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Concept: Devonian

189

A nautiloid conch containing many disarticulated exoskeletons of Omegops cornelius (Phacopidae, Trilobita) was found in the Upper Devonian Hongguleleng Formation of the northwestern margin of the Junggar Basin, NW China. The similar number of cephala, thoraces and pygidia, unbroken thoraces, explicit exuviae, and lack of other macrofossils in the conch, indicate that at least seven individual trilobites had moulted within the nautiloid living chamber, using the vacant chamber of a dead nautiloid as a communal place for ecdysis. This exuvial strategy manifests cryptic behaviour of trilobites, which may have resulted from the adaptive evolution induced by powerful predation pressure, unstable marine environments, and competition pressure of organisms occupying the same ecological niche in the Devonian period. The unusual presence of several trilobites moulting within a nautiloid conch is possibly associated with social behaviours in face of a serious crisis. New materials in this study open a window for understanding the survival strategy of marine benthic organisms, especially predator-prey interactions and the behavioural ecology of trilobites in the middle Palaeozoic.

Concepts: Arthropod, Predation, Ecology, Carboniferous, Ordovician, Trilobite, Devonian, Ecdysis

181

gen. et sp. nov. is described from the Famennian Worange Point Formation; the holotype is amongst the largest tristichopterids and sarcopterygians documented by semi-articulated remains from the Devonian Period. The new taxon has dentary fangs and premaxillary tusks, features assumed to be derived for large Northern Hemisphere tristichopterids (, , ). It resembles in ornament, but is distinguished by longer proportions of the parietal compared to the post-parietal shield, and numerous differences in shape and proportions of other bones. Several characters (accessory vomers in the palate, submandibulars overlapping ventral jaw margin, scales ornamented with widely-spaced deep grooves) are recorded only in tristichopterids from East Gondwana (Australia-Antarctica). On this evidence gen. nov. is placed in an endemic Gondwanan subfamily Mandageriinae within the Tristichopteridae; it differs from the nominal genotype in its larger size, less pointed skull, shape of the orbits and other skull characters. The hypothesis that tristichopterids evolved in Laurussia and later dispersed into Gondwana, and a derived subgroup of large Late Devonian genera dispersed from Gondwana, is inconsistent with the evidence of the new taxon. Using oldest fossil and most primitive clade criteria the most recent phylogeny resolves South China and Gondwana as areas of origin for all tetrapodomorphs. The immediate outgroup to tristichopterids remains unresolved - either from Greenland as recently proposed, or from Gondwana, earlier suggested to be the sister group to all tristichopterids. Both taxa combine two characters that do not co-occur in other tetrapodomorphs (extratemporal bone in the skull; non-cosmoid round scales with an internal boss). Recently both ‘primitive’ and ‘derived’ tristichopterids have been discovered in the late Middle Devonian of both hemispheres, implying extensive ghost lineages within the group. Resolving their phylogeny and biogeography will depend on a comprehensive new phylogenetic analysis.

Concepts: Phylogenetic nomenclature, Phylogenetics, Cladistics, Vertebrate, Sarcopterygii, Tetrapod, Carboniferous, Devonian

68

The vertebrate recovery after the end-Permian mass extinction can be approached through the ichnological record, which is much more abundant than body fossils. The late Olenekian (Early Triassic) tetrapod ichnoassemblage of the Catalan Pyrenean Basin is the most complete and diverse of this age from Western Tethys. This extensional basin, composed of several depocenters, was formed in the latest phases of the Variscan orogeny (Pangea breakup) and was infilled by braided and meandering fluvial systems of the red-beds Buntsandstein facies. Abundant and diverse tetrapod ichnites are recorded in these facies, including Prorotodactylus mesaxonichnus isp. nov. (tracks possibly produced by euparkeriids), cf. Rotodactylus, at least two large chirotheriid morphotypes (archosauriform trackmakers), Rhynchosauroides cf. schochardti, two other undetermined Rhynchosauroides forms, an undetermined Morphotype A (archosauromorph trackmakers) and two types of Characichnos isp. (swimming traces, here associated to archosauromorph trackmakers). The Pyrenean ichnoassemblage suggests a relatively homogeneous ichnofaunal composition through the late Early Triassic of Central Pangea, characterized by the presence of Prorotodactylus and Rotodactylus. Small archosauromorph tracks dominate and present a wide distribution through the different fluviatile facies of the Triassic Pyrenean Basin, with large archosaurian footprints being present in a lesser degree. Archosauromorphs radiated and diversified through the Triassic vertebrate recovery, which ultimately lead to the archosaur and dinosaur dominance of the Mesozoic.

