe the surface chemistry of the curved and inhomogeneous cuticle of the African flower scarab. The analysis shows the distribution of organic and inorganic surface species while also hinting at the presence of aragonite at the dorsal protrusion region of the Eudicella gralli head, in line with its biological function.

Piaractus orinoquensis, a new species of serrasalmid fish, is described from the Orinoco River basin. The new species differs from congeners by having a slenderer body, relatively smaller head and snout, more compressed mid-body, fewer scales above and below the lateral line and diagnostic molecular characters in the coI mitochondrial gene region. We also provide a re-description of Piaractus brachypomus, restricting its geographic distribution to the Amazon River basin. Both species are economically important in their respective basins and need to be independently managed as distinct species., (© 2019 The Fisheries Society of the British Isles.)

The skin surface structure of squamate reptiles varies greatly among species, likely because it plays a key role in a range of tasks, such as camouflage, locomotion, self-cleaning, mitigation of water loss and protection from physical damage. Although we have foundational knowledge about squamate skin morphology, we still know remarkably little about how intraspecific variation in skin surface structure translates to functional variation. This gap in our understanding can be in part traced back to: (i) our lack of knowledge on how body size determines skin surface structure; and (ii) the lack of means to perform high-throughput and detailed analysis of the three-dimensional (3D) anatomy of reptilian skin surfaces in a non-destructive manner. To fill this gap, we explored the possibilities of a new imaging technique, termed gel-based stereo-profilometry, to visualize and quantify the 3D topography of reptilian skin surface structure. Using this novel approach, we investigated intra-specific and intra-individual variation in the skin surface morphology of a focal lizard species, Anolis cristatellus. We assessed how various characteristics of surface topography (roughness, skew and kurtosis) and scale morphology (area, height, width and shape) scale with body size across different body regions. Based on an ontogenetic series of A. cristatellus males, we show that skin roughness increases with body size. Skin patches on the ventral body region of lizards were rougher than on the dorsum, but this was a consequence of ventral scales being larger than dorsal scales. Dorsal surface skew and kurtosis varied with body size, but surfaces on the ventral skin showed no such relationship. Scale size scaled isometrically with body size, and while ventral scales differed in shape from dorsal scales, scale shape did not change with ontogeny. Overall, this study demonstrates that gel-based stereo-profilometry is a promising method to rapidly assess the 3D surface structure of reptilian skin at the microscopic level. Additionally, our findings of the explanatory power of body size on skin surface diversity provide a foundation for future studies to disentangle the relationships among morphological, functional and ecological diversity in squamate reptile skin surfaces., (© 2019 Anatomical Society.)

Salmo salar post-smolts were reared in seawater under controlled laboratory conditions for 12 weeks. The fish were exposed to three constant temperature treatments (15, 10.5 and 6°C) and four feeding treatments (constant feeding, food withheld for 7 days, food withheld for 14 days and food withheld intermittently for four periods of 7 days). Scale growth was proportional to fish growth across all treatments, justifying the use of scale measurements as a proxy for growth during the early marine phase. The rate of circuli deposition was dependant on temperature and feeding regime and was generally proportional to fish growth but with some decoupling of the relationship at 15°C. Deposition rates varied from 4.8 days per circulus at 15°C (constant feeding) to 15.1 days per circulus at 6°C (interrupted feeding). Cumulative degree day ( ° D) was a better predictor of circuli number than age, although the rate of circuli deposition ° D -1 was significantly lower at 6°C compared with 15 and 10.5°C. Inter-circuli distances were highly variable and did not reflect growth rate; tightly packed circuli occurred during periods without food when growth was depressed, but also during periods of rapid growth at 15°C. The results further current understanding of scale growth properties and can inform investigations of declining marine growth in S. salar based on interpretations of scale growth patterns., (© 2019 The Fisheries Society of the British Isles.)

