Per Ahlberg

Uppsala University | UU · Department of Organismal Biology

Trilobites are among the most iconic of fossils and formed a prominent component of marine ecosystems during most of their 270-million-year-long history from the early Cambrian period to the end Permian period1. More than 20,000 species have been described to date, with presumed lifestyles ranging from infaunal burrowing to a planktonic life in...

Bone is an evolutionary novelty of vertebrates, likely to have first emerged as part of ancestral dermal armor that consisted of osteogenic and odontogenic components. Whether these early vertebrate structures arose from mesoderm or neural crest cells has been a matter of considerable debate. To examine the developmental origin of the bony part of...

The Middle Devonian (Givetian) Valentia Slate Formation in the Iveragh Peninsula, southwest Ireland, is more renowned for the second oldest record of tetrapod trackways in the world than for its heavily metamorphosed bone remains. The present study focuses on new discoveries of non-tetrapod sarcopterygian fish fossils from the Valentia Slate Format...

Jensen et al. (1) question evidence presented of a chambered heart within placoderms, citing its small size and apparently ventral atrium. However, they fail to note the belly-up orientation of the placoderm within one nodule, and the variability of heart morphology within extant taxa. Thus, we remain confident in our interpretation of the minerali...

The recently discovered mass mortality of fishes from the Tanis Site in the North Dakota portion of the Late Cretaceous Hell Creek Formation contains many well-preserved, three-dimensional skeletons. Among these are representatives of two acipenseriform families, Acipenseridae (sturgeons) and Polyodontidae (paddlefishes). This paper describes two n...

Material of the antiarch placoderm Bothriolepis from the middle Givetian of the Valentia Slate Formation in Iveragh Peninsula, Ireland, is described and attributed to a new species, B. dairbhrensis sp. nov. A revision of the genus Bothriolepis is proposed, and its taxonomic content and previous phylogenetic analyses are reviewed, as well as the val...

We describe the largest bony fish in the Late Devonian (late Famennian) fossil assemblage from Waterloo Farm near Makhanda/Grahamstown, South Africa. It is a giant member of the extinct clade Tristichopteridae (Sarcopterygii: Tetrapodomorpha) and most closely resembles Hyneria lindae from the late Famennian Catskill Formation of Pennsylvania, USA....

In a recent paper by DePalma et al., the End-Cretaceous mass extinction was calibrated to spring/summer on the basis of stable isotopic analyses and other data. An independent study, which was concurrently in review and known to DePalma et al,. reached the same conclusion using osteohistology and stable isotope analysis. Notwithstanding this agreem...

Molecular studies suggest that the origin of jawed vertebrates was no later than the Late Ordovician period (around 450 million years ago (Ma))1,2. Together with disarticulated micro-remains of putative chondrichthyans from the Ordovician and early Silurian period3–8, these analyses suggest an evolutionary proliferation of jawed vertebrates before,...

The origin and early diversification of jawed vertebrates involved major changes to skeletal and soft anatomy. Skeletal transformations can be examined directly by studying fossil stem gnathostomes; however, preservation of soft anatomy is rare. We describe the only known example of a three-dimensionally mineralized heart, thick-walled stomach, and...

The early Tournaisian (Carboniferous) stage represents a key episode in the evolution of vertebrates. It follows the end-Devonian Hangenberg extinction event, which led to a major perturbation to both terrestrial and aquatic vertebrate ecosystems, and resulted in a significant restructuring of assemblages. However, few faunal associations of this a...

The lobe-finned fish, lungfish (Dipnoi, Sarcoptergii), have persisted for ~400 million years from the Devonian Period to present day. The evolution of their dermal skull and dentition is relatively well understood, but this is not the case for the central nervous system. While the brain has poor preservation potential and is not currently known in...

The spiracular region, comprising the hyomandibular pouch together with the mandibular and hyoid arches, has a complex evolutionary history. In living vertebrates, the embryonic hyomandibular pouch may disappear in the adult, develop into a small opening between the palatoquadrate and hyomandibula containing a single gill-like pseudobranch, or crea...

