Weak trophic interactions and the balance of nature (original) (raw)

Recent discoveries of previously unknown fossil forms have dramatically transformed understanding of many aspects of the fish-tetrapod transition. Newer paleobiological approaches have also contributed to changed views of which animals were involved and when, where, and how the transition occurred. This review summarizes major advances made and reevaluates alternative interpretations of important parts of the evidence.We begin with general issues and concepts, including limitations of the Paleozoic fossil record.We summarize important features of paleoclimates, paleo- environments, paleobiogeography, and taphonomy. We then review the history of Devonian tetrapods and their closest stem group ancestors within the sarcopterygian fishes. It is now widely accepted that the first tetrapods arose from advanced tetrapodomorph stock (the elpistostegalids) in the Late Devonian, probably in Euramerica. However, truly terrestrial forms did not emerge until much later, in geographically far-flung regions, in the Lower Carboniferous. The complete transition occurred over about 25 million years; definitive emergences onto land took place during the most recent 5 million years. The sequence of character acquisition during the transition can be seen as a five-step process involving: (1) higher osteichthyan (tetrapodomorph) diversification in the Middle Devonian (beginning about 380 million years ago [mya]), (2) the emergence of “prototetrapods” (e.g., Elginerpeton) in the Frasnian stage (about 372 mya), (3) the appearance of aquatic tetrapods (e.g., Acanthostega) sometime in the early to mid- Famennian (about 360 mya), (4) the appearance of “eutetrapods” (e.g., Tulerpeton) at the very end of the Devonian period (about 358 mya), and (5) the first truly terrestrial tetrapods (e.g., Pederpes) in the Lower Carboniferous (about 340 mya). We discuss each of these steps with respect to inferred functional utility of acquired character sets. Dissociated heterochrony is seen as the most likely process for the evolutionarily rapid morphological transformations required. Developmental biological processes, including paedomorphosis, played important roles. We conclude with a discussion of phylogenetic interpretations of the evidence.

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 fossil evidence. The trace fossils demonstrate that tetrapods were capable of performing subaerial lateral sequence walks before the end of the Middle Devonian. The derived morphological characters of elpistostegids and Devonian tetrapods are related to substrate locomotion, weight support and aerial vision, and thus to terrestrial competence, but the retention of lateral-line canals, gills and fin rays shows that they remained closely tied to the water. Elpistostegids and tetrapods both evolved no later than the beginning of the Middle Devonian. The earliest tetrapod records come from inland river basins, sabkha plains and ephemeral coastal lakes that preserve few, if any, body fossils; contemporary elpistostegids occur in deltas and the lower reaches of permanent rivers where body fossils are preserved. During the Frasnian, elpistostegids disappear and these riverine-deltaic environments are colonised by tetrapods. This replacement has, in the past, been misinterpreted as the origin of tetrapods.

All living tetrapods have a one-to-two branching pattern in the embryonic proximal limb skeleton, with a single element at the base of the limb (the humerus or femur) that articulates distally with two parallel radials (the ulna and radius or the tibia and fibula). This pattern is also seen in the fossilized remains of stem-tetrapods, including the fishlike members of the group, in which despite the absence of digits, the proximal parts of the fin skeleton clearly resemble those of later tetrapods. However, little is known about the developmental mechanisms that establish and canalize this highly conserved pattern. We describe the well-preserved pelvic fin skeleton of Rhizodus hibberti, a Carboniferous sarcopterygian (lobe-finned) fish, and member of the tetrapod stem group. In this specimen, three parallel radials, each robust with a distinct morphology , articulate with the femur. We review this unexpected morphology in a phylogenetic and developmental context. It implies that the developmental patterning mechanisms seen in living tetrapods, now highly constrained, evolved from mechanisms flexible enough to accommodate variation in the zeugopod (even between pectoral and pelvic fins), while also allowing each element to have a unique morphology. zeugopod | pelvis | limb patterning | tetrapodomorph | rhizodontid

Study of a growth series of twenty-seven specimens from the Upper Devonian of Escuminac Bay, Que´bec documents a complex pattern of vertebral development in the osteolepiform fish Eusthenopteron foordi. Ossification begins with elements associated with the caudal, anal, and second dorsal fins. Development of the haemal arches, caudal radials, and caudal neural arches continues anteriorly and posteriorly from near the level of the anterior margin of the caudal fin. Trunk neural arches ossify later than the caudal neural arches and as a separate sequence. Trunk intercentra most likely begin ossification posteriorly and continue forward after the ossification of haemal arches is complete. Comparisons of many different patterns of vertebral development within the modern actinopterygians demonstrates that the sequence of development in Eusthenopteron foordi is unique. The diverse patterns of vertebral development observed in fossil and modern fish presumably result from an interplay between the inherent anterior to posterior sequence of development controlled by the Hox genes, and varying selective forces imposed by the physical and biological environment in which the fish develop. Initiation of vertebral development in the caudal region of Eusthenopteron foordi can be attributed to selection for early function of the tail in propulsion. In contrast, vertebral development in Carboniferous amphibians typically proceeds from anterior to posterior. This may reflect development in the still water of ponds and lakes in contrast with the coastal environment inhabited by the hatchlings of Eusthenopteron foordi. The sequences of vertebral development seen in Carboniferous labyrinthodonts and lepospondyls are divergently derived from that observed in Eusthenopteron foordi.

The relationship of the three living groups of sarcopterygians or lobe-finned fish (tetrapods, lungfish and coelacanths) has been a matter of debate1–5. Although opinions still differ, most recent phylogenies suggest that tetrapods are more closely related to lungfish than to coelacanths6–10. However, no previously known fossil taxon exhibits a concrete character combination approximating the condition expected in the last common ancestor of tetrapods and lungfish—and it is still poorly understood how early sarcopterygians diverged into the tetrapod lineage (Tetrapodomorpha)7 and the lungfish lineage (Dipnomorpha)7. Here we describe a fossil sarcopterygian fish, Styloichthys changae gen. et sp. nov., that possesses an eyestalk and which exhibits the character combination expected in a stem group close to the last common ancestor of tetrapods and lungfish. Styloichthys from the Lower Devonian of China bridges the morphological gap between stem-group sarcopterygians (Psarolepis and Achoania)10 and basal tetrapodomorphs/basal dipnomorphs. It provides information that will help in the study of the relationship of early sarcopterygians, and which will also help to resolve the tetrapod–lungfish divergence into a documented sequence of character acquisition.

About half of all vertebrate species today are ray-finned fishes (Actinopterygii), and nearly all of them belong to the Neopterygii (modern ray-fins). The oldest unequivocal neopterygian fossils are known from the Early Triassic. They appear during a time when global fish faunas consisted of mostly cosmopolitan taxa, and contemporary bony fishes belonged mainly to non-neopterygian (“paleopterygian”) lineages. In the Middle Triassic (Pelsonian substage and later), less than 10 myrs (million years) after the Permian-Triassic boundary mass extinction event (PTBME), neopterygians were already species-rich and trophically diverse, and bony fish faunas were more regionally differentiated compared to the Early Triassic. Still little is known about the early evolution of neopterygians leading up to this first diversity peak. A major factor limiting our understanding of this “Triassic revolution” is an interval marked by a very poor fossil record, overlapping with the Spathian (late Olenekia...