Recent studies on the analysis of the evolutionary relationships between the various taxa of the teleost fishes are based on the evaluation of the efficacy of different approaches, verifying the taxonomical congruence between the hypotheses inferred from molecular data and those from morphological data. The first objective of this study was to investigate the macro- and microscopic characterization of the body key scales for the sixteen Aphanius species, and to discuss their taxonomic significance. In addition, to reveal the phylogenetic origins of particular adaptations and to identify a possible cases of convergent evolution in scales of Aphanius species, we used a phylogenetic character mapping procedure that uses an independent data source, namely molecular data that is presumed to be functionally independent of the scale traits. Comparison of the resulted phylogenetic trees and scale data dendrogram implied that neither scale shape/scale surface morphology, nor scale surface microstructure can be used for the species identification and inferring phylogenetic relationships in the genus Aphanius. This conclusion confirms previous studies, indicating that the scale shape and ornamentations varies with habitat, age and size of the fish. We also concluded that the polygonal state in the scales of Aphanius species was a primitive shared character state., (Copyright © 2019 Elsevier Ltd. All rights reserved.)

Many cichlid species in the shallow-shore of Lake Tanganyika suffer damage from attacks by the scale-eater Perissodus microlepis . Many prey fish engage in warning behaviors to this predator. It has been hypothesized that, if prey fish have difficulty employing such behavioral tactics, morphological defenses against scale-eating, such as hard scales, will evolve. The shrimp-eating cichlids, Altolamprologus compressiceps ( Ac ) and Neolamprologus fasciatus ( Nf ), exhibit hunting behaviors in which they remain motionless for up to 10 seconds while aiming at prey, when they are vulnerable to scale-eating predators; thus, these fish have likely evolved morphological defenses against "scale-attacks". We tested this hypothesis in Ac and Nf , as well as three other predatory fish, Lamprologus callipterus , Lepidiolamprologus elongatus and Lep. attenuatus , that are not motionless for such a long time. Under natural conditions, Ac and Nf were rarely attacked, while the other three species were attacked frequently. When freshly killed specimens of these five species were displayed underwater in the presence of P. microlepis , Ac was rarely attacked, while Nf and the three other species were attacked frequently. Among the five fish species, the force required to tear off scales was highest for Ac , and this force was negatively correlated with the frequency of attacks on the displayed fish. These results support the hypothesis that the hard scales of Ac function as an anti-scale-attack measure, although it remains unclear why free-swimming Nf were rarely attacked while aiming at prey, despite the fact that the force required to tear off its scales was not large., (© 2019 Zoological Society of Japan.)

We describe and interpret the functional morphology of skin of the Yellow-bellied sea snake, Hydrophis platurus. This is the only pelagic sea snake, and its integument differs from what is known for other species of snakes. In gross appearance, the scales of H. platurus consist of non-overlapping, polygonal knobs with flattened outer surfaces bearing presumptive filamentous sensillae. The deep recesses between scales ('hinge') entrap and wick water over the body surface, with mean retention of 5.1 g/cm of skin surface, similar to that determined previously for the roughened, spiny skin of marine file snakes, Acrochordus granulatus. This feature possibly serves to maintain the skin wet when the dorsal body protrudes above water while floating on calm oceanic slicks where they forage. In contrast with other snakes, including three species of amphibious, semi-marine sea kraits (Laticauda spp.), the outer corneous β-protein layer consists of a syncytium that is thinner than seen in most other species. The subjacent α-layer is also thin, and lipid droplets and lamellar bodies are seen among the immature, cornifying α-cells. A characteristic mesos layer, comprising the water permeability barrier, is either absent or very thin. These features are possibly related to (1) permeability requirements for cutaneous gas exchange, (2) reduced gradient for water efflux compared with terrestrial environments, (3) less need for physical protection in water compared with terrestrial ground environments, and (4) increased frequency of ecdysis thought to be an anti-fouling mechanism. The lipogenic features of the α-layer possibly compensate for the reduced or absent mesos layer, or produce layers of cells that comprise what functionally might be termed a mesos layer, but where the organization of barrier lipids nonetheless appears less robust than what is characteristically seen in squamates., (© 2019 Wiley Periodicals, Inc.)

