The structure and phylogenetic significance of the conifer Pseudohirmerella delawarensis nov. comb. from the Upper Triassic of North America

The conifer genus Pseudohirmerella Arndt is based on ovulate cone scales of the type species, Pseudohirmerella platysperma (Mägdefrau) Arndt, from the Upper Triassic of Germany, which bear five distal lobes and two paired structures represented by bulges on the adaxial surface. The paired structures were first described as seeds, but have recently been re-interpreted as arils by Arndt. Similar scales from the Passaic Formation at Milford, NJ, USA were described as Glyptolepis delawarensis Bock, and the new combination Pseudohirmerella delawarensis is proposed. A relatively complete concept of Pseudohirmerella is presented, which includes ovulate cones, possible associated pollen cones, associated shoots with Pagiophyllum-Brachyphyllum morphotype leaves and associated, anatomically preserved wood. Based on the ovulate cone morphology and the presence of organic matter lining the concavities on the scale rather than the corresponding bulges on the counterparts, it is likely that the supposed seeds or arils of Pseudohirmerella are actually casts of empty, seed-bearing depressions. Cheirolepidiaceous affinities are likely based on ovulate cone scale morphology, persistent pollen cones, foliage type and details of wood anatomy. The derived ovulate cone scale morphology of Pseudohirmerella indicates a substantial, but mostly undocumented, Triassic diversification of the Cheirolepidiaceae.     

Angiosperm growth habit,dispersal and diversification reconsidered

Summary Previous studies have sought to elucidate the relationship between dispersal mode (biotic versus abiotic) and the taxonomic diversification of angiosperm families, but with ambiguous results. In this study, we propose the hypothesis that the combination of (1) the large seed size required of plants germinating in closed, light-poor environments and (2) the necessity to move disseminules away from the maternal plant in order to avoid intraspecific competition, predation and pathogens should favour biotically-dispersed relative to abiotically-dispersed woody arborescent angiosperms, resulting in higher diversification of the former. In this paper, we seek patterns of diversification that support this hypothesis. We examine the association between dispersal mode, growth habit and taxonomic richness of monocotyledon and dicotyledon families using (1) contingency table analyses to detect the effect of dispersal mode on the relative abundances and diversification of woody versus herbaceous taxa and (2) non-parametric analyses of variance to detect the statistical effect of dispersal mode on taxonomic diversification (mean number of species per genus, genera per family and species per family) in monocot and dicot families dominated by biotic or abiotic dispersal. We found a clear statistical effect of dispersal mode on diversification. Among families of woody dicots, dispersal by vertebrates is associated with significantly higher levels of species per genus, genera per family and species per family than is abiotic dispersal. The same pattern is observed among woody monocots, but is not significant at the 0.05 level. Among families of herbaceous monocots and dicots, the situation is reversed, with abiotically-dispersed families exhibiting higher levels of diversification than vertebrate-dispersed families. When woody and herbaceous families are pooled, there is no association between dispersal mode and diversification. These data coincide with evidence from the fossil record to suggest vertebrate dispersal has positively contributed to the diversification of woody angiosperms. We suggest that vertebrate dispersal may have promoted the diversity of extant taxa by reducing the probability of extinction over evolutionary time, rather than by elevating speciation rates. Our results suggest vertebrate dispersal has contributed to, but does not explainin toto, the diversity of living angiosperms.     

果实类型和种子传播的进化:系统发育和生态学简论

Success of flowering plants is greatly dependent on effective seed dispersal. Specific fruit types aid different mechanisms of seed dispersal. However, little is known about what evolutionary forces have driven the diversification of fruit types and whether there were phylogenetic constraints on fruit evolution among angiosperm lineages. To address these questions, we first surveyed the orders and families of angiosperms for fruit types and found no clear association between fruit types and major angiosperm lineages, suggesting there was little phylogenetic constraint on fruit evolution at this level. We then surveyed fruit types found in two contrasting habitats an open habitat including the Indian desert and North American plains and prairies, and a closed forest habitat of Australian tropical forest. The majority of genera in the survey of tropical forests in Australia were fleshy fruit trees, whereas the majority of genera in the survey of prairies and plains in central North America were herbs with capsules and achenes. Both capsules and achenes are frequently dispersed by wind in the open, arid habitat, whereas fleshy fruits are generally dispersed by animals. Since desert and plains tend to provide continuous wind to aid dispersal and there are more abundant mammal and bird dispersers in the closed forest, this survey suggests that fruit evolution was driven at least in part by dispersal agents abundant in particular habitats.     