We present an updated time frame for the 30 m thick late Miocene sedimentary Trachilos section from the island of Crete that contains the potentially oldest hominin footprints. The section is characterized by normal magnetic polarity. New and published foraminifera biostratigraphy results suggest an age of the section within the Mediterranean biozo...

Sharks are iconic predators in today’s oceans, yet their modern diversity has ancient origins. In particular, present hypotheses suggest that a combination of mass extinction, global climate change, and competition has regulated the community structure of dominant mackerel (Lamniformes) and ground (Carcharhiniformes) sharks over the last 66 million...

Lungfish (Dipnoi) are lobe-finned fish (Sarcopterygii) that have persisted for over 400 million years from the Devonian Period to present day. They are the extant sister group to tetrapods and thus have the ability to provide unique insight into the condition of the earliest tetrapods as well as their own evolutionary history. The evolution of thei...

Sharks (Selachimorpha) are iconic marine predators that have survived multiple mass extinctions over geologic time. Their prolific fossil record is represented mainly by isolated shed teeth, which provide the basis for reconstructing deep time diversity changes affecting different selachimorph clades. By contrast, corresponding shifts in shark ecol...

Significance Some lineages of organisms have undergone major evolutionary radiations, while others have not. Establishing why is a central goal of evolutionary research. Whole-genome duplication (WGD) is often proposed as having caused the spectacular evolutionary radiation of teleost fishes. However, due to the absence of genetic data for fossil s...

Teleost fishes comprise one-half of all vertebrate species and possess a duplicated genome. This whole-genome duplication (WGD) occurred on the teleost stem lineage in an ancient common ancestor of all living teleosts and is hypothesized as a trigger of their exceptional evolutionary radiation. Genomic and phylogenetic data indicate that WGD occurr...

The Triassic was a crucial period for the early evolution and diversification of insects, including Coleoptera the most diverse order of organisms on Earth. The study of Triassic beetles, however, relies almost exclusively on flattened fossils with limited character preservation. Using synchrotron microtomography, we investigated a fragmentary Uppe...

Jennifer Clack (née Agnew) dedicated her entire research career of more than 40 years to the fish-tetrapod transition, the evolutionary process during the Devonian and Carboniferous periods that transformed a lineage of lobe-finned fishes into the earliest land vertebrates. She was widely regarded as the world leader in this field. During an expedi...

Our understanding of the earliest evolution of jawed vertebrates depends on a credible phylogenetic framework for the jawed stem gnathostomes collectively known as “placoderms”.¹ However, their relationships, and whether placoderms represent a single radiation or a paraphyletic array, remain contentious.2, 3, 4, 5, 6, 7, 8, 9, 10, 11, 12, 13 This u...

Sharks (Selachimorpha) are iconic marine predators that have survived multiple mass extinctions over geologic time. Their fossil record is represented by an abundance of teeth, which traditionally formed the basis for reconstructing large-scale diversity changes among different selachimorph clades. By contrast, corresponding patterns in shark ecolo...

The ontogenetic trajectory of a marginal jawbone of Lophosteus superbus (Late Silurian, 422 Million years old), the phylogenetically most basal stem osteichthyan, visualized by synchrotron microtomography, reveals a developmental relationship between teeth and dermal odontodes that is not evident from the adult morphology. The earliest odontodes ar...

[This corrects the article DOI: 10.1098/rspa.2020.0355.].

Tides are a major component of the interaction between the marine and terrestrial environments, and thus play an important part in shaping the environmental context for the evolution of shallow marine and coastal organisms. Here, we use a dedicated tidal model and palaeogeographic reconstructions from the Late Silurian to early Late Devonian (420 M...

Our understanding of the earliest evolution of jawed vertebrates depends on a credible phylogenetic assessment of the jawed stem gnathostomes collectively known as ‘placoderms’. However, their relationships, and even whether ‘placoderms’ represent a single radiation or a paraphyletic array, remain contentious. Here we describe the endocranial cavit...