The insects of subfamily Triatominae exhibit many morphological features used for taxonomic identification. In some species, however, these features are very similar. The authors have proposed by first time the description of the structure referred as urotergite I process. The objective of the study was analyse the use of urotergite I process in the species of the genus Triatoma as a taxonomic feature that may contribute to the more precise and correct identification of these Trypanosoma cruzi Chagas, 1909 vectors. Scanning electron microscopy (SEM) was used to describe the structure of urotergite I process in ten Triatoma (Hemiptera, Reduviidae, Triatominae) species: Triatoma brasiliensis Neiva, 1911; Triatoma circummaculata Stål, 1859; Triatoma infestans (Klug, 1834); Triatoma maculata (Erichson, 1848); Triatoma pseudomaculata Corrêa & Espínola, 1964; Triatoma rubrovaria (Blanchard, 1843); Triatoma sordida (Stål, 1859); Triatoma tibiamaculata (Pinto, 1926); Triatoma vitticeps (Stål, 1859) and Triatoma wygodzinskyi Lent, 1951. The morphological description of the urotergite I process reflects the taxonomic value of the structure for separating and identifying species of ten specimens of the genus Triatoma. The morphological pattern of the urotergite I process on these ten species was compared, and interspecific variability was observed. We suggest the use of the urotergite I process as a complementary character to identify insects of the subfamily Triatominae. In the future, the use of the urotergite I process in dichotomic keys to identify triatomines may contribute to the improvement of the entomological surveillance of Chagas' disease., (Copyright © 2019 Elsevier B.V. All rights reserved.)

Scale morphology and squamation play an important role in the study of fish phylogeny and classification. However, as the scales of the earliest osteichthyans or bony fishes are usually found in a disarticulated state, research into squamation patterns and phylogeny has been limited. Here we quantitatively describe the scale morphology of the oldest articulated osteichthyan, the 425-million-year-old Guiyu oneiros, based on geometric morphometrics and high-resolution computed tomography. Based on the cluster analysis of the scales in the articulated specimens, we present a squamation pattern of Guiyu oneiros, which divides the body scales into 4 main belts, comprising 16 areas. The new pattern reveals that the squamation of early osteichthyans is more complicated than previously known, and demonstrates that the taxa near the crown osteichthyan node in late Silurian had a greater degree of squamation zonation compared to more advanced forms. This study offers an important reference for the classification of detached scales of early osteichthyans, provides new insights into the early evolution of osteichthyan scales, and adds to our understanding of the early osteichthyan body plan.

Natural dermal armors are serving as a source of inspiration in the pursuit of "next-generation" structural materials. Although the dynamic strain response of these materials is arguably the most relevant to their performance as armors, limited work has been performed in this area. Here, uniaxial tension and transverse puncture tests were performed on specimens obtained from the scales of Asian carp over strain rates spanning seven decades, from 10 -4 to 10 3 s -1 . The importance of anatomical variations was explored by comparing the performance of scales from the head, middle and tail regions. In both loading orientations, the scales exhibited a significant increase in the resistance to failure with loading rate. The rate sensitivity was substantially higher for transverse loading than for in-plane tension, with average strain rate sensitivity exponents for measures of the toughness of 0.35 and 0.08, respectively. Spatial variations in the properties were largest in the puncture responses, and scales from the head region exhibited the greatest resistance to puncture overall. The results suggest that the layered microstructure of fish scales is most effective at resisting puncture, rather than in-plane tension, and its effectiveness increases with rate of loading. X-ray microCT showed that delamination of plies in the internal elasmodine and stretching of the fibrils were key mechanisms of energy dissipation in response to puncture loading. Understanding contributions from the microstructure to this behavior could guide the development of flexible engineered laminates for penetration resistance and other related applications., (Copyright © 2018 Elsevier Ltd. All rights reserved.)

