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The evolution of different spore size classes, or heterospory, is a fundamental reproductive innovation in land plants. The appearance of heterospory is particularly notable during the Devonian, when most known origins of the trait occur. Here we provide a perspective on the evolution of heterospory during this time interval, particularly from the late Early Devonian through the Middle Devonian (Emsian to Givetian Stages; 408–383 Ma), which shows an unusually high concentration of heterospory origins. We use theoretical considerations and compilations of fossil and extant spore sizes to suggest that the basic features of most heterosporous lineages, large spores and gametophytes that mature within the spore wall, are difficult to evolve in combination, because large spores disperse poorly but small spores cannot support a functional gametophyte developing within their walls; evolving spores between 100 and 200 microns in diameter appears to represent a particularly important barrier for the evolution of heterospory. We then discuss why this barrier may have been lower in the Devonian, noting that the appearance and spread of heterospory is coincident with the emergence of peat-accumulating wetland habitats. We suggest that more widespread wetland habitats would have generally lowered barriers to the evolution of heterospory by reducing dispersal limitation in larger spores. Ultimately, we suggest that the initial evolution of heterospory may be explained by major changes in sedimentology, thought to have been driven by plant evolution itself, that increased the diversity of terrestrial depositional environments and led to a greater number of habitats where large spores could be successful.

PREMISE Seed cones of extant Pinaceae exhibit two mechanisms of seed release. In "flexers" the cone scales remain attached to the central axis, while flexing and separating from each other to release the seeds. In "shedders" scales are shed from the axis, with the seeds either remaining attached to the scale or becoming detached. The early fossil history of Pinaceae from the Jurassic to Early Cretaceous is dominated by flexing seed cones, while the systematic information provided by shedding fossil cones has been overlooked and rarely integrated with data based on compression and permineralized specimens. We describe the earliest and best-documented evidence of a "shedder" seed cone from the Aptian-Albian of Mongolia. METHODS Lignite samples from Tevshiin Govi locality were disaggregated in water, washed, and dried in air. Fossils were compared to material of extant Pinaceae using LM and CT scans. RESULTS Lepidocasus mellonae gen. et sp. nov. is characterized by a seed cone that disarticulated at maturity and shed obovate bract-scale complexes that have a distinctive ribbed surface and an abaxial surface covered with abundant trichomes. The ovuliferous scale has ca. 30-40 resin canals, but only scarce xylem near the attachment to the cone axis. Resin vesicles are present in the seed integument. Phylogenetic analysis places Lepidocasus as sister to extant Cedrus within the abietoid grade. CONCLUSIONS The exquisite preservation of the trichomes in L. mellonae raises questions about their potential ecological function in the cones of fossil and living Pinaceae. Lepidocasus mellonae also shows that a shedding dispersal syndrome, a feature that has often been overlooked, evolved early in the history of Pinaceae during the Early Cretaceous.

Premise of research. Heterospory, or the production of different spore size classes, is a key aspect of advanced vascular plant reproductive strategies. Heterospory is widespread across plant linea...

Premise of research. Serotiny is a specialized dispersal mechanism in which seeds are retained on the parent plant and released following a disturbance event. This strategy has evolved repeatedly i...

New Caledonia was, until recently, considered an old continental island harbouring a rich biota with outstanding Gondwanan relicts. However, deep marine sedimentation and tectonic evidence suggest complete submergence of the island during the latest Cretaceous to the Paleocene. Molecular phylogenies provide evidence for some deeply-diverging clades that may predate the Eocene and abundant post-Oligocene colonisation events. Extinction and colonization biases, as well as survival of some groups in refuges on neighbouring paleo-islands, may have obscured biogeographic trends over long time scales. Fossil data are therefore crucial for understanding the history of the New Caledonian biota, but occurrences are sparse and have received only limited attention. Here we describe five exceptional fossil assemblages that provide important new insights into New Caledonia’s terrestrial paleobiota from three key time intervals: prior to the submersion of the island, following re-emergence, and prior to Pleistocene climatic shifts. These will be of major importance for elucidating changes in New Caledonia’s floristic composition over time.

