The tessellated pattern of dermal armour in the Heterostraci

The earliest and most primitive heterostraeans possessed a tessellated carapace. Isolated dentine tubercles scattered in the skin formed primordia around which concentric rings of further tubercles were laid down. This pattern of growth produced a characteristic terrazzo. From this stage the gradual elimination of tesserae can be traced in several groups. Beginning with areally growing or cyclomorial tesserae, the individual units appear simultaneously or synchronomorially, thereafter they become fused into a system of large discrete plates. Finally these synehroriomorial units appear earlier and earlier in ontogeny with progressively wider zones of cyclomorial growth being added on to them. Thus a pattern of cyclomorial plates is eventually produced.
In the psammosteids there was a redevelopment of tesserae so that the latest stages were comparable to the very early stages, although the tesserae were developed on an already existing pattern of large plates.
The possible origin of the tessellated pat tern of dermal armour is discussed. The apparent macular nature of dermal structures is not considered to be an inherent property of skin but instead due to simple physico-chemical factors.     

Histologic studies of ostracoderms, placoderms and fossil elasmobranchs. 6. Hard tissues of Ordovician vertebrates

Size and distribution of dermal elements in Ordovician eriptychiids and astraspids are considered relative to phases of skeletal assimilation and regression. The phyletic significance of acellularity in aspidin is discussed, as also is the alleged relationship claimed to exist in initial developmental stages between that hard tissue and dentine proper. To judge from hard tissue histology the astraspids may not, like the eriptychiids, have belonged to the heterostracans but to another group, of early agnathans, still incompletely known.     

Histology and affinity of the earliest armoured vertebrate

Arandaspids are the earliest skeletonizing vertebrates known from articulated remains. Despite a wealth of data, their affinity remains questionable because they exhibit a random mixture of primitive and derived characteristics. We constrain the affinity of arandaspids by providing the first detailed characterization of their dermoskeleton which is revealed to be three-layered, composed of a basal laminated, cancellous middle and tubercular superficial layers. All three layers are composed of acellular bone but the superficial layer also includes dentine and enameloid, comprising the tubercles. As such, the composition of the arandaspid dermoskeleton is common to heterostracans and astraspids, supporting existing hypotheses of early vertebrate phylogeny. This emphasizes the peculiarity of existing interpretations of aranadaspid anatomy and there is need for a complete reappraisal of the existing anatomical data.     

DEVELOPMENT AND EVOLUTIONARY ORIGINS OF VERTEBRATE SKELETOGENIC AND ODONTOGENIC TISSUES

This review deals with the following seven aspects of vertebrate skeletogenic and odontogenic tissues.