The first dinosaur embryos found inside megaloolithid eggs from Auca Mahuevo, Patagonia, were assigned to sauropod dinosaurs that lived approximately 80 million years ago. Discovered some 25 years ago, these considerably flattened specimens still remain the only unquestionable embryonic remains of a sauropod dinosaur providing an initial glimpse in...

Ontogenetic data obtained by synchrotron microtomography of a marginal jawbone of Lophosteus superbus (Late Silurian, 422 Million years old), the phylogenetically basalmost stem osteichthyan, reveal developmental relationships between teeth and ornament that are not obvious from the adult morphology. The earliest odontodes are two longitudinal foun...

Teeth and jaws The first vertebrates were jawless, much like a modern hagfish. There has been a lot of interest in how these forms transitioned to having jaws like most of their descendants, including humans. Much of our understanding of this process has focused on how the teeth are replaced relative to the jaw. Previous theories suggested that too...

Porolepiform scales from the Lower Devonian of the Holy Cross Mountains, Poland, described from the ‘placoderm sandstone’ of the Daleszyce area, are revised. The aim of the present article is to organize the recently collected, but not formally described, porolepiform material from the Holy Cross Mountains, as well as specimens from erratic boulder...

A new genus and species of Devonian tetrapod, Brittagnathus minutus gen. et sp. nov., is described from a single complete right lower jaw ramus recovered from the Acanthostega mass-death deposit in the upper part of the Britta Dal Formation (upper Famennian) of Stensiö Bjerg, Gauss Peninsula, East Greenland. Visualization by propagation phase contr...

Palaeontologist who described how vertebrates moved from water to land. Palaeontologist who described how vertebrates moved from water to land.

Our understanding of the earliest evolution of jawed vertebrates depends on a credible phylogenetic framework for the jawed stem gnathostomes collectively known as "placoderms".(1) However, their relationships, and whether placoderms represent a single radiation or a paraphyletic array, remain contentious.(2-13) This uncertainty is compounded by an...

In-situ minor, trace and rare earth element (REE) compositions have been measured in conodonts from the lower Silurian Vesiku Bone Bed (Estonia), with the aim to visualize and to quantify conodont apatite elemental geochemistry, to evaluate compositional differences between conodont histologies, and to evaluate the use of conodont chemistry as a pa...

The known diversity of tetrapods of the Devonian period has increased markedly in recent decades, but their fossil record consists mostly of tantalizing fragments1–15. The framework for interpreting the morphology and palaeobiology of Devonian tetrapods is dominated by the near complete fossils of Ichthyostega and Acanthostega; the less complete, b...

Diets of pterosaurs have mainly been inferred from indirect evidence such as comparative anatomy, associations of co-occurring fossils, and functional morphology. Gut contents are rare, and until now there is only a single coprolite (fossil dropping), with unidentified inclusions, known. Here we describe three coprolites collected from a palaeosurf...

Diets of extinct animals can be difficult to analyse if no direct evidence, such as gut contents, is preserved in association with body fossils. Inclusions from coprolites (fossil faeces), however, may also reflect the diet of the host animal and become especially informative if the coprolite producer link can be established. Here we describe, base...

Diets of extinct animals can be difficult to analyse if no direct evidence, such as gut contents, is preserved in association with body fossils. Inclusions from coprolites (fossil faeces), however, may also reflect the diet of the host animal and become especially informative if the coprolite producer link can be established. Here we describe, base...

Fossils, function and phylogeny: Papers on early vertebrate evolution in honour of Professor Jennifer A. Clack – Introduction - Volume 109 Issue 1-2 - Marcello RUTA, Per E. AHLBERG, Timothy R. SMITHSON

Here we present evidence for osteophagy in the Late Triassic archosaur Smok wawelski Niedźwiedzki, Sulej and Dzik, 2012, a large theropod-like predator from Poland. Ten medium to large-sized coprolites are matched, by their dimensions and by association with body fossils and footprints, to S. wawelski. The coprolites contain fragments of large serr...