Evolutionary pulses of complexity The evolution of land plants has been marked by an increase in the complexity of reproductive structures. Leslie et al . analyzed the temporal pattern of this increase in a study of fossil and extant land plants across the entire paleontological record. Reproductive complexity increased in two widely separated pulses corresponding to key innovations in reproductive biology: the origin of seeds in the very late Devonian and the origin of flowering plants in the mid-Cretaceous almost 250 million years later. After the origin of flowers, there was a rapid expansion in the morphological complexity of flowering plants. In contrast to many aspects of animal morphological diversity, which expanded early in evolutionary history, most complexity in plants was achieved relatively late. —AMS

Premise of research. The Devonian was a crucial period in plant history for both vegetative and reproductive evolution. We focus on reproductive structures during this time period and ask whether a...

Plant-derived diterpenoids are commonly used as conifer-specific biomarkers and for chemotaxonomic assignment or confirmation. Numerous studies have reported on the utility of diterpenoids as chemotaxonomic indicators, but few have quantitatively analyzed diterpenoid concentrations, especially with respect to conifer phylogeny. In addition, the carbon isotope values (δ13C) of diterpenoids provide a means to track conifer-specific δ13C values, which is useful for tracking water availability and the carbon cycle. To expand on previous work, we measured diterpenoid concentrations and carbon isotopes of 43 conifer species, and Ginkgo biloba, collected at the University of California Botanical Garden at Berkeley. In this collection, all extant conifer families and almost two-thirds of extant genera are present, including many subtropical and Southern Hemisphere groups that were once common and widespread components of fossil assemblages. Overall, we found diterpenoid concentrations were highly variable among species and families. Despite this variability, there is coarse family-level phylogenetic structuring with the lowest concentrations in Pinaceae. When diterpenoid concentrations are fitted to a model of trait evolution (Brownian motion), we however find that there is no phylogenetic signal. In contrast, when terpenoids are analyzed by the proportion of diterpenoid compound structure classes (abietanes, labdanes, pimaranes, tetracyclics, and totarols/sempervirol), there was significant phylogenic signal for the abietane and tetracyclic structures. Diterpenoid biosynthetic carbon isotope fractionation, as expressed between diterpenoids and leaf tissue (editerpenoid), also contained a phylogenetic signal, as well as the broad phylogenetic structuring observed in total diterpenoid concentrations. Overall, these results indicate that the Pinaceae is unique among conifer clades with respect to terpenoid structure classes, concentrations and editerpenoid. When diterpenoids are applied to taxonomic assignment of fossils, it would be useful to combine several traits (concentration, proportion, and editerpenoid). In this context, it should be possible to broadly distinguish three major conifer groups: Cupressaceae, Podocarpaceae and Pinaceae. However, based on these results, we recommend against assuming that closely related species have similar diterpenoid compositions.

BACKGROUND AND AIMS: Structures that simultaneously perform many functional roles are likely to show a variety of morphological solutions to these demands, and thus probably exhibit high morphological disparity. In contrast, specialization for a few simple functions should result in a more limited suite of morphologies. We explore this idea using lycopsid reproductive structures, which, throughout their history, have performed a limited set of functional roles compared with the reproductive structures of other plant groups such as seed plants. METHODS: We scored living and fossil lycopsid taxa for 18 discrete character measurements and several continuous traits, including sporangium size, supporting axis diameter, and strobilus length and width. We used the discrete characters to construct a multivariate morphospace for lycopsid reproductive morphology through time, and the continuous characters to test whether fossil and extant lycopsids show similar patterns of tissue allocation within reproductive structures. RESULTS: Lycopsids occupy similar areas of reproductive morphospace and show similar patterns of tissue allocation over most of their history, alternating between diffuse fertile zones with leaf-like sporophylls and compact strobili with specialized sporophylls that allow sporangia to be closely packed while also protected during their development. Growth habit also plays an important role in lycopsid reproductive evolution, broadly influencing the size and shape of reproductive structures. CONCLUSIONS: Lycopsid reproductive structures are primarily specialized for densely packaging sporangia, and are consistent with the idea that performing limited functional roles is associated with reduced morphological disparity. Morphologies similar to lycopsid strobili are also found in other groups with simple, wind-dispersed propagules, suggesting that the same processes occur across plant lineages.