• 1 The evolutionary sequence in which the tissues appeared amongst the lower craniate taxa.
• 2 The topographic association between skeletal (cartilage, bone) and dental (dentine, cement, enamel) tissues in the oldest vertebrates of each major taxon.
• 3 The separate developmental origin of the exo- and endoskeletons.
• 4 The neural-crest origin of cranial skeletogenic and odontogenic tissues in extant vertebrates.
• 5 The neural-crest origin of trunk dermal skeletogenic and odontogenic tissues in extant vertebrates.
• 6 The developmental processes that control differentiation of skeletogenic and odontogenic tissues in extant vertebrates.
• 7 Maintenance of developmental interactions regulating skeletogenic/odontogenic differentiation across vertebrate taxa. We derive twelve postulates, eight relating to the earliest vertebrate skeletogenic and odontogenic tissues and four relating to the development of these tissues in extant vertebrates and extrapolate the developmental data back to the evolutionary origin of vertebrate skeletogenic and odontogenic tissues. The conclusions that we draw from this analysis are as follows.
• 8 The dermal exoskeleton of thelodonts, heterostracans and osteostracans consisted of dentine, attachment tissue (cement or bone), and bone.
• 9 Cartilage (unmineralized) can be inferred to have been present in heterostracans and osteostracans, and globular mineralized cartilage was present in Eriptychius, an early Middle Ordovician vertebrate unassigned to any established group, but assumed to be a stem agnathan.
• 10 Enamel and possibly also enameloid was present in some early agnathans of uncertain affinities. The majority of dentine tubercles were bare.
• 11 The contemporaneous appearance of cellular and acellular bone in heterostracans and osteostracans during the Ordovician provides no clue as to whether one is more primitive than the other.
• 12 We interpret aspidin as being developmentally related to the odontogenic attachment tissues, either closer to dentine or a form of cement, rather than as derived from bone.
• 13 Dentine is present in the stratigraphically oldest (Cambrian) assumed vertebrate fossils, at present some only included as Problematica, and is cladistically primitive, relative to bone.
• 14 The first vertebrate exoskeletal skeletogenic ability was expressed as denticles of dentine.
• 15 Dentine, the bone of attachment associated with dentine, the basal bone to which dermal denticles are fused and cartilage of the Ordovician agnathan dermal exoskeleton were all derived from the neural crest and not from mesoderm. Therefore the earliest vertebrate skeletogenic/odontogenic tissues were of neural-crest origin.
• 16 Given the separate developmental and evolutionary origin of the cranial exo- and endoskeletons (both derivatives of the cranial neural crest) we conclude that bone (of attachment) was the primary skeletogenic tissue in the exoskeleton (cartilage being secondary), but that uncalcified cartilage was the primary skeletogenic tissue in the endoskeleton (bone – perichondral – being secondary).
• 17 Using evidence from developmental biology we conclude that the trunk neural crest of Ordovician agnathans was odontogenic, forming both dentine and bone of attachment of the trunk dermal exoskeleton.
• 18 Initiation of differentiation of skeletogenic and odontogenic tissues is controlled epigenetically by one or more epithelial-mesenchymal interactions in epigenetic cascades.
• 19 Changes in timing of steps in these epigenetic cascades provides an evolutionary mechanism for altering the types of skeletogenic/odontogenic tissues and/or structures formed.
• 20 The appearance of epithelial-mesenchymal interactions and the origin of the skeletogenic/odontogenic neural crest at the outset of vertebrate evolution provided the developmental basis for the evolutionary origin of vertebrate skeletogenic and odontogenic tissues and for the appearance and evolution of the vertebrate skeleton.

     

Histology of the heterostracan dermal skeleton: Insight into the origin of the vertebrate mineralised skeleton

Living vertebrates are divided into those that possess a fully formed and fully mineralised skeleton (gnathostomes) versus those that possess only unmineralised cartilaginous rudiments (cyclostomes). As such, extinct phylogenetic intermediates of these living lineages afford unique insights into the evolutionary assembly of the vertebrate mineralised skeleton and its canonical tissue types. Extinct jawless and jawed fishes assigned to the gnathostome stem evidence the piecemeal assembly of skeletal systems, revealing that the dermal skeleton is the earliest manifestation of a homologous mineralised skeleton. Yet the nature of the primitive dermal skeleton, itself, is poorly understood. This is principally because previous histological studies of early vertebrates lacked a phylogenetic framework required to derive evolutionary hypotheses. Nowhere is this more apparent than within Heterostraci, a diverse clade of primitive jawless vertebrates. To this end, we surveyed the dermal skeletal histology of heterostracans, inferred the plesiomorphic heterostracan skeleton and, through histological comparison to other skeletonising vertebrate clades, deduced the ancestral nature of the vertebrate dermal skeleton. Heterostracans primitively possess a four‐layered skeleton, comprising a superficial layer of odontodes composed of dentine and enameloid; a compact layer of acellular parallel‐fibred bone containing a network of vascular canals that supply the pulp canals (L1); a trabecular layer consisting of intersecting radial walls composed of acellular parallel‐fibred bone, showing osteon‐like development (L2); and a basal layer of isopedin (L3). A three layered skeleton, equivalent to the superficial layer L2 and L3 and composed of enameloid, dentine and acellular bone, is possessed by the ancestor of heterostracans + jawed vertebrates. We conclude that an osteogenic component is plesiomorphic with respect to the vertebrate dermal skeleton. Consequently, we interpret the dermal skeleton of denticles in chondrichthyans and jawless thelodonts as independently and secondarily simplified. J. Morphol. 276:657–680, 2015. © 2015 The Authors Journal of Morphology Published by Wiley Periodicals, Inc.     

