Gavin C. Young

An exceptional Devonian fish from Australia sheds light on tetrapod origins

The transition from fishes to tetrapods was one of the most dramatic events in the evolution of vertebrates, but many pivotal fossils are incomplete, resulting in gaps in the data that are used for phylogenetic reconstruction. Here we present new observations from the most complete, acid-prepared Devonian tetrapodomorph fish yet discovered, Gogonasus, which was previously placed just crownward of Kenichthys and rhizodontids, the most primitive taxa on the tetrapod lineage. Unexpectedly, Gogonasus shows a mosaic of plesiomorphic and derived tetrapod-like features. Whereas the braincase and dermal cranial skeleton exhibit generalized morphologies with respect to Eusthenopteron or Panderichthys, taxa that are traditionally considered to be phyletically close to tetrapods, the presence of a deeply invaginated, wide spiracle, advanced internal spiracular architecture and near-horizontal hyomandibula are specialized features that are absent from Eusthenopteron. Furthermore, the pectoral fin skeleton of Gogonasus shares several features with that of Tiktaalik, the most tetrapod-like fish. A new phylogenetic analysis places Gogonasus crownward of Eusthenopteron as the sister taxon to the Elpistostegalia. Aspects of the basic tetrapod limb skeleton and middle ear architecture can now be traced further back within the tetrapodomorph radiation.

Live birth in the Devonian period

The extinct placoderm fishes were the dominant group of vertebrates throughout the Middle Palaeozoic era, yet controversy about their relationships within the gnathostomes (jawed vertebrates) is partly due to different interpretations of their reproductive biology. Here we document the oldest record of a live-bearing vertebrate in a new ptyctodontid placoderm, Materpiscis attenboroughi gen. et sp. nov., from the Late Devonian Gogo Formation of Australia (approximately 380 million years ago). The new specimen, remarkably preserved in three dimensions, contains a single, intra-uterine embryo connected by a permineralized umbilical cord. An amorphous crystalline mass near the umbilical cord possibly represents the recrystallized yolk sac. Another ptyctodont from the Gogo Formation, Austroptyctodus gardineri, also shows three small embryos inside it in the same position. Ptyctodontids have already provided the oldest definite evidence for vertebrate copulation, and the new specimens confirm that some placoderms had a remarkably advanced reproductive biology, comparable to that of some modern sharks and rays. The new discovery points to internal fertilization and viviparity in vertebrates as originating earliest within placoderms.

The most primitive osteichthyan braincase

Most living vertebrates, from teleosts to tetrapods, are osteichthyans (bony fishes), but the origin of this major group is poorly understood. The actinopterygians (ray-finned bony fishes) are the most successful living vertebrates in terms of diversity. They appear in the fossil record in the Late Silurian but are poorly known before the Late Devonian. Here we report the discovery of the oldest and most primitive actinopterygian-like osteichthyan braincase known, from 400–million-year-old limestone in southeastern Australia. This specimen displays previously unknown primitive conditions, in particular, an opening for a cartilaginous eyestalk. It provides an important and unique counterpart to the similarly aged and recently described Psarolepis from China and Vietnam. The contrasting features of these specimens, and the unusual anatomy of the new specimen in particular, provide new insights into anatomical conditions close to the evolutionary radiation of all modern osteichthyan groups.

A primitive actinopterygian neurocranium from the Early Devonian of southeastern Australia

Abstract A well-preserved neurocranium with attached partial skull roof is described from the Early Devonian Taemas Formation at Wee Jasper, southern New South Wales, Australia. The dermal ornament, skull roof pattern, and general proportion and structure of the endocranium are typically actinopterygian and the specimen is tentatively assigned to the actinopterygian genus Ligulalepsis. Other features more closely resemble some other groups, such as sarcopterygians (widely-spaced orbital walls, and short, broad telencephalon cavity), acanthodians (position of hyomandibular facet), and placoderms (many similarities including position of foramina for oculomotor, profundus, and trigeminal nerves, pituitary vein, and ophthalmic and orbital arteries in and around the orbit). This specimen is the first early osteichthyan to demonstrate the presence of an eyestalk, previously known only in placoderms and chondrichthyans. The unusual mix of characters and presence of an eyestalk provide new insights into primitive osteichthyan anatomy.

Ordovician microvertebrate remains from the Amadeus Basin, central Australia

ABSTRACT Microvertebrate remains are described from five formations (Pacoota Sandstone, Horn Valley Siltstone, Stairway Sandstone, Stokes Formation, Carmichael Sandstone) in the Ordovician sequence of the Amadeus Basin, central Australia. Two new genera and species are erected for scales and other elements with distinctive morphology and histology. Other material is referred to indeterminate species of the agnathan genera Arandaspis and Porophoraspis previously described from central Australia, and Sacabambaspis previously described from the Ordovician of Bolivia. The most common vertebrate from the Stokes Formation is referred to Areyonga oervigi, gen. et sp. nov., a form taxon provisionally assigned to the Chondrichthyes. Scales of this species apparently lack a base, and are made of an atubular laminar surface tissue of uncertain histology. Mode of scale growth was similar to that of the Early Devonian form Polymerolepis. If correctly interpreted, this taxon is the oldest chondrichthyan and/or gnathost...