The early-middle Eocene Fossil Hill flora from King George Island in the Antarctic Peninsula regions is one of the most diverse Cenozoic plant assemblages in Antarctica. It represents a rich Nothof...

BACKGROUND AND AIMS: Form and function relationships in plant reproductive structures have long fascinated biologists. Although the intricate associations between specific pollinators and reproductive morphology have been widely explored among animal-pollinated plants, the evolutionary processes underlying the diverse morphologies of wind-pollinated plants remain less well understood. Here we study how this diversity may have arisen by focusing on two conifer species in the pine family that have divergent reproductive cone morphologies at pollination. METHODS: Standard histology methods, artificial wind pollination assays and phylogenetic analyses were used in this study. KEY RESULTS: A detailed study of cone ontogeny in these species reveals that variation in the rate at which their cone scales mature means that pollination occurs at different stages in their development, and thus in association with different specific morphologies. Pollination experiments nevertheless indicate that both species effectively capture pollen. CONCLUSIONS: In wind-pollinated plants, morphological diversity may result from simple variation in development among lineages rather than selective pressures for any major differences in function or performance. This work also illustrates the broader importance of developmental context in understanding plant form and function relationships; because plant reproductive structures perform many different functions over their lifetime, subtle differences in development may dramatically alter the specific morphologies that they use to meet these demands.

Bonacorsi et al. describe a new fossil from the Early Devonian that provides the earliest clear evidence for more advanced reproductive biology in land plants. The plant produced multiple spore size classes, which is an essential innovation necessary for all advanced plant reproductive strategies, including seeds and flowers.

Biologists often study morphological evolution through form and function relationships. But biological structures can perform multiple functional roles, complicating efforts to understand the evolutionary significance of any one relationship. Plant reproductive organs perform multiple roles in a sequence, however, which provides a unique opportunity to understand how structures evolve to meet multiple functional demands. Using conifers as a study group, we discuss how a shared developmental trajectory links the performance of sequential functional roles. Variation in development among lineages can underlie morphological diversity; pollination-stage seed cones in Pinaceae conifers function similarly but show diverse forms reflecting differences in developmental rate. As cones develop further, the morphologies that they use to perform later functional roles are influenced by the specific developmental patterns used to meet earlier demands, which may ultimately limit morphological diversity. However, we also show how selective pressures relating to the final functional stage (seed dispersal) may influence cone anatomy and morphology over all previous stages, highlighting the complex linkages among form, function, and development. We end by discussing the potential relationships between functional ontogeny and morphological disparity in plant reproductive structures more broadly, suggesting that the complex functional roles associated with seed plant reproduction probably underlie the high disparity in this group.

Summary Seed size varies tremendously in plants and its evolution is influenced by multiple ecological and biological factors that are difficult to disentangle. In this study, we focus on understanding the role of seed dispersal by animals in the evolution of seed size in conifers, the most diverse extant nonflowering seed plant group. Relationships among seed size, dispersal syndrome, climate and cone morphology were analyzed across conifers using quantitative models of character evolution and phylogenetic regression techniques. Dispersal syndrome is a more consistent predictor of seed size within major extant conifer clades than climate. Seeds are generally larger in animal-dispersed than wind-dispersed species, and particular cone morphologies are consistently associated with specific ranges in seed size. Seed size and cone morphology evolve in a correlated manner in many animal-dispersed conifers, following a trade-off that minimizes the total size of the dispersal unit. These relationships are also present in other nonflowering seed plant groups, and have been important in the evolution of seeds and cones at least over the Cenozoic and perhaps over much of the later Mesozoic.

