The structure and mechanical design of rhinoceros dermal armour.

The collagenous dermis of the white rhinoceros forms a thick, protective armour that is highly specialized in its structure and material properties compared with other mammalian skin. Rhinoceros skin is three times thicker than predicted allometrically, and it contains a dense and highly ordered three-dimensional array of relatively straight and highly crosslinked collagen fibres. The skin of the back and flanks exhibits a steep stress-strain curve with very little 'toe' region, a high elastic modulus (240 MPa), a high tensile strength (30 MPa), a low breaking strain (0.24) and high breaking energy (3 MJm-3) and work of fracture (78 kJm-2). By comparison, the belly skin is somewhat less stiff, weaker, and more extensible. In compression, rhinoceros skin withstands average stresses and strains of 170 MPa and 0.7, respectively, before yielding. As a biological material, rhinoceros dorsolateral skin has properties that are intermediate between those of 'normal' mammalian skin and tendons. This study shows that the dermal armour of the rhinoceros is very well adapted to resist blows from the horns of conspecifics, as might occur during aggressive behaviour, due to specialized material properties as well as its great thickness.     

The presence of a spiracle in the Heterostraci (Agnatha)

Study of new fossil material suggests that spiracles were present in very early Agnatha. This leads to the supposition that jaws evolved earlier than was formerly supposed.     

Affinities of the Heterostraci (Agnatha)

From a study of the perforations in the heterostracan carapace and the impressions on the inner surface of the bony plates it is suggested that the Heterostraci in most features belonged to a structural stage from which it was possible to derive both the gnathostomes and the cyclostomes (including the cephalaspids and anaspids).
However, the structure of the gills and the presence of a spiracle in the Heterostraci indicate that they were more closely related to the gnathostome stock.     

Role of the dermal tracts in the pigment pattern of the frog.

The pigment pattern of the ventral skin of the frog Rana esculenta is compared in skin fragments grown for 24 hr with or without antiserum directed to fibronectin (anti-FN). Melanocyte-stimulating hormone (MSH) was added to the medium during the last hour in culture in order to enhance visibility of melanophores in the ventral region of the frog skin. Comparison of these two treatments provides information regarding the precise localization of melanophores in the dermal tracts and their involvement in the pigment pattern of the ventral frog skin. In this regard, the whitish pigment pattern of skin fragments is compared to the tiny black spots found on anti-FN treated skin fragments and the abundant blotchy spots found on skin cultured alone. The distribution of melanophores in the dermal tracts observed in vertical semithin sections is found to be related to the three different levels of the dermal tracts. This report demonstrates the importance of fibronectin as a substrate for the melanophore migration, the importance of the tract level for the melanophore localization both involved in the pigment pattern of the ventral skin.     

Alterations in the secretory pattern of dermal dendritic cells following melanin uptake

Under a variety of circumstances, melanin occurs in the dermal compartment of the skin, being mostly observed in cells that have been termed melanophages, some of which have been identified as dermal dendritic cells. We analysed changes in the expression and secretion pattern of cytokines by dendritic cells after the uptake of melanin from various sources. Dendritic cells were derived from human primary blood monocytes or from the human monocytic cell line THP-1. Melanin uptake increased the secretion of the chemokines MIP-1β (CCL4) and MCP-1 (CCL2). The higher MIP-1β secretion was accompanied by higher MIP-1β gene expression. Elevation of MIP-1β secretion was dependent on the uptake of melanin but could not be induced by the phagocytosis of latex beads, indicating that the phagocytic process itself was not sufficient to increase the secretion of this cytokine. The data thus show that the uptake of melanin changes the cytokine expression and secretion pattern of dendritic-like cells.     

Competition for territories does not explain allopaternal care in the tessellated darter

Extensive research has focused on understanding the evolution of parental care, with fishes providing important model systems for understanding patterns of variation within and between species. Classic theory predicts that individuals will care for offspring when the fitness benefits through increased offspring survival and growth outweigh the cost to the parents through decreased future reproductive opportunities. Yet, a puzzling observation not explained by this basic theory is the fact that in some species individuals defend and provision unrelated offspring and thus exhibit alloparental care. The tessellated darter, Etheostoma olmstedi, represents one of the first known examples of allopaternal care in fishes. In this species, males often clean and guard eggs fertilized but deserted by other males. Allopaternal care has been argued to occur in the tessellated darter because of competition for a limited number of mating sites where less dominant males accept territories with eggs when other breeding sites are not available. Here, we test this hypothesis using male territory choice experiments. When allowed to choose between two otherwise identical territories either containing eggs fertilized by another male or with no eggs, males spent significantly more time at territories with eggs. This demonstrates that competition for mating territories is not the primary factor explaining the existence of allopaternal care in the tessellated darter. Instead, males of this species may exhibit allopaternal care to dilute predator pressure on their own eggs or because females prefer to mate with males whose territories contain eggs.     