Fish scales are laminated composites that consist of plies of unidirectional collagen fibrils with twisted-plywood stacking arrangement. Owing to their composition, the toughness of scales is dependent on the intermolecular bonding within and between the collagen fibrils. Adjusting the extent of this bonding with an appropriate stimulus has implications for the design of next-generation bioinspired flexible armours. In this investigation, scales were exposed to environments of water or a polar solvent (i.e. ethanol) to influence the extent of intermolecular bonding, and their mechanical behaviour was evaluated in uniaxial tension and transverse puncture. Results showed that the resistance to failure of the scales increased with loading rate in both tension and puncture and that the polar solvent treatment increased both the strength and toughness through interpeptide bonding; the largest increase occurred in the puncture resistance of scales from the tail region (a factor of nearly 7×). The increase in strength and damage tolerance with stronger intermolecular bonding is uncommon for structural materials and is a unique characteristic of the low mineral content. Scales from regions of the body with higher mineral content underwent less strengthening, which is most likely the result of interference posed by the mineral crystals to intermolecular bonding. Overall, the results showed that flexible bioinspired composite materials for puncture resistance should enrol constituents and complementary processing that capitalize on interfibril bonds.

The crustacean cuticle forms skeletal elements consisting of chitin-protein fibrils reinforced by amorphous and crystalline calcium carbonate and phosphate minerals. The edges of skeletal elements are of particular interest. They are subject to repeated strain and stress because they form transitions to the arthrodial membranes connecting them. These allow for relative movements of skeletal elements. In this study, we investigate structure, chemical composition, mineral organization and local mechanical properties of the anterior and posterior edges of the tergite cuticle in the conglobating beach isopod Tylos europaeus and compare these with the protective dorsal region of the tergites. The distribution of mineral phases at the edges resembles that of dorsal regions of the tergites. At the transition with the unmineralized arthrodial membrane the calcite containing distal exocuticle is replaced by epicuticular material and the subjacent cuticular layers containing amorphous calcium carbonate become enriched with amorphous calcium phosphate. At the edges, the local elastic modulus and hardness values are significantly lower compared to dorsal regions of the tergite cuticle, for both, the calcite and the amorphous mineral containing layers. The calcite within the tergite cuticle is assembled in different texture patterns: (i) almost random co-orientation, (ii) almost single crystalline calcite, and (iii) a graded organization. Calcite organization and co-orientation strength is highly variable, not only on very few tens of micrometres, but also between regions with different skeletal functionality. Our results show that besides structure and composition, patterns of calcite organization contribute to the hierarchical architecture and functionality of biological composites., (Copyright © 2018 Elsevier Inc. All rights reserved.)

A new freshwater goby, Rhinogobius immaculatus sp. nov., is described here from the Qiantang River in China. It is distinguished from all congeners by the following combination of characters: second dorsal-fin rays I, 7-9; anal-fin rays I, 6-8; pectoral-fin rays 14-15; longitudinal scales 29-31; transverse scales 7-9; predorsal scales 2-5; vertebrae 27 (rarely 28); preopercular canal absent or with two pores; a red oblique stripe below eye in males; branchiostegal membrane mostly reddish-orange, with 3-6 irregular discrete or connected red blotches on posterior branchiostegal membrane and lower operculum in males; caudal-fin base with a median black spot; and no black blotch on anterior part of first dorsal fin in males.

Background: The shedding of epithelial squames (skin scales) by staff in operating theatre air is an important source of deep infection following joint replacement surgery. This is a serious complication, resulting in significant morbidity for the patient and substantial cost implications for healthcare systems. Much effort has been put into providing clean air in operating theatres, yet little attention has been given to reducing the shedding of surface skin scales at source., Aim: To develop a novel method for calculating surface skin scale density using surface microscopy, and to use it to evaluate the effect of a skincare regimen on operating theatre staff., Methods: Surface microscopy with Z-stacked imaging was used to visualize the effect of a skincare regimen involving three stages: washing with soap; exfoliation; and application of emollient. A USB microscope was then used in a field study to take images of the skin of operating theatre staff who applied the regimen to one lower limb the night before testing. The other limb was used as a control. Two blinded assessors analysed scale density., Results: Z-stack images from the surface microscope enabled observations of the skincare regimen. The USB microscope also provided adequate images that enabled assessment of skin scale density. In the operating theatre staff, a 72.1% reduction in visible skin scales was observed following application of the skincare regimen., Conclusions: Further work is required to demonstrate how this effect correlates with dispersion of skin particles in a cleanroom, and subsequently in live operating theatre studies., (Copyright © 2018 The Healthcare Infection Society. Published by Elsevier Ltd. All rights reserved.)