PREMISE OF THE STUDY Plants preserved in different fossil modes provide complementary data concerning the paleobiology and evolutionary relationships among plant groups. New material from the Early Jurassic of Patagonia shows the importance of combining these sources of information, as we describe the first compression/impression fossils of Pararaucaria, a genus of the extinct conifer family Cheirolepidiaceae previously known from permineralized fossils. These fossils extend the temporal range of this genus and may allow its wider recognition in the fossil record. METHODS We studied fossil plants from the Early Jurassic (Pleinsbachian-Toarcian) locality of Taquetren in Patagonia, Argentina using standard paleobotanical preparation and description techniques. KEY RESULTS Pararaucaria taquetrensis consists of isolated ovuliferous scales and small seed cones with helically arranged bract-scale complexes attached to scale-leaf foliage. Bract-scale complexes consist of separated bracts and ovuliferous scales with two seeds and three broad distal lobes. CONCLUSIONS Pararaucaria taquetrensis represents the oldest known Cheirolepidiaceae seed cones from the Southern Hemisphere, and this material highlights the importance of compression and impression fossils in understanding the distribution of fossil taxa. This material also suggests that Cheirolepidiaceae cone scales can be easily confused with those of another common conifer family, the Araucariaceae, which has important implications for accurately understanding Mesozoic conifer diversity and paleoecology.

Strap-shaped, parallel-veined leaves of Podozamites and Pseudotorellia are among the most common fossils in Late Triassic to Early Cretaceous floras from the Northern Hemisphere. Podozamites is considered to be the leaf of a voltzian conifer, but its cuticle is poorly known, whereas Pseudotorellia, with a thick cuticle and haplocheilic stomata, is commonly considered to be the leaf of a ginkgoalean. Here we describe the leaf morphology, cuticle and stomata of Podozamites and Pseudotorellia based on excellently preserved material from the Early Cretaceous of central Mongolia. Podozamites harrisii sp. nov. has transversely oriented, paracytic (probably syndetocheilic) stomata that are regularly arranged in longitudinal files. Pseudotorellia resinosa sp. nov. and Pseudotorellia palustris sp. nov. have scattered, longitudinally oriented stomata in which the two guard cells are sunken and surrounded by 2–5 specialized lateral subsidiary cells and 1–3 unspecialized polar cells. Association evidence and similari...

Premise of research. The two living species of Cunninghamia form a clade that is sister to the rest of the Cupressaceae sensu lato, within which the monotypic extant genus Taiwania is sister to all Cupressaceae except Cunninghamia. Seed cones of both Cunninghamia and Taiwania bear helically arranged bract-scale complexes, but a free ovuliferous scale is present only in Cunninghamia. Here, we describe two new genera of fossil seed cones similar to those of Cunninghamia and Taiwania.Methodology. Early Cretaceous (Aptian-Albian) bulk lignite samples collected from the Tevshiin Govi locality were disaggregated in water, cleaned with hydrochloric and hydrofluoric acids, washed, and dried in air.Pivotal results. Pentakonos gen. nov. has seed cones that are up to 7 mm long with ca. 25–30 helically arranged, papery bract-scale complexes that have minute teeth. Adaxially, the free tip of the membranous ovuliferous scale is denticulate, and the scale bears five small, thin-winged seeds. Pentakonos adds to the diver...