From an initial isolated position as the oldest evolutionary prototype of a bird, Archaeopteryx has, as a result of recent fossil discoveries, become embedded in a rich phylogenetic context of both more and less crownward stem-group birds. This has prompted debate over whether Archaeopteryx is simply a convergently bird-like non-avialan theropod. H...

Evolution and Development of Fishes - edited by Zerina Johanson January 2019

Hyneria lindae is one of the largest Devonian sarcopterygians. It was found in the Catskill Formation (late Famennian) of Pennsylvania, USA. The current study focuses on the palaeohistology of the humerus of this tristichopterid and supports a low ossification rate and a late ossification onset in the appendicular skeleton. In addition to anatomica...

The subdivision of the gnathostome neurocranium into an anterior neural crest-derived moiety and a posterior mesodermal moiety has attracted the interest of researchers for nearly two centuries. We present a synthetic scenario for the evolution of this structure, uniting developmental data from living cyclostomes and gnathostomes with morphological...

Ichthyosaurs are extinct marine reptiles that display a notable external similarity to modern toothed whales. Here we show that this resemblance is more than skin deep. We apply a multidisciplinary experimental approach to characterize the cellular and molecular composition of integumental tissues in an exceptionally preserved specimen of the Early...

Significance The fossil fish Rhizodus hibberti , a member of the tetrapod stem group, shows a unique skeletal pattern in the pelvic fin. Rather than the highly conserved one-to-two pattern of a femur, tibia, and fibula (seen in all known tetrapods, including the extinct, fishlike members of the group), the fin of Rhizodus comprises a femur articula...

The hypothesis that tetrapods evolved from elpistostegids during the Frasnian, in a predominantly aquatic context, has been challenged by the discovery of Middle Devonian tetrapod trackways predating the earliest body fossils of both elpistostegids and tetrapods. Here I present a new hypothesis based on an overview of the trace fossil and body foss...

The Cretaceous-Palaeogene (K-Pg) mass extinction profoundly altered vertebrate ecosystems and prompted the radiation of many extant clades [1, 2]. Sharks (Selachimorpha) were one of the few larger-bodied marine predators that survived the K-Pg event and are represented by an almost-continuous dental fossil record. However, the precise dynamics of t...

The first half of the Mississippian or Early Carboniferous (Tournaisian to mid- Viséan), an interval of about 20 million years, has become known as “Romer's Gap” because of its poor tetrapod record. Recent discoveries emphasise the differences between pre-“Gap” Devonian tetrapods, unambiguous stem-group members retaining numerous “fish” characters...

Out of Antarctica When we think of Devonian tetrapods, the ancestors of all modern vertebrates, we tend to picture amphibian-like creatures emerging from the water into a wet tropical forest or swamp. Indeed, all previously described specimens of this group have been recovered from the tropics. Gess and Ahlberg now describe two fossil tetrapods fro...

The skull of 'Ligulalepis' from the Early Devonian of Australia (AM-F101607) has significantly expanded our knowledge of early osteichthyan anatomy, but its phylogenetic position has remained uncertain. We herein describe a second skull of 'Ligulalepis' and present micro-CT data on both specimens to reveal novel anatomical features, including crani...

Opinions differ on whether the evolution of tetrapods (limbed vertebrates) from lobe-finned fishes was directly linked to terrestrialization. The earliest known tetrapod fossils, from the Middle Devonian (approximately 390 million years old) of Zachełmie Quarry in Poland, are trackways made by limbs with digits; they document a direct environmental...

There are two types of dermal skeletons in jawed vertebrates: placoderms and osteichthyans carry large bony plates (macromery), whereas chondrichthyans and acanthodians are covered by small scales (micromery). Fin spines are one of the last large dermal structures found on micromeric taxa and offer a potential source of histology and morphology tha...