Species-specific growth responses of favositid corals to soft-bottom substrates

Gibson, Michael A. & Broadhead, Thomas W. 1989 07 15: Species-specific growth responses of favositid corals to soft-bottom substrates. Lethaia , Vol. 22, pp. 287–299. Oslo. ISSN 0024–1164.
Species of favositid corals from the Upper Silurian and Lower Devonian of Tennessee, USA, exhibit structural modifications related to corallum geometry, interfacial skeletal material, and biotic associations that enabled them to survive in terrigenous mud rich environments. Favosites conicus Hall (Lower Devonian) had a flat, holotheca-covered base and a radial pattern of colony growth, but apparently had a short life span and may not have survived beyond the first reproductive cycle (monocarpous). It was adapted for living between major episodes of terrigenous mud influx. F. foerstei (Lower Devonian) had a convex, pseudoholotheca-covered base and a modified axial pattern of colony growth. Its large size, in comparison to that of F. conicus , suggests a longer lived colony (polycarpous), in which continued upward and outward growth enabled it to survive episodic sediment influx. F. forbesi (Upper Silurian) exhibited radial growth to form either (1) a globose corallum that was symbiotic with the stalks of living crinoids permitting the colony to live entirely above the substrate, or (2) a Gorallum with a steeply convex, holotheca-covered base that represents a bottom-dwelling colony in which the rate of growth probably only slightly exceeded the rate of sediment accumulation. * Functional morphology, astogeny, paleoecology, Tabulata .     

Mineralized cartilage in the skeleton of chondrichthyan fishes

The cartilaginous endoskeleton of chondrichthyan fishes (sharks, rays, and chimaeras) exhibits complex arrangements and morphologies of calcified tissues that vary with age, species, feeding behavior, and location in the body. Understanding of the development, evolutionary history and function of these tissue types has been hampered by the lack of a unifying terminology. In order to facilitate reciprocal illumination between disparate fields with convergent interests, we present levels of organization in which crystal orientation/size delimits three calcification types (areolar, globular, and prismatic) that interact in two distinct skeletal types, vertebral and tessellated cartilage. The tessellated skeleton is composed of small blocks (tesserae) of calcified cartilage (both prismatic and globular) overlying a core of unmineralized cartilage, while vertebral cartilage usually contains all three types of calcification.     

Ausgangsform und Entwicklung der rhombischen Schuppen der Osteichthyes (Pisces)

Ganoid and cosmoid scales, the two types of rhombic scales within the osteichthyans, can be traced back to a primitive scale similar to the scales ofLophosteus. The primitive rhombic scale did not have a peg-and-socket articulation, it is composed of lamellar bone superposed by many layers of spongy bone + dentine. That kind of superposition of layers of spongy bone + dentine (+ enamel) has been retained in the cosmoid scale; in contrast in the ganoid scale the growth of the dentine has become restricted to the lateral surface, the growth of ganoin to the outer surface and the growth of bone to the inner surface of the scale. The scales ofAndreolepis have a position between the primitive rhombic scale and the ganoid scale. — Scales from the Gedinnian (Lower Devonian) of New Sibirian Islands, USSR, are described asDialipina markae n. sp. The morphological features are very typical for the genus, but the histology is different from the type speciesD. salgueiroensis. Within the two Devonian palaeoniscoid generaDialipina andOrvikuina acellular bone with irregular non-vascular canals of Williamson has developed twice from cellular bone.     

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.     

Histological structure of the cancellous bone layer in Bothriolepis canadensis (Antiarchi, Placodermi)

The Placodermi are extinct basal gnathostomes which had extensive dermal and perichondral bone, but which lacked the endochondral bone which characterizes the more derived bony fishes. Thin sections of bone from a specimen of the antiarch placoderm Bothriolepis canadensis, from the Escuminac Formation (Frasnian, Upper Devonian), Québec, Canada, reveal that part of the cancellous layer in its dermal and endoskeletal bone formed from perichondral bone trabeculae growing around cartilage spheres. The resultant structure mimics that of osteichthyan endochondral bone. The layout and dimensions of this polygonal mosaic patterning of the bone trabeculae and flattened cartilage spheres resemble those of the prismatic layers of calcified cartilage in chondrichthyans. If the lack of endoskeletal bone in chondrichthyans is a derived character, then the structure identified in B. canadensis could represent a 'template' for the formation of prismatic calcified cartilage in the absence of bone.     

