Paleobiology

The Siluro-Devonian primary radiation of land biotas is the terrestrial equivalent of the much-debated Cambrian "explosion" of marine faunas. Both show the hallmarks of novelty radiations (phenotypic diversity increases much more rapidly than species diversity across an ecologically undersaturated and thus low-competition landscape), and both ended with the formation of evolutionary and ecological frameworks analogous to those of modern ecosystems. Profound improvements in understanding early land plant evolution reflect recent liberations from several research constraints: Cladistic techniques plus DNA sequence data from extant relatives have prompted revolutionary reinterpretations of land plant phylogeny, and thus of systematics and character-state acquisition patterns. Biomechanical and physiological experimental techniques developed for extant 263 0066-4162/98/1120-0263$08.00 and complicated by the recent knowledge that global landmass positions, currents, climates, and atmospheric compositions have been profoundly variable (and thus nonuniformitarian) through the Phanerozoic. Combining phylogenetic and paleoecological data offers potential insights into the identity and function of key innovations, though current evidence suggests the importance of accumulating within lineages a critical mass of phenotypic character. Challenges to further progress include the lack of sequence data and paucity of phenotypic features among the early land plant clades, and a fossil record still inadequate to date accurately certain crucial evolutionary and ecological events.

s Abstract Late Carboniferous and Early Permian strata record the transition from a cold interval in Earth history, characterized by the repeated periods of glaciation and deglaciation of the southern pole, to a warm-climate interval. Consequently, this time period is the best available analogue to the Recent in which to study patterns of vegetational response, both to glacial-interglacial oscillation and to the appearance of warm climate. Carboniferous wetland ecosystems were dominated by spore-producing plants and early gymnospermous seed plants. Global climate changes, largely drying, forced vegetational changes, resulting in a change to a seed plant-dominated world, beginning first at high latitudes during the Carboniferous, reaching the tropics near the Permo-Carboniferous boundary. For most of this time plant assemblages were very conservative in their composition. Change in the dominant vegetation was generally a rapid process, which suggests that environmental thresholds were crossed, and involved little mixing of elements from the wet and dry floras.

A change to global greenhouse conditions following deglaciation occurred during the late Paleozoic. The deep-past data set preserved in the stratigraphic record can serve as a model system to understand vegetational responses during this kind of climatic change, especially in the tropics. No other time in Earth history so mimics the late Cenozoic or provides the long-term data set from which generalizations can be deduced. Two long-term glacial cycles have been identified in Permian-Carboniferous time. The waxing and waning of glaciers during the height of either ice age resulted mainly in spatial displacement of vegetation, and also in minor variations in tropical climate. Brief intervals of rapid deglaciation at the end of the Middle Pennsylvanian (Westphalian) and mid-Early Permian (Sakmarian) were accompanied by major changes in plant assemblages, including extinctions, changes in the • umM GSA TODAY

■ Abstract Studies of plant and animal assemblages from both the terrestrial and the marine fossil records reveal persistence for extensive periods of geological time, sometimes millions of years. Persistence does not require lack of change or the absence of variation from one occurrence of the assemblage to the next in geological time. It does, however, imply that assemblage composition is bounded and that variation occurs within those bounds. The principal cause for these patterns appears to be species-, and perhaps clade-level, environmental fidelity that results in long-term tracking of physical conditions. Other factors that influence persistent recurrence of assemblages are historical, biogeographic effects, the "law of large numbers," niche differentiation, and biotic interactions. Much research needs to be done in this area, and greater uniformity is needed in the approaches to studying the problem. However, great potential also exists for enhanced interaction between paleoecology and neoecology in understanding spatiotemporal complexity of ecological dynamics.

Further developing a controversial neoGoldschmidtian paradigm that we first published in 1994, we here narrowly define saltational evolution as a genetic modification that is expressed as a profound phenotypic change across a single generation and results in a potentially independent evolutionary lineage termed a prospecies (the 'hopeful monster' of Richard Goldschmidt). Of several saltational and parasaltational mechanisms previously discussed by us, the most directly relevant to evolutionary-developmental genetics is dichotomous saltation, which is driven by mutation within a single ancestral Uneage. It can result not only in instantaneous speciation but also in the simultaneous origin of a profound phenotypic novelty more likely to be treated as a new supraspecific taxon. Saltational events form unusually long branches on morphological phylogenies, which follow a punctuated equilibrium pattern, but at the time of origin they typically form zero length branches on the contrastingly gradualistic molecular phylogenies. Our chosen casestudies of heterotopy (including homeosis) and heterochrony in fossil seed-ferns and extant orchids indicate that vast numbers of prospecies are continuously generated by mutation of key developmental genes that control morphogenesis. First principles suggest that, although higher plant mutants are more likely to become established than higher animals, the fitness of even plant prospecies is in at least most cases too low to survive competition-mediated selection. The establishment of prospecies is most Hkely under temporary release from selection in environments of low biotic competition for resources, followed by honing to competitive fitness by gradual reintroduction to neoDarwinian selection. Unfike neoDarwinism alone, this two-phase evolutionary paradigm is consistent with the recent results of (a) whole-genome sequencing, which have revealed a surprisingly small total number of genes per model species, and (b) of Quantitative Trait Locus analyses, which indicate that major phenotypic features are determined by one or two homeotic genes of major phenotypic effect and only a handful of genes of lesser effect. Evolutionarydevelopmental genetics has already proved beyond reasonable doubt the credibility of the initial 'generative phase' of the neoGoldschmidtian hypothesis (though further investigation is needed of the effects of canalisation and epigenesis following both gain and loss of function). Unfortunately, far fewer data are currently available to test the subsequent 'estabhshment phase'; this deficiency places a premium on monitoring the ecological progress of many prospecies, in both artificial and natural