Recruitment trade-offs and the evolution of dispersal mechanisms in plants

In this study we place seed size vs. seed number trade-offs in the context of plant dispersal ability. The objective was to suggest explanations for the evolution of different seed dispersal mechanisms, in particular fleshy fruits, wind dispersal and the maintenance of unassisted dispersal. We suggest that selection for improved dispersal may act either by increasing the intercept of a dispersal curve (log seed number vs. distance) or by flattening the slope of the curve. 'Improved dispersal' is defined as a marginal increase in the number of recruits sited at some (arbitrary) distance away from the parent plant. Increasing the intercept of the dispersal curve, i.e. producing more seeds, is associated with a reduction in seed size, which in turn affects the recruitment ability, provided that this ability is related to seed size. If recruitment is related to seed size there will be a recruitment cost of evolving increased seed production. On the other hand, a flattening of the slope by evolving dispersal attributes is likely to be associated with a fecundity cost. An exception is wind dispersal where smaller (and hence more numerous) seeds may lead to more efficient dispersal. We derive two main predictions: If recruitment is strongly related to seed size, selection for improved dispersal acts on the slope of the dispersal curve, i.e. by favouring evolution of dispersal attributes on seeds or fruits. If, on the other hand, recruitment is only weakly related to seed size (or not related, or negatively related), selection for improved dispersal favours increased seed production. Despite its simplicity, the model suggests explanations for (i) why so many plant species lack special seed dispersal attributes, (ii) differences in dispersal spectra among plant communities, and (iii) adaptive radiation in seed size and dispersal attributes during angiosperm evolution. This revised version was published online in July 2006 with corrections to the Cover Date.     

Ichnology of an ephemeral lacustrine/alluvial plain system: Jurassic East Berlin Formation,Hartford Basin,USA

A trace fossil assemblage from the Lower Jurassic East Berlin Formation of the Newark Supergroup, Hartford Basin, New England, USA, includes: Scoyenia gracilis, Skolithos ichnosp., Palaeophycus striatus, Planolites montanus, Fuersichnus ichnosp., fusiform burrows, pelleted material, an escape structure, and large burrows.

This assemblage is assigned to the Scoyenia ichnofacies. Specific lebensspuren are not limited to specific lithofacies; instead, their initial distribution seems to have been influenced principally by water availability within an ephemeral lacustrine/alluvial plain system. Other factors in distribution may have included amounts of organic matter, patterns of sedimentation, sediment grain size, biotic factors (settling from invertebrate drift, competition), and additional abiotic factors (wind deflation, waves, currents, desiccation, soft‐sediment deformation, evaporite formation, pedoturbation).

Extreme environmental conditions within the original depositional setting strongly influenced the availability of water which, in turn, strongly influenced the paleoecology of burrowing invertebrates in this nonmarine system.     

Seed release by invasive thistles: the impact of plant and environmental factors

Dispersal is a key process in biological studies of spatial dynamics, but the initiation of dispersal has often been neglected, despite strong indications that differential timing of dispersal can significantly affect dispersal distances. To investigate which plant and environmental factors determine the release of plumed seeds by the invasive thistles Carduus acanthoides and Carduus nutans, we exposed 192 flower heads of each species to increasing wind speeds in a full-factorial wind tunnel experiment with four air flow turbulence, three flower head wetness and two flower head temperature levels. The number of seed releases was highest under dry and turbulent conditions and from heads that had already lost a considerable number of seeds, but was not affected by flower head size, head angle or temperature. Inspection of the trials on video showed that higher wind speeds were needed to meet the seed release threshold in laminar flows and for C. acanthoides heads that had been wet for a longer time. Species differences were minimal, although seed release was more sensitive to lower levels of turbulence in the larger-headed and more open C. nutans heads. Knowledge of seed release biases towards weather conditions favourable for long-distance dispersal improves our understanding of the spread of invaders and allows managers to increase the efficiency of their containment strategies by applying them at crucial times.     