Schizothoracine is the predominant wild fish subfamily of the Tibetan plateau (TP). Their scales, pharyngeal teeth and barbels have gradually regressed with increasing altitude. Schizothoracine have been divided into three groups: primitive, specialized and highly specialized. Ectodysplasin-A ( Eda ) has been considered as a major gene that contributes to the development of skin appendages. The present study cloned the Eda genes of 51 Schizothoracine fish species which represent the three groups and five Barbinae species. Phylogenetic analyses indicated that Eda may have acted as the genetic trigger for scale loss in the Schizothoracine. Furthermore, 14 single nucleotide polymorphisms (SNPs) and two deletions (18 bp and 6 bp in size), were also detected in the Eda coding sequence of the highly specialized group compared to the primitive group. The same SNPs and two indels result in four non-synonymous and two G-X-Y and 1 XY motif indels, which possibly contribute to significant structure changes in the Eda gene. The domain including (G-X-Y) n motif in the Eda gene is relatively conserved amongst teleosts. Based on the above results, we hypothesize that the evolution of Eda gene might be associated with the scale loss in Schizothoracine fishes in response to the phased uplift of the TP.

Ectothermic vertebrates are sensitive to thermal fluctuations in the environments where they occur. To buffer these fluctuations, ectotherms use different strategies, including the integument, which is a barrier that minimizes temperature exchange between the inner body and the surrounding air. In lizards, this barrier is constituted by keratinized scales of variable size, shape and texture, and its main function is protection, water loss avoidance and thermoregulation. The size of scales in lizards has been proposed to vary in relation to climatic gradients; however, it has also been observed that in some groups of Iguanian lizards it could be related to phylogeny. Thus, here, we studied the area and number of scales (dorsal and ventral) of 60 species of Liolaemus lizards distributed in a broad latitudinal and altitudinal gradient to determine the nature of the variation of the scales with climate, and found that the number and size of scales are related to climatic variables, such as temperature and geographical variables as altitude. The evolutionary process that best explained how these morphological variables evolved was the Ornstein-Uhlenbeck model. The number of scales seemed to be related to common ancestry, whereas dorsal and ventral scale areas seemed to vary as a consequence of ecological traits. In fact, the ventral area is less exposed to climate conditions such as ultraviolet radiation or wind and is, thus, under less pressure to change in response to alterations in external conditions. It is possible that scale ornamentation, such as keels and granulosity, may bring some more information in this regard., (© 2018 International Society of Zoological Sciences, Institute of Zoology/Chinese Academy of Sciences and John Wiley & Sons Australia, Ltd.)

Biological surfaces with unique wettability in nature have provided an enormous innovation for scientists and engineers. More specifically, materials possessing various wetting properties have drawn considerable attention owing to their promising application prospects. Recently, great efforts have been concentrated on the researches on wetting-induced drag-reduction materials inspired by biology because of their ability to save energy. In this work, the drag-reduction characteristics of the bionic surface with delicate water-trapping microstructures of fish Ctenopharyngodon idellus scales were explored by experimental method. Firstly, the resistance of smooth surface and bionic surface experimental sample at different speeds was carefully tested through the testing system for operation resistance. Then, the contact angle (CA) of fish scale surface was measured by means of the contact angle measuring instrument. It was discovered that the bionic surface created a rewarding drag-reduction effect at a low speed, and the drag-reduction rate significantly displayed a downward trend with the increase in flow speed. Thus, when the rate was 0.66 m/s, the drag-reduction effect was at the optimum level, and the maximum drag reduction rate was 2.805%, which was in concordance with the simulated one. Furthermore, a contact angle (CA) of 11.5° appeared on the fish scale surface, exhibiting fine hydrophilic property. It further manifested the spreading-wetting phenomenon and the higher surface energy for the area of apical of fish scales, which played an important role in drag-reduction performance. This work will have a great potential in the engineering and transportation field.