Seed dispersal is critical for plants, but the evolution of mechanisms that actually release seeds from their parents is not well understood. We use the reproductive cones of conifers, specifically the Pinaceae clade, to explore the factors driving the evolution of different release mechanisms in plants. We combine comparative anatomical and phylogenetic analyses to test whether fundamental trade-offs in the mechanical and hydraulic properties of vasculature underlie the evolution of two seed release mechanisms: cone scale flexion and cone scale shedding. We then test whether these mechanisms are linked with differences in seed size, dispersal syndrome and reproductive allocation. Cone scale xylem in flexing species is tough, but poorly conductive. Xylem in shedding species is less extensive, fragile and highly conductive; its thin-walled tracheids allow scales to easily fracture at maturity. Shedding is also consistently associated with large, densely packed seeds. Pinaceae cones exploit a well-known trade-off in xylem mechanical strength vs hydraulic efficiency to generate release mechanisms that allow seeds of various sizes to leave the protecting cone. The linkage among release mechanisms, vascular anatomy and seed traits illustrates how a wide variety of selective pressures may influence the function and physiology of reproductive structures.

Corystosperms, a key extinct group of Late Permian to Early Cretaceous plants, are important for understanding seed plant phylogeny, including the evolution of the angiosperm carpel and anatropous bitegmic ovule. Here, we describe a new species of corystosperm seed-bearing organ, Umkomasia mongolica sp. nov., based on hundreds of three-dimensionally preserved mesofossils from the Early Cretaceous of Mongolia. Individual seed-bearing units of U. mongolica consist of a bract subtending an axis that bifurcates, with each fork (cupule stalk) bearing a cupule near the tip. Each cupule is formed by the strongly reflexed cupule stalk and two lateral flaps that partially enclose an erect seed. The seed is borne at, or close to, the tip of the reflexed cupule stalk, with the micropyle oriented towards the stalk base. The corystosperm cupule is generally interpreted as a modified leaf that bears a seed on its abaxial surface. However, U. mongolica suggests that an earlier interpretation, in which the seed is borne directly on an axis (shoot), is equally likely. The 'axial' interpretation suggests a possible relationship of corystosperms to Ginkgo. It also suggests that the cupules of corystosperms may be less distinct from those of Caytonia than has previously been supposed.

Leaf waxes (e.g. n-alkanes, n-alkanoic acids) and their carbon isotopes (δ13C) are commonly used to track past changes in the carbon cycle, water availability, and plant ecophysiology. Previous studies indicated that conifers have lower n-alkane concentrations than angiosperms and that 13C fractionation during n-alkane synthesis (en-alkane) is smaller than in angiosperms. These prior studies, however, sampled a limited phylogenetic and geographic subset of conifers, leaving out many important subtropical and Southern Hemisphere groups that were once widespread and common components of fossil assemblages. To expand on previous work, we collected 43 conifer species (and Ginkgo biloba) from the University of California Botanical Garden at Berkeley, sampling all extant conifer families and almost two-thirds of extant genera. We find that Pinaceae, including many North American species used in previous studies, have very low or no n-alkanes. However, other conifer groups have significant concentrations of n-alkanes, especially Southern Hemisphere Araucariaceae and Podocarpaceae (monkey puzzles, Norfolk Island pines, and yellowwoods), and many species of Cupressaceae (junipers and relatives). Within the Cupressaceae, we find total n-alkane concentrations are high in subfamilies Cupressoideae and Callitroideae, but significantly lower in the early diverging taxodioid lineages (including bald cypress and redwood). Individual n-alkane chain lengths have a weak phylogenetic signal, except for n-C29 alkane, but when combined using average chain length (ACL), a strong phylogenetic signal emerges. The strong phylogenetic signal in ACL, observed in the context of a common growth environment for all plants we sampled, suggests that ACL is strongly influenced by factors other than climate. An analysis of en-alkane indicates a strong phylogenetic signal in which the smallest biosynthetic fractionation occurs in Pinaceae and the largest in Taxaceae (yews and relatives). The relationship between phylogeny and en-alkane may be related to differences in carbon metabolism among conifer clades. These results have important implications for interpreting n-alkane δ13C values in sedimentary archives, especially outside of North America.

Premise of research. Abundant fossil and molecular evidence suggests that all extant conifer families were established by the Early Cretaceous. However, the recognition and understanding of the lineages that lead to the evolution of these extant families remain incomplete. Late Paleozoic to Early Mesozoic Voltziales conifers