Concepts: Dinosaur, Carboniferous, Pyrenees, Triassic, Archosaur, Permian, Archosauromorpha, Devonian

42

Tetrapods evolved from sarcopterygian fishes in the Devonian and were the first vertebrates to colonize land. The locomotor component of this transition can be divided into four major events: terrestriality, the origins of digited limbs, solid substrate-based locomotion, and alternating gaits that use pelvic appendages as major propulsors. As the sister group to tetrapods, lungfish are a morphologically and phylogenetically relevant sarcopterygian taxon for understanding the order in which these events occurred. We found that a species of African lungfish (Protopterus annectens) uses a range of pelvic fin-driven, tetrapod-like gaits, including walking and bounding, in an aquatic environment, despite having a derived limb endoskeleton and primitively small, muscularly supported pelvis. Surprisingly, given these morphological traits, P. annectens also lifts its body clear of the substrate using its pelvic fins, an ability thought to be a tetrapod innovation. Our findings suggest that some fundamental features of tetrapod locomotion, including pelvic limb gait patterns and substrate association, probably arose in sarcopterygians before the origin of digited limbs or terrestriality. It follows that the attribution of some of the nondigited Devonian fossil trackways to limbed tetrapods may need to be revisited.

Concepts: Fish, Vertebrate, Sarcopterygii, Tetrapod, Fossil, Coelacanth, Osteichthyes, Devonian

40

Pentastomids (tongue worms) are worm-like arthropods known today from ∼140 species [1]. All but four are parasitic on vertebrates. Their life cycle typically involves larval development in an intermediate host followed by maturation in the respiratory tract of a definitive terrestrial host. Fossil pentastomids are exceedingly rare and are known only from isolated juveniles [2-6]. The identity of the possible hosts of fossil pentastomids and the origin of their lifestyle have generated much debate. A new, exceptionally preserved species, described based on adults from 425-million-year-old marine rocks, is the only known fossil pentastomid associated with a host, in this case a species of ostracod crustacean. The pentastomids are preserved near eggs within the ostracod and also, uniquely for any fossil or living pentastomid, are attached externally to the host. This discovery affirms the origin of pentastomids as ectoparasitic on marine invertebrates. The terrestrialization of pentastomids may have occurred in parallel with the vertebrate invasion of land.

Concepts: Species, Insect, Crustacean, Vertebrate, Annelid, Invertebrate, Maxillopoda, Devonian

36

Enamel, the hardest vertebrate tissue, covers the teeth of almost all sarcopterygians (lobe-finned bony fishes and tetrapods) as well as the scales and dermal bones of many fossil lobe-fins. Enamel deposition requires an organic matrix containing the unique enamel matrix proteins (EMPs) amelogenin (AMEL), enamelin (ENAM) and ameloblastin (AMBN). Chondrichthyans (cartilaginous fishes) lack both enamel and EMP genes. Many fossil and a few living non-teleost actinopterygians (ray-finned bony fishes) such as the gar, Lepisosteus, have scales and dermal bones covered with a proposed enamel homologue called ganoine. However, no gene or transcript data for EMPs have been described from actinopterygians. Here we show that Psarolepis romeri, a bony fish from the the Early Devonian period, combines enamel-covered dermal odontodes on scales and skull bones with teeth of naked dentine, and that Lepisosteus oculatus (the spotted gar) has enam and ambn genes that are expressed in the skin, probably associated with ganoine formation. The genetic evidence strengthens the hypothesis that ganoine is homologous with enamel. The fossil evidence, further supported by the Silurian bony fish Andreolepis, which has enamel-covered scales but teeth and odontodes on its dermal bones made of naked dentine, indicates that this tissue originated on the dermal skeleton, probably on the scales. It subsequently underwent heterotopic expansion across two highly conserved patterning boundaries (scales/head-shoulder and dermal/oral) within the odontode skeleton.

Concepts: Fish, Vertebrate, Chordate, Actinopterygii, Sarcopterygii, Tetrapod, Osteichthyes, Devonian