Four species of the genus Osteobrama collected from five different sampling sites of Chindwin and Barak river basin of North-East India, namely O. belangeri, O. cotio, O. cunma and O. feae, were characterized. Meristic study showed differences in anal fin rays count and lateral line scales. Morphometric analysis revealed significant differences among Osteobrama species. In Truss analysis, a clear pattern of differentiation was observed among the four species with discriminant function analysis assigning 100% correctly to the particular species. In molecular analysis, four Osteobrama species collected were barcoded with COI and 16S rRNA sequences and phylogenetically these four species formed two distinct clusters, O. belangeri form one separate cluster, from the other three species i.e. O. cotio, O. cunma and O. feae. High-resolution clusters generated (NJ trees) aided the groupings of species corresponding to their genera and families which are in confirmation to the values generated by COI Automatic Barcode Gap Discovery bioinformatics platform.

Size-selective mortality owing to lack of energy reserves during the first marine winter has been suggested to be a result of juvenile salmon failing to reach a critical size or condition by the end of their first marine summer and not surviving the following winter due to this presumed energy deficit. This hypothesis implies strong size dependency of mortality, and is subject to empirical data support for acceptance. Scale circulus spacing has been interpreted as an index for body size, and we reviewed the effect of size-selective mortality with a knife-edge mortality function on descriptive statistics for a scale circulus spacing index (SCSI). In order to invoke size selection as an important driver of mortality during the first year of ocean rearing, it is necessary to demonstrate not only that size-selective mortality is directed towards the smaller members of the population, but that the selective nature of the mortality can account for a substantial portion of the observed mortality. If the assumption is made that a random sample of a single juvenile population has been obtained, then studies that employ a SCSI to infer size-selective mortality coupled with a critical size limit must demonstrate a shift toward larger values of the SCSI, but also a concomitant reduction in the variance and range of the SCSI and an increase in the skewness and kurtosis of the SCSI values. Through simulation we found that the percentage of adults that displayed a SCSI value greater than the maximum observed in the juvenile sample was highly dependent on the initial juvenile sample size and size-selective mortality rate. Geographical distributions of juvenile Pacific salmon can be stratified by size, with larger individuals migrating earlier from local ocean entry locations than smaller individuals, and thus differential timing migration of juveniles based upon body size prior to the collection of the marine juvenile sample may be a more plausible explanation of published trends in the SCSI, rather than invoking substantial size-selective mortality and a critical size limit., Competing Interests: The authors have declared that no competing interests exist.

Tunas of the genus Thunnus possess many morphological and physiological adaptations for their high-performance epipelagic ecology. Although Thunnus anatomy has been studied, there are no quantitative studies on the structure of their scales. We investigated the scales of bigeye tuna (Thunnus obesus) from ten regions of the body using micro computed tomography (µCT)-scanning and histology to quantitatively and qualitatively compare regional scale morphology. We found a diversity of scale sizes and shapes across the body of bigeye tuna and discriminant function analysis on variables derived from µCT-data showed that scales across the body differ quantitatively in shape and size. We also report the discovery of a novel scale type in corselet, tail, and cheek regions. These modified scales are ossified shells supported by internal trabeculae, filled with fat, and possessing an internal blood supply. Histological analysis showed that the outer lamellar layers of these thickened scales are composed of cellular bone, unexpected for a perciform fish in which bone is typically acellular. In the fairing region of the anterior body, these fat-filled scales are stacked in layers up to five scales deep, forming a thickened bony casing. Cheek scales also possess a fat-filled internal trabecular structure, while most posterior body scales are more plate-like and similar to typical teleost scales. While the function of these novel fat-filled scales is unknown, we explore several possible hypotheses for their function., (© 2018 Wiley Periodicals, Inc.)