Primate origins and the evolution of angiosperms

Traditionally, the morphological traits of primates were assumed to be adaptations to an arboreal way of life. However, Cartmill [1972] pointed out that a number of morphological traits characteristic of primates are not found in many other arboreal mammals. He contends that orbital convergence and grasping extremities indicate that the initial divergence of primates involved visual predation on insects in the lower canopy and undergrowth of the tropical forest. However, recent research on nocturnal primates does not support the visually-oriented predation theory. Although insects were most likely important components of the diets of the earliest euprimates, it is argued here that visual predation was not the major impetus for the evolution of the adaptive traits of primates. Recent paleobotanical research has yielded evidence that a major evolutionary event occurred during the Eocene, involving the angiosperms and their dispersal agents. As a result of long-term diffuse coevolutionary interactions with flowering plants, modern primates, bats, and plant-feeding birds all first arose around the Paleocene-Eocene boundary and became the major seed dispersers of modern tropical flora during the Eocene. Thus, it is suggested here that the multitude of resources available on the terminal branches of the newly evolved angiosperm, rain forest trees led to the morphological adaptations of primates of modern aspect.     

How important is long‐distance seed dispersal for the regional survival of plant species?

Long‐distance seed dispersal is generally assumed to be important for the regional survival of plant species. In this study, we quantified the importance of long‐distance seed dispersal for regional survival of plant species using wind dispersal as an example. We did this using a new approach, by first relating plant species’ dispersal traits to seed dispersal kernels and then relating the kernels to regional survival of the species. We used a recently developed and tested mechanistic seed dispersal model to calculate dispersal kernels from dispersal traits. We used data on 190 plant species and calculated their regional survival in two ways, using species distribution data from 36,800 1 km²‐grid cells and 10,754 small plots covering the Netherlands during the largest part of the 20th century. We carried out correlation and stepwise multiple regression analyses to quantify the importance of long‐distance dispersal, expressed as the 99‐percentile dispersal distance of the dispersal kernels, relative to the importance of median‐distance dispersal and other plant traits that are likely to contribute to the explanation of regional survival: plant longevity (annual, biennial, perennial), seed longevity, and plant nutrient requirement. Results show that long‐distance dispersal plays a role in determining regional survival, and is more important than median‐distance dispersal and plant longevity. However, long‐distance dispersal by wind explains only 1–3% of the variation in regional survival between species and is equally important as seed longevity and much less important than nutrient requirement. In changing landscapes such as in the Netherlands, where large‐scale eutrophication and habitat destruction took place in the 20th century, plant traits indicating ability to grow under the changed, increasingly nutrient‐rich conditions turn out to be much more important for regional survival than seed dispersal.     

Lower Jurassic Arthropod Resting Trace from the Hartford Basin of Massachusetts,USA

Cheliceratichnus lockleyi ichnogen. nov. et ichnosp. nov. is a new ichnotaxon of arthropod resting trace (cubichnium) from the Lower Jurassic (Hettangian) East Berlin Formation in Holyoke, Massachusetts, USA. The trace fossil is preserved as showing many of the external anatomical features of the exoskeleton, which resemble those of some chelicerates, notably sun spiders (Solifugae). The resting trace is directly associated with a trackway of the ichnospecies Acanthichnus cursorius Hitchcock. This is the first described fossil resting trace of a solifugan-like arthropod, and the first direct evidence of a trackmaker of A. cursorius.     