A major extinction of terrestrial plants occurred at the end of the Westphalian (Middle Pennsylvanian) in the lowland tropics of North America. Approximately 67% of the species in peat-forming mires, and at least half the species in clastic wetlands were eliminated by changing climatic conditions, probably protracted moisture deficits or exaggeration of seasonal dryness. Independent studies suggest that the end of the Westphalian was marked by deglaciation in Gondwana and increase in global temperature.

Early Permian (late Leonardian Series) plant assemblages from King, Knox, and Stonewall Counties of North-Central Texas are dominated by seed plants, some apparently congeneric with taxa heretofore known only from the Late Permian or the Mesozoic. Conifers are the dominant elements, including one or more species of Ullmannia, Pseudovoltzia liebeana, both known from the Late Permian Zechstein flora of Germany and England, Podozamites sp., characteristic of the Mesozoic, and Walchia sp., abundant in Early Permian floras. Locally common are Taeniopteris cf. eckardtii, a Zechstein species, an unidentified plant represented by pinnulelike laminae with fine parallel veins, similar to pinnules of some Mesozoic cycads, and calamite stems. Rarely encountered are leaf fragments of the Paleozoic ginkgophyte Dicranophyllum, flabellate ginkgophyte leaves, leaves with a broad midvein and narrow, fimbriate lamina, and Wattia, typical of the Early Permian. Associated with these foliar remains are ovulate reproductive structures including the presumed cycad megasporophyll Dioonitocarpidium, known only from the Mesozoic, a voltzialean cone scale similar to Swedenborgia, and a variety of seeds, some remarkably similar to Agathis, of Cretaceous age. The assemblage includes only rare scraps of foliage and seeds possibly attributable to the pteridophyllous elements (gigantopterids, callipterids, and ferns) that dominate the Permian. The fossil plants occur in multistorey, fining-upwards, tidal-channel deposits that also include pelecypods and fragmentary palaeoniscoid fish. The occurrence of derived lineages in xeric habitats during the Early Permian indicates that some supposed Mesozoic groups actually preceded and survived the end-Permian extinction, reappearing in basinal lowlands during the mid-Mesozoic.

Pennsylvanian-age wetland plant communities and landscape gradients exhibit persistent species composition and ecomorphic structure. Such patterns are attributable in large part to strong phylogenetic partitioning of ecological resource space at the level of higher taxa. Each of four major class-rank clades in tropical wetlands was centered in a physically distinct part of the lowland ecosystem. Once established in these physical settings clades tended to resist displacement until removed by physically driven extinction. Lycopsid trees were the principal dominants of the wettest habitats and had subpartitioned these environments along generic lines. Seed plants were the dominants on well to poorly drained clastic substrates, a diverse set of habitats reflected in high diversity of species and architectures. Sphenopsids were most abundant in aggradational environments subject to high levels of physical stress. Ferns initially were interstitial opportunists and colonizers of disturbed areas in a variety of environments. These ecological patterns were estabUshed in concert with the architectural radiation of the vascular plants, which occurred during the Devonian-Mississippian transition and estabhshed the major classes. Within the wetlands, the replacement of species by close relatives drawn from the same families or genera contributed significantly to persistence of communities and landscape gradients. Replacement was more likely to occur from within a clade where similar ecologies already existed, than from across major clades between which the basic ecologies were different. Possibly interacting with the phylogenetic factors were community and landscape level multispecies effects that may have placed limits on species replacement patterns. The existence of such emergent properties of multispecies assemblages is suggested by a breakdown of the system beginning with major, climatically induced extinctions at the Middle-Late Pennsylvanian transition. Following extinctions of the major Middle Pennsylvanian trees, opportunistic ferns gave rise to dominants in many parts of the wetlands, perhaps due to loss of some aspects of system self-regulation. As climatic drying continued seed plants began their largely passive rise to dominance in most kinds of habitats.

The Late Carboniferous glaciation reached its peak during the latter part of the Westphahan (late Desmoinesian). In the tropical Illinois basin this was the time of deposition of the Carbondale and lower Modesto Formations, characterized by the cyclic repetition of major coal deposits, black shales, limestones, gray shales and sandstones. These lithological changes evidence repeated major fluctuations in climate and sealevel. Fossil floras from the tropical ever-wet to seasonally wet terrestrial hthofacies, coals and shales above coals, remain compositionally similar throughout the 1-1.5 m.y. time interval.