A new camuropiscid arthrodire (Pisces: Placodermi) from Gogo, Western Australia

A new camuropiscid arthrodire, Latocamurus coulthardi gen. et sp. nov., is described from the Upper Devonian Gogo Formation, Western Australia. Latocamurus , known from two complete specimens, is recognized as a camuropiscid by its narrow, spindle-shaped armour, deep postnasal plates participating in the orbits, preorbital plates which meet mesially, cheek unit firmly sutured to skull roof, posterior check plates tightly interconnected and much reduced, and the robust durophagous dentition. It is characterized by its downturned snout, broad, flat rostral plate, and narrow, deep parasphenoid. It is placed phyletically as the plesiomorphic sister taxon to all other camuropiscids which are more derived in having, inter alia , an anterior lateral plate which anteriorly contacts the anterior ventrolateral plate and pointed rostral plates. The family Camuropiscidae Dennis & Miles 1979b is redefined to incorporate features of the new genus. Camuropiscids and Incisoscutum are closely related by features of the postnasal plate and cheek.     

Histochemistry of lower vertebrate calcified structures. I. Enamel of the dogfish Squalus acanthius compared with mammalian enamel and homologous dentine

The enameloid and dentine of Squalus acanthius have been compared histochemically with those of Bos taurus. Squalus enameloid is much less reactive to a variety of stains or reagents than dentine or bovine immature enamel but it does have positive reactions with picromethyl blue, Mallory's and Van Gieson's stains, and Alcian blue. It stains faintly with Biebrich scarlet, indicating some anionic groups. Specific reactions for tyrosine, tryptophane, lysine, histidine, arginine, and cysteine are negative. Bos immature enamel is positive for cationic, anionic, and aromatic reactive groups by all test procedures, and dentine was positive for the anionic components. Bovine maturing enamel, however, is more similar in terms of lack of reactivity to Squalus enameloid but differed because the bovine enamel was moderately positive for tyrosine; tryptophane, and anionic groups and negative with Mallory's picromethyl blue and Van Gieson's stains. A fibrous transitional area between Squalus dentine and enameloid has staining reactions characteristic of both collagen and keratins.     

The fish swimming trace Undichna unisulca from the Silurian of Sweden: probably the oldest vertebrate locomotion trace fossil

The fish swimming trace Undichna unisulca is reported from a sandstone block collected from the Upper Silurian (Ludlow) Burgsvik Formation of Gotland, Sweden. It represents the oldest record of a vertebrate trace (apart from coprolites), predating previous finds from the Lower Devonian by at least 10 Ma. A thelodont or acanthodian fish may have produced the sinusoidal trails with the aid of their caudal or anal fins while browsing slowly over the sediment whilst searching for food.     

Devonian floral assemblages and plant megafossils from the Iberian Peninsula: A review

This paper presents a review and critical analysis of the literature on Devonian floras of the Iberian Peninsula. Although the known outcrops of Devonian strata in the Iberian Peninsula are marine, in some cases, a few fragmentary remains of vascular plants are associated with faunal remains. Records include largely specimens from the Lower Devonian of Barrancos (Alentejo, Portugal) and the Upper Devonian of Sierra de Hornachos in Badajoz province, southwest Spain; the remainder consists of drifted plant fragments from scattered sites in the Iberian Peninsula ranging in age from Lochkovian to Upper Devonian-Earliest Carboniferous. The vegetation inferred for the Lower Devonian of the Iberian Peninsula is mainly based on palynological data and corresponds to herbaceous types characterised by bryophytes, rhyniophytes (Horneophyton, Cooksonia, Rhynia), trimerophytes (Psilophyton, Pertica and Hostinella), primitive lycophytes (Drepanophycus) and incertae sedis such as Nothia and Chaleuria, all flora that developed near the coast in low-lying and, at least periodically, wet areas. In the Middle Devonian, two vegetation strata can be recognised: herbaceous (Psilophyton) and semi-arboreal (Cladoxylales). Although three levels of vegetation in the Upper Devonian, have been described from outcrops worldwide, the scarce available data from the Iberian Peninsula only indicate an arborescent lycopsid vegetation and species with uncertain botanical affinity such as Sphenopteridium keilhauii Nathorst.     

First Mesozoic Scutigeromorph Centipede, from the Lower Cretaceous of Brazil

The first Mesozoic scutigeromorph centipede (Myriapoda: Chilopoda), Fulmenocursor tenax gen. et sp. nov., is described from the Lower Cretaceous (Aptian) Crato Formation of north-east Brazil. Previously described fossil Scutigeromorpha are known from Dominican and Baltic amber, the Carboniferous (Westphalian D) Francis Creek Shale of Mazon Creek, Illinois, the Silurian and Devonian of Britain, and the Devonian of New York State.     