Origins of bone repair in the armour of fossil fish: response to a deep wound by cells depositing dentine instead of dermal bone

The outer armour of fossil jawless fishes (Heterostraci) is, predominantly, a bone with a superficial ornament of dentine tubercles surrounded by pores leading to flask-shaped crypts (ampullae). However, despite the extensive bone present in these early dermal skeletons, damage was repaired almost exclusively with dentine. Consolidation of bone, by dentine invading and filling the vascular spaces, was previously recognized in Psammolepis and other heterostracans but was associated with ageing and dermal shield wear (reparative). Here, we describe wound repair by deposition of dentine directly onto a bony scaffold of fragmented bone. An extensive wound response occurred from massive deposition of dentine (reactionary), traced from tubercle pulp cavities and surrounding ampullae. These structures may provide the cells to make reparative and reactionary dentine, as in mammalian teeth today in response to stimuli (functional wear or damage). We suggest in Psammolepis, repair involved mobilization of these cells in response to a local stimulatory mechanism, for example, predator damage. By comparison, almost no new bone is detected in repair of the Psammolepis shield. Dentine infilling bone vascular tissue spaces of both abraded dentine and wounded bone suggests that recruitment of this process has been evolutionarily conserved over 380 Myr and precedes osteogenic skeletal repair.     

Role of drainage and barriers in the genetic structuring of a tessellated darter population

While population genetic structuring is easily identified, the causes of the structure can be difficult to determine. Habitat fragmentation in aquatic systems has often been identified as a major source of increased population structure and decreased genetic diversity in fish, including benthic resident species such as darters. However, these findings are often not replicated across natural and manmade barriers and come from endangered or threatened populations where the genetic structure is likely already compromised due to small population size. To evaluate the factors involved in structuring a healthy darter population, we genotyped 506 tessellated darters from 18 sites in three different river drainages and one large lake. Sites were all in the same watershed but separated from one another by one or more of three different types of barriers: dams, natural fall lines and causeways. We found that while diversity and allele frequency varied largely by drainage, within drainage variation was minimal even across multiple barriers. No single barrier type appeared to be more formidable than any other. Our results indicate that healthy populations of darters may naturally be structured by drainage, but likely disperse across barriers enough to retain drainage-wide homogeneity.     

Phylogeny of the Pteraspidiformes (Heterostraci), Silurian–Devonian jawless vertebrates

The Pteraspidiformes are an order of armoured agnathans of Silurian and Devonian age composed of five families: the Anchipteraspididae, Protopteraspididae, Pteraspididae, Protaspididae and Psammosteidae. Relationships within the order are poorly known and although past attempts have been made to represent the state of knowledge in this area none have attempted a rigorous computer-based analysis. Here we present the results of the cladistic analysis on a matrix comprising 25(+1) taxa and 59 characters using W inclada . The first analysis was run without any representative of the Psammosteidae and supports the presence of five groups within the order although one, the 'Protopteraspididae', is paraphyletic and a second, the Gigantaspididae, is newly recognized. In a second analysis the addition of the psammosteid Drepanaspis gemuendenensis results in a better resolved tree in which the basal family, the Anchipteraspididae, are clearly monophyletic and in which the Psammosteidae, as represented by Drepanaspis , is the next less inclusive. This is contrary to the previous view that the psammosteids were an advanced clade within the Pteraspidiformes.     