33

Our understanding of early gnathostome evolution has been hampered by a generally scant fossil record beyond the Devonian. Recent discoveries from the late Silurian Xiaoxiang Fauna of Yunnan, China, have yielded significant new information, including the earliest articulated osteichthyan fossils from the Ludlow-aged Kuanti Formation. Here we describe the partial postcranium of a new primitive bony fish from the Kuanti Formation that represents the second known taxon of pre-Devonian osteichthyan revealing articulated remains. The new form, Sparalepis tingi gen. et sp. nov., displays similarities with Guiyu and Psarolepis, including a spine-bearing pectoral girdle and a placoderm-like dermal pelvic girdle, a structure only recently identified in early osteichthyans. The squamation with particularly thick rhombic scales shares an overall morphological similarity to that of Psarolepis. However, the anterior flank scales of Sparalepis possess an unusual interlocking system of ventral bulges embraced by dorsal concavities on the outer surfaces. A phylogenetic analysis resolves Sparalepis within a previously recovered cluster of stem-sarcopterygians including Guiyu, Psarolepis and Achoania. The high diversity of osteichthyans from the Ludlow of Yunnan strongly contrasts with other Silurian vertebrate assemblages, suggesting that the South China block may have been an early center of diversification for early gnathostomes, well before the advent of the Devonian “Age of Fishes”.

Concepts: Fish, Vertebrate, Gnathostomata, Sarcopterygii, Fossil, Trilobite, Devonian, Silurian

33

Devonian tetrapods (limbed vertebrates), known from an increasingly large number of localities, have been shown to be mainly aquatic with many primitive features. In contrast, the post-Devonian record is marked by an Early Mississippian temporal gap ranging from the earliest Carboniferous (Tournaisian and early Viséan) to the mid-Viséan. By the mid-Viséan, tetrapods had become effectively terrestrial as attested by the presence of stem amniotes, developed an essentially modern aspect, and given rise to the crown group. Up to now, only two localities have yielded tetrapod specimens from the Tournaisian stage: one in Scotland with a single articulated skeleton and one in Nova Scotia with isolated bones, many of uncertain identity. We announce a series of discoveries of Tournaisian-age localities in Scotland that have yielded a wealth of new tetrapod and arthropod fossils. These include both terrestrial and aquatic forms and new taxa. We conclude that the gap in the fossil record has been an artifact of collection failure.

Concepts: Sarcopterygii, Tetrapod, Synapsid, Fossil, Paleontology, Amphibian, Carboniferous, Devonian

29

Osteichthyans comprise two divisions, each containing over 32,000 living species [1]: Sarcopterygii (lobe-finned fishes and tetrapods) and Actinopterygii (ray-finned fishes). Recent discoveries from China highlight the morphological disparity of early sarcopterygians and extend their origin into the late Silurian [2-4]. By contrast, the oldest unambiguous actinopterygians are roughly 30 million years younger, leaving a long temporal gap populated by fragments and rare body fossils of controversial phylogenetic placement [5-10]. Here we reinvestigate the enigmatic osteichthyan Meemannia from the Early Devonian (∼415 million years ago) of China, previously identified as an exceptionally primitive lobe-finned fish [3, 7, 11, 12]. Meemannia combines “cosmine”-like tissues taken as evidence of sarcopterygian affinity with actinopterygian-like skull roof and braincase geometry, including endoskeletal enclosure of the spiracle and a lateral cranial canal. We report comparable histological structures in undoubted ray-finned fishes and conclude that they are general osteichthyan features. Phylogenetic analysis places Meemannia as an early-diverging ray-finned fish, resolving it as the sister lineage of Cheirolepis [13] plus all younger actinopterygians. This brings the first appearance of ray-fins more in line with that of lobe-fins and fills a conspicuous faunal gap in the otherwise diverse late Silurian-earliest Devonian vertebrate faunas of the South China Block [4].

Concepts: Fish, Vertebrate, Chordate, Actinopterygii, Sarcopterygii, Tetrapod, Osteichthyes, Devonian

27

The oldest-known air-breathing land animal is the millipede Pneumodesmus newmani, found in the Cowie Harbour Fish Bed at Stonehaven, Scotland. Here we report the youngest, most concordant 238U-206Pb zircon age from ash below the fish bed of 413.7±4.4 Ma (±2σ), whereas the youngest age from a tuffaceous sandstone above the fish bed is statistically indistinguishable at 414.3±7.1 Ma. The Cowie Harbour Fish Bed thus appears to be lowermost Devonian (Lochkovian), contrary to the previously accepted mid-Silurian age based on palynomorphs from adjacent exposures. This has implications for the evolutionary timetable of land colonization, as the Cowie ages overlap late Lochkovian zircon ages reported elsewhere for andesite below the nearby (~50 mi) Rhynie Chert, which has more advanced terrestrial biota. The results postdate the possible late Silurian Ludford Lane locality in Shropshire, England. Pneumodesmus newmani is thus not the earliest air-breathing land animal, unless the Ludford Lane locality is younger than presently assigned.

Concepts: Insect, Trilobite, Devonian, Silurian, Millipede, Pneumodesmus newmani, Stonehaven, Ludlow