Thin section (S2368) of a dermal plate of Romundina, deposited at the Natural History Museum of Stockholm. Arrows mark the cell lacunae and arrowheads mark the dentine tubules. Note the different colors of the outer layer and the inner layer, both of which have dentine tubules and cell lacunae. (TIF)

The fossils assigned to the tetrapod stem group document the evolution of terrestrial vertebrates from lobe-finned fishes. During the past 18 years the phylogenetic structure of this stem group has remained remarkably stable, even when accommodating new discoveries such as the earliest known stem tetrapod Tungsenia and the elpistostegid (fish-tetra...

We describe late Miocene tetrapod footprints (tracks) from the Trachilos locality in western Crete (Greece), which show hominin-like characteristics. They occur in an emergent horizon within an otherwise marginal marine succession of Messinian age (latest Miocene), dated to approximately 5.7 Ma (million years), just prior to the Messinian Salinity...

In order to understand the ecological dimension of the fish-tetrapod transition, which occurred within the tetrapod stem group during the Devonian Period (419-359 million years ago), we need life-history data from transitional forms. Only recently have serious attempts begun to utilize limb-bone histology as a source of such data. Here we present h...

Recently, attempts have been made to utilize limb-bone histology as a source of data for understanding the ecological dimension of the fish-tetrapod transition. Here we present histological life history data from a humerus (ANSP 21483) of Hyneria lindae, a fish member of the tetrapod stem group from the Late Devonian Catskill Formation (Pennsylvani...

Phylogenetic analysis of early tetrapod evolution has resulted in a consensus across diverse data sets in which the tetrapod stem group is a relatively homogenous collection of medium- to large-sized animals showing a progressive loss of 'fish' characters as they become increasingly terrestrial, whereas the crown group demonstrates marked morpholog...

Coprolites (fossil faeces) reveal clues to ancient trophic relations, and contain inclusions representing organisms that are rarely preserved elsewhere. However, much information is lost by classical techniques of investigation, which cannot find and image the inclusions in an adequate manner. We demonstrate that propagation phase-contrast synchrot...

The numerous cushion-shaped tooth-bearing plates attributed to the stem group osteichthyan Lophosteus superbus, which are argued here to represent an early form of the osteichthyan inner dental arcade, display a previously unknown and presumably primitive mode of tooth shedding by basal hard tissue resorption. They carry regularly spaced, recumbent...

The taxonomy of Early Devonian placoderm material from the Lochkovian and Pragian of the Prague basin, previously attributed to the genera Radotina and Holopetalichthys, is revised. The Pragian species Radotina tesselata Gross 1958 shares detailed similarities with the holotype of the Lochkovian Radotina kosorensis Gross 1950, which is also the hol...

We describe the first known occurrence of a Devonian coelacanth specimen from the lower Famennian of the Holy Cross Mountains, Poland, with a conodont element preserved in its digestive tract. A small spiral and phosphatic coprolite (fossil excrement) containing numerous conodont elements and other unrecognized remains was also found in the same de...

Placoderms are considered as the first jawed vertebrates and constitute a paraphyletic group in the stem-gnathostome grade. The acanthothoracid placoderms are among the phylogenetically most basal and morphologically primitive gnathostomes, but their neurocranial anatomy is poorly understood. Here we present a near-complete three-dimensional skull...

Anatomical abbreviations used in the figures. (XLS)

Romundina stellina, MNHN.F.CPW1, filling of the internal structures and rotation around the specimen. (0400–0750 (Nouveau).avi: Maya Autodesk; 25 MB. (AVI)

Romundina stellina, MNHN.F.CPW1, rotation of the endocranial cavity. olaf_002: Mimics; 5,28 MB. (AVI)

Romundina stellina, MNHN.F.CPW1, rotation of the right inner ear. olaf_006: Mimics; 8,61 MB. (AVI)

Remark concerning the myodomes and the extrinsic muscles in the orbit. (DOC)

Romundina stellina, MNHN.F.CPW1, complete run-through (contrasted) (complete_runthrough_contrasted_avi.avi; 19,8 MB). ImageJ. (AVI)