New insights into early land ecosystems: a glimpse of a lilliputian world

Small, relatively uncompressed, very fragmentary plant remains (mesofossils) are described from a Silurian (P ídolí) and a Lower Devonian (Lochkovian) locality in the Welsh Borderland. Excellent cellular preservation provides characters leading to the demonstration of diversity in plants of simple gross morphology and allows deliberations on functional anatomy (e.g. of stomata), and reproductive biology (including development and dehiscence of sporangia). A survey of in-situ spores is presented, and preliminary comparisons made with dispersed spore assemblages especially in relation to reconstruction of vegetation on local and regional scales. The earliest body fossils of unequivocal terrestrial arthropods isolated from the same locality as the P ídolí plants suggest that the decomposer/microherbivore/predator soil and litter communities found in the Lower and Middle Devonian extend back at least into the Silurian. Evidence for plant—animal interaction in the Lower Devonian comes from spore-dominated coprolites believed to have been produced by litter-feeding myriapods.     

华南中泥盆世艾菲尔期浅海相四射珊瑚的组合特征和古生物地理区系

虽然在华南中泥盆世艾菲尔期(Eifelian)较深水或斜坡相地层中产有许多国际标准的牙形类带化石,但在广阔的浅海相地层中却很难寻觅到其踪影。华南浅海相沉积地区是否缺失艾菲尔阶?在野外众多剖面的实地考察中表明,从下泥盆统埃姆斯阶到中泥盆统吉维特阶都是连续沉积,中间并没有发现地层的缺失或间断现象。浅海相的艾菲尔阶与其上覆的吉维特阶和下伏的埃姆斯阶都是连续沉积,说明可能是由于海水太浅,不太适合那些国际标准分子的生存而已。艾菲尔期中期末发生一次生物灭绝事件(Mid-Eifelian event),favositids,heliolitids和许多古老类型的珊瑚在地球上灭绝。华南艾菲尔期四射珊瑚可以划分成下、上两个完全不同的组合:1)Utara-tuiasinen sis-Sociop hyllum minor组合(牙形类partitus带—costatus带);2)Columnaria spinosa-Dendrostella praerhenana组合(牙形类australis带—kocklianus带)。     

Classification of uppermost Ordovician to Lower Devonian tubular and filamentous macerals from the Anglo-Welsh Basin

Dispersed tubes and filaments from the uppermost Ordovician, Silurian and Lower Devonian of the Anglo-Welsh Basin of Great Britain are described and classified using the artificial Anteturma system instituted for spores. The new Anteturma Trichomiformis is erected containing two new Turmae, Tubiformis and Filiformis. Within Tubiformis, the Infraturma Laevimurali, Endomurali and Extramurali are described, each containing a single genus with four, two and one species respectively. Within Filiformis, the Infraturma Ornatimurali is erected, based on a monospecific genus. Two assemblages of tubes and filaments are recognized. The older is found in uppermost Ordovician through to middle Silurian strata and is typified by a low diversity of smooth-walled tubes. The younger ranges from the middle Silurian at least to the Lower Devonian and is typified by various smooth-walled tubes, tubes with internal and external thickenings, and filaments. The majority of tubes and filaments are considered derived from the Nematophytales Lang.     

The histology and affinities of sinacanthid fishes: primitive gnathostomes from the Silurian of China

On the basis of well preserved specimens from the Lower Silurian of the Tarim Basin, Xinjiang Uygur Autonomous Region and Shiqian County, Guizhou Province, People's Republic of China we describe in detail the histological structure of sinacanthid spines, the only known remains of a group of fishes common in Silurian strata from China. The sinacanthids have previously been assigned either to the acanthodians or to the chondrichthyans. The spine structure is composed of an outer layer of atubular dentine and an inner layer of globular calcified cartilage, and the nature and distribution of these tissues indicates that the spines were formed as a result of interaction between the endoskeleton and dermoskeleton. The tissue distribution and style of growth described herein places the sinacanthids crownwards of the placoderms, and possibly within the total group Chondrichthyes. However, before they can be firmly placed within a phylogenetic scheme, further evidence is required both on the general anatomy of sinacanthids and on the nature of chondrichthyan apomorphies.  © 2005 The Linnean Society of London, Zoological Journal of the Linnean Society , 2005, 144 , 379–386.