Background: The experimental approach to the evolution and development of the vertebrate skeleton has to a large extent relied on "direct-developing" amniote model organisms, such as the mouse and the chicken. These organisms can however only be partially informative where it concerns secondarily lost features or anatomical novelties not present in their lineages. The widely used anamniotes Xenopus and zebrafish are "indirect-developing" organisms that proceed through an extended time as free-living larvae, before adopting many aspects of their adult morphology, complicating experiments at these stages, and increasing the risk for lethal pleiotropic effects using genetic strategies., Results: Here, we provide a detailed description of the development of the osteology of the African mouthbrooding cichlid Astatotilapia burtoni, primarily focusing on the trunk (spinal column, ribs and epicentrals) and the appendicular skeleton (pectoral, pelvic, dorsal, anal, caudal fins and scales), and to a lesser extent on the cranium. We show that this species has an extremely "direct" mode of development, attains an adult body plan within 2 weeks after fertilization while living off its yolk supply only, and does not pass through a prolonged larval period., Conclusions: As husbandry of this species is easy, generation time is short, and the species is amenable to genetic targeting strategies through microinjection, we suggest that the use of this direct-developing cichlid will provide a valuable model system for the study of the vertebrate body plan, particularly where it concerns the evolution and development of fish or teleost specific traits. Based on our results we comment on the development of the homocercal caudal fin, on shared ontogenetic patterns between pectoral and pelvic girdles, and on the evolution of fin spines as novelty in acanthomorph fishes. We discuss the differences between "direct" and "indirect" developing actinopterygians using a comparison between zebrafish and A. burtoni development.

We adopted light and electron microscopy to understand the structure of the skin of two species of caecilians, Ichthyophis tricolor and Uraeotyphlus cf. oxyurus, from Western Ghats of Kerala, India. The surface of the skin of these caecilians contains an irregular pattern of microridges. Oval, round and polymorphic glandular openings are randomly distributed all over the skin surface. Most of the openings are funnel shaped. The epithelial cells along the rim of the opening descend into the tunnel of the duct. A few glandular openings protrude slightly above the epithelium of the duct. The skin is formed of epidermis and dermis. Small flat disk-like dermal scales, composed of a basal plate of several layers of unmineralized collagen fibers topped with a discontinuous layer of mineralized globular squamulae, are lodged in pouches in the transverse ridges of the skin. Each pouch contains 1-4 scales, which might differ in size. The scales are almost similar between species, yet the difference can be useful in distinguishing between the two species. Flask cells and Merkel cells are present in the epidermis. Two types of glands, mucous and granular, are present in the dermis. The mucous glands are densely packed with mucous vesicles. Darkly stained mucous producing cells are located around the periphery of the gland. Secretory mucous vesicles differ in their organization and distribution. The granular glands are located perpendicular to the skin surface. The granule producing cells of the gland are located near the periphery. There are differently stained spherical secretory granules of various sizes in the cytoplasm. Thus, the use of different microscopic techniques contributed fascinatingly to the first ever understanding of organization of the skin of two selected caecilian species from Western Ghats, revealing certain features to differ between them., (Copyright © 2018 Elsevier Ltd. All rights reserved.)