Allopaternal care in the tessellated darter,Etheostoma olmstedi (Pisces: Percidae)

Synopsis Large male tessellated darters, Etheostoma olmstedi, defend flat-bottomed rocks, the undersides of which serve as spawning substrates. Because females attach eggs directly onto bare stone, a spawning bout necessarily decreases the surface area available for further breeding at a nest. In apparent response to the female preference of ovipositing on rocks with the most bare surface, fathers abandon nests with their attached offspring to search for rocks with more uncovered surface. Egg abandonment also results after roving fathers intercept and spawn with ripe females and remain at a different rock to care for a new clutch. In essence, large males appear to maximize the number of eggs they fertilize by sequentially monopolizing breeding patches that are temporarily most desirable to females. The costs to a father of abandoning his eggs appear to be low for two reasons: (1) non-territorial males, created by a scarcity of suitable rocks, readily occupy vacant nests, and clean and aerate the abandoned eggs; and (2) after water hardening, eggs are less vulnerable to cannibalism. The maintenance of eggs by nonparental males may be both incidental to scrubbing the ceiling for their own breeding and adaptive in that viable eggs stimulate females to spawn; neither would appear to involve parental investment costs. Thus, allopaternal care in the tessellated darter may have evolved because it is performed in the selfish pursuit of spawning opportunities while entailing little, if any, of the costs normally subsumed within parental investment.     

Genetics of body shape and armour variation in threespine sticklebacks

Patterns of genetic variation and covariation can influence the rate and direction of phenotypic evolution. We explored the possibility that the parallel morphological evolution seen in threespine stickleback (Gasterosteus aculeatus) populations colonizing freshwater environments is facilitated by patterns of genetic variation and covariation in the ancestral (marine) population. We estimated the genetic (G) and phenotypic (P) covariance matrices and directions of maximum additive genetic (g(max) ) and phenotypic (p(max) ) covariances of body shape and armour traits. Our results suggest a role for the ancestral G in explaining parallel morphological evolution in freshwater populations. We also found evidence of genetic constraints owing to the lack of variance in the ancestral G. Furthermore, strong genetic covariances and correlations among traits revealed that selective factors responsible for threespine stickleback body shape and armour divergence may be difficult to disentangle. The directions of g(max) and p(max) were correlated, but the correlations were not high enough to imply that phenotypic patterns of trait variation and covariation within populations are very informative of underlying genetic patterns.     

The dermal chromatophore unit

Rapid color changes of amphibians are mediated by three types of dermal chromatophores, xanthophores, iridophores, and melanophores, which comprise a morphologically and physiologically distinct structure, the dermal chromatophore unit. Xanthophores, the outermost element, are located immediately below the basal lamella. Iridophores, containing light-reflecting organelles, are found just beneath the xanthophores. Under each iridophore is found a melanophore from which processes extend upward around the iridophore. Finger-like structures project from these processes and occupy fixed spaces between the xanthophores and iridophores. When a frog darkens, melanosomes move upward from the body of the melanophore to fill the fingers which then obscure the overlying iridophore. Rapid blanching is accomplished by the evacuation of melanosomes from these fingers. Pale coloration ranging from tan to green is provided by the overlying xanthophores and iridophores. Details of chromatophore structure are presented, and the nature of the intimate contact between the chromatophore types is discussed.     

The identification of the dermal bones of the head

The discovery of the ichthyostegid Amphibia in Upper Devonian rocks by Säve-Söderbergh (1932) introduced further difficulties into the already complex problems of the dermal bones of the skull roof. For some years previously ideas about the origin of the tetrapods had been dominated by Watson's (1926) Croonian Lecture in which he had demonstrated beyond reasonable doubt that the crossopterygian fishes and not the Dipnoi were their ancestors, and had attempted to show that many of the features of the Carboniferous labyrinthodonts were a direct inheritance from these fishes. It was to be expected, therefore, that any Amphibia from the Upper Devonian would be intermediate in their structures between the Middle Devonian osteolepids and the Carboniferous labyrinthodonts, but when discovered the ichthyostegids did not conform at all well to this expectation. While their skulls showed some very primitive features which might have been expected, the pattern of the dermal bones did not conform to plan, for these new animals had lost, it seemed, the intertemporals, bones found in both the osteolepids and nearly all early labyrinthodonts, and had a single postparietal bone in place of the paired bones of all other early Amphibia. The osteolepid skull had many more bones than these earliest Amphibia.     

Protopteraspis Gosseleti (Vertebrata: Pteraspidomorphi: Heterostraci) from the Lower Devonian of Shropshire, England

New Protopteraspis material from Shropshire is sufficiently similar to Protopteraspis leathensis from the Welsh Borderland, and the type-species Protopteraspis gosseleti from France, to justify transfering P. leathensis to P. gosseleti . The Anglo-Welsh, biostratigraphic leathensis Zone is changed to the Protopteraspis Zone ( sensu Blieck and Janvier 1989). Possible lifestyles are discussed for Protopteraspis gosseleti .