HISTORY OF ICHNOLOGY—Ichnological Eccentrics: The Curious Case of Dr. Joseph Barratt of Middletown,Connecticut

Joseph Barratt was a British-born and educated physician who settled in the United States in 1819. He had a great interest in natural history, collecting both plants and insects and studying geology, mycology, ornithology, chemistry, meteorology, Native Americans, and local history. He was apparently a man of great energy and ambition but one who could not focus to see a project to completion. Barratt was active in the early history of the discovery of vertebrate footprints in the Newark Supergroup in the eastern United States but latter developed some very strange theories regarding the age and significance of these deposits. In his latter years, Barratt's mental state deteriorated and he became even more obsessed with trying to publicize his outlandish theories. Dr. Joseph Barratt is remembered in ichnology for basically two things; he sold Edward Hitchcock a superb specimen of vertebrate tracks; and he may have the most elaborate ichnological tombstone ever constructed.     

Which factors have stronger explanatory power for primary wind dispersal distance of winged diaspores: the case of Zygophyllum xanthoxylon (Zygophyllaceae)?

Aims How seed dispersal distance is related to various factors is a major challenge for seed ecologists. However, there are different answers as to which factor is most important in determining wind dispersal distance. This study is to quantitatively describe the relationship between various factors and primary wind dispersal distance of winged diaspores.Methods The dispersal distances of five morphologies of winged diaspores in Zygophyllum xanthoxylum (Zygophyllaceae) were measured under controlled conditions in a wind tunnel. The explanatory power of environmental factor (i.e. wind speed), plant trait (i.e. release height) and diaspore attributes (i.e. wing loading (the ratio of diaspore mass to projected area), settlement-velocity, shape index (the variance of diaspore length, width and thickness)) to the variation in dispersal distance was assessed by releasing diaspores at varying wind speeds and release heights.Important findings Wind speed and seed release height were the strongest explanatory factors to dispersal distance, contributing 41.1% and 24.8% (P < 0.01) to total variation in dispersal distance, respectively. Wind speed accounted more for relatively light disc-shaped seeds than for relatively heavy spherical seeds. Wing loading, shape index and settlement-velocity explained 9.0% (P < 0.01), 1.4% (P < 0.01) and 0.9% (not significant) of the variation in dispersal distance, respectively. From disc-shaped to four-winged diaspores, relative contributions of wing loading and shape index decreased but contribution of settlement-velocity increased. The relative contributions of various factors to wind seed dispersal distance may change with the change in seed morphology.     

A new apparatus to measure the rate of fall of seeds

1. Seed dispersal is widely recognized to be of crucial importance in the ecology of plant communities, yet characterization of dispersal capacity rarely goes beyond two categories: 'good' and 'poor' dispersers, defined on the basis of dispersule morphology.
2. We describe a new apparatus which determines the rate of fall of seeds in air by detecting their passage through two fans of laser light.
3. This apparatus is able to measure rate of fall in smaller seeds than previously published methods and reveals that there is no justification for the arbitrary division of species into two wind dispersal categories.     

Comparative evolution of flower and fruit morphology

Angiosperm diversification has resulted in a vast array of plant morphologies. Only recently has it been appreciated that diversification might have proceeded quite differently for the two key diagnostic structures of this clade, flowers and fruits. These structures are hypothesized to have experienced different selective pressures via their interactions with animals in dispersal mutualisms, resulting in a greater amount of morphological diversification in animal-pollinated flowers than in animal-dispersed fruits. I tested this idea using size and colour traits for the flowers and fruits of 472 species occurring in three floras (St John, Hawaii and the Great Plains). Phylogenetically controlled analyses of nearest-neighbour distances in multidimensional trait space matched the predicted pattern: in each of the three floras, flowers were more divergent from one another than were fruits. In addition, the spacing of species clusters differed for flowers versus fruits in the flora of St John, with clusters in flower space more divergent than those in fruit space. The results are consistent with the idea that a major driver of angiosperm diversification has been stronger selection for divergent floral morphology than for divergent fruit morphology, although genetic, physiological and ecological constraints may also play a role.