Body scales in teleost fish are the first line of defense in protecting the fish from invading pathogens. However, the relationship between the adaptive evolution of immune-related genes and changes in the body scale-covering of fish has not been previously studied. Schizothoracine fish, characterized by progressive reduction of body scales from the primitive to the highly specialized species, are a good group to investigate this relationship. We obtained 11 IRF genes (IRF1-11) from 14 of schizothoracine fish representing primitive, specialized, and highly specialized species, of which seven IRF genes (IRF2, IRF3, IRF5, IRF6, IRF7, IRF8 and IRF9) contained the complete CDS. Sequence analysis demonstrated the deletion or insertion of 4-7 amino acids in IRF2, IRF3, IRF6 and IRF9, which seems to be a common phenomenon in the schizothoracine fish. Selection pressure analysis supported the hypothesis that positive selection has been driving the rapid evolution of IRFs in specific lineages of the schizothoracine fish. This, however, is not correlated with body scale reduction or loss in the evolution of these IRFs. Remarkably, the deletion or insertion found in IRF protein sequences presented a regular pattern corresponding to the scale-covering changes in schizothoracine fish. Our study provides evidence for positive selection in the IRF family, contributing to a better understanding of the adaptive evolution of immune-related genes in schizothoracine fish in response to environmental changes of the Qinghai-Tibetan Plateau., (Copyright © 2017 Elsevier Ltd. All rights reserved.)

Fish scales serve as a flexible natural armor that have received increasing attention across the materials community. Most efforts in this area have focused on the composite structure of the predominately organic elasmodine, and limited work addresses the highly mineralized external portion known as the Limiting Layer (LL). This coating serves as the first barrier to external threats and plays an important role in resisting puncture. In this investigation the structure, composition and mechanical behavior of the LL were explored for three different fish, including the arapaima (Arapaima gigas), the tarpon (Megalops atlanticus) and the carp (Cyprinus carpio). The scales of these three fish have received the most attention within the materials community. Features of the LL were evaluated with respect to anatomical position to distinguish site-specific functional differences. Results show that there are significant differences in the surface morphology of the LL from posterior and anterior regions in the scales, and between the three fish species. The calcium to phosphorus ratio and the mineral to collagen ratios of the LL are not equivalent among the three fish. Results from nanoindentation showed that the LL of tarpon scales is the hardest, followed by the carp and the arapaima and the differences in hardness are related to the apatite structure, possibly induced by the growth rate and environment of each fish., Statement of Significance: The natural armor of fish, turtles and other animals, has become a topic of substantial scientific interest. The majority of investigations have focused on the more highly organic layer known as the elasmodine. The present study addresses the highly mineralized external portion known as the Limiting Layer (LL). Specifically, the structure, composition and mechanical behavior of the LL were explored for three different fish, including the arapaima (Arapaima gigas), the tarpon (Megalops atlanticus) and the carp (Cyprinus carpio). Results show that there are significant differences in the surface morphology of the LL from posterior and anterior regions in the scales, and between the three species. In addition, the composition of the LL is also unique among the three fish. Results from nanoindentation showed that the LL of tarpon scales is the hardest, followed by the carp and the arapaima and the differences in hardness are related to the apatite structure, possibly induced by the growth rate and environment of each fish. In addition, a new feature was indentified in the LL, which has not been discussed before. As such, we feel this work is unique and makes a significant contribution to the field., (Copyright © 2017 Acta Materialia Inc. Published by Elsevier Ltd. All rights reserved.)

A re-evaluation of the armoured catfish species of Hypostomus in the Rio Jaguaribe, north-eastern Brazil, was prompted by the discovery of specimens with pale spots on a dark background collected from that basin c. 1936 and deposited at the Academy of Natural Sciences of Philadelphia. Recent field collections in the Rio Jarguaribe basin confirmed the presence of the pale-spotted specimens and its distinctiveness as a new species. Hypostomus sertanejo n. sp. is diagnosed from other species of Hypostomus by having fins and dermal-plated regions of head and body with pale spots or vermiculations on darker background, teeth slender, asymmetrically bicuspid and numerous (34-75) on dentary and premaxilla, depressed dorsal-fin spine not reaching adipose spine, unbranched pectoral-fin spine longer than unbranched pelvic-fin ray, seven branched dorsal-fin rays and one (rarely two) predorsal plate(s) bordering supraoccipital. Ancistrus salgadae Fowler 1941 is hypothesized to be a junior synonym of Hypostomus carvalhoi (Miranda-Ribeiro, 1937), a dark-spotted Hypostomus described from the Rio Granjeiro, a tributary to the upper Rio Salgado., (© 2017 The Fisheries Society of the British Isles.)