Copulation in antiarch placoderms and the origin of gnathostome internal fertilization

Reproduction in jawed vertebrates (gnathostomes) involves either external or internal fertilization. It is commonly argued that internal fertilization can evolve from external, but not the reverse. Male copulatory claspers are present in certain placoderms, fossil jawed vertebrates retrieved as a paraphyletic segment of the gnathostome stem group in recent studies. This suggests that internal fertilization could be primitive for gnathostomes, but such a conclusion depends on demonstrating that copulation was not just a specialized feature of certain placoderm subgroups. The reproductive biology of antiarchs, consistently identified as the least crownward placoderms and thus of great interest in this context, has until now remained unknown. Here we show that certain antiarchs possessed dermal claspers in the males, while females bore paired dermal plates inferred to have facilitated copulation. These structures are not associated with pelvic fins. The clasper morphology resembles that of ptyctodonts, a more crownward placoderm group, suggesting that all placoderm claspers are homologous and that internal fertilization characterized all placoderms. This implies that external fertilization and spawning, which characterize most extant aquatic gnathostomes, must be derived from internal fertilization, even though this transformation has been thought implausible. Alternatively, the substantial morphological evidence for placoderm paraphyly must be rejected.

Fossil musculature of the most primitive jawed vertebrates

From Jawless to Jawed The earliest vertebrates were jawless. Past reconstructions have assumed that the primitive jawed condition was much like that found in sharks. Trinajstic et al. (p. 160, published online 13 June; see the Perspective by Kuratani) describe fossil musculature from the early jawed placoderms (an extinct class of armored prehistoric fish) that show that the basal structure was distinct from that found in sharks, having a notable dermal joint between the skull and shoulder girdle. Fossilized muscles in an ancient armored fish reveal a distinctive neck musculature that differs from that of sharks. [Also see Perspective by Kuratani] The transition from jawless to jawed vertebrates (gnathostomes) resulted in the reconfiguration of the muscles and skeleton of the head, including the creation of a separate shoulder girdle with distinct neck muscles. We describe here the only known examples of preserved musculature from placoderms (extinct armored fishes), the phylogenetically most basal jawed vertebrates. Placoderms possess a regionalized muscular anatomy that differs radically from the musculature of extant sharks, which is often viewed as primitive for gnathostomes. The placoderm data suggest that neck musculature evolved together with a dermal joint between skull and shoulder girdle, not as part of a broadly flexible neck as in sharks, and that transverse abdominal muscles are an innovation of gnathostomes rather than of tetrapods.

New discoveries of Devonian vertebrates from South America, and implications for Gondwana-Euramerica contact.

A new fossil fish fauna from western Venezuela provides the first South American Devonian record of antiarch and phyllolepid placoderms, and sarcopterygians including lungfishes, groups widely distributed on other continents. Endemic elements in the fish fauna indicate Gondwana affinities, and the age of the phyllolepid placoderm occurrence is consistent with a model of biotic dispersal between Gondwana and Euramerica near the Frasnian–Famennian boundary. A narrow marine barrier separating northern and southern landmasses is indicated, in contrast to the wide equatorial ocean for the Late Devonian postulated from palaeomagnetic data.Abstract A new fossil fish fauna from western Venezuela provides the first South American Devonian record of antiarch and phyllolepid placoderms, and sarcopterygians including lungfishes, groups widely distributed on other continents. Endemic elements in the fish fauna indicate Gondwana affinities, and the age of the phyllolepid placoderm occurrence is consistent with a model of biotic dispersal between Gondwana and Euramerica near the Frasnian–Famennian boundary. A narrow marine barrier separating northern and southern landmasses is indicated, in contrast to the wide equatorial ocean for the Late Devonian postulated from palaeomagnetic data.

PRIMITIVE JAW STRUCTURE IN AN ARTICULATED BRACHYTHORACID ARTHRODIRE (PLACODERM FISH; EARLY DEVONIAN) FROM SOUTHEASTERN AUSTRALIA

Abstract A unique ‘buchanosteid’ arthrodire specimen from Emsian limestones at Burrinjuck represents the complete articulated remains of the head and cheek, and most of the trunk armor bones from a single fish, together with well-ossified braincase and jaw cartilages. The structure of the toothplates and jaws are described, and compared with primitive brachythoracid material from Saudi Arabia of similar age. Both the palatoquadrate and meckelian cartilages are well preserved, and perichondrally ossified as single elements. As previously described in some phlyctaeniids, the dermal gnathal elements carry crowded denticles similar to the normal tubercular ornament, enlarged along anterior and lateral margins. Six characters concerning gnathal plates of brachythoracids, which have been used in phylogenetic analysis, are discussed and reformulated.

An articulated phyllolepid fish (Placodermi) from the Devonian of central Australia: implications for non-marine connections with the Old Red Sandstone continent

A second species of the placoderm genus Placolepis ( Pl. harajica sp. nov.), based on a single articulated specimen from Givetian–Frasnian strata in the MacDonnell Ranges, demonstrates the occurrence of this taxon across the Australian craton. Placolepis (order Phyllolepida) is endemic to east Gondwana, and other phyllolepids are widespread in the Givetian and younger of Gondwana (Australia, Antarctica, Turkey, Venezuela), but do not occur until Late Devonian (Famennian) time in the Northern Hemisphere (Europe, Russia, Greenland, North America). The disjunct space–time distribution of the Phyllolepida is inconsistent with palaeomagnetic evidence indicating a wide equatorial ocean between Gondwana and Laurussia in Late Devonian time. This new species provides additional evidence supporting a Gondwana origin for the group, and later access to northern landmasses resulting from closure of the ocean between Gondwana and Laurussia and continental connection at or near the Frasnian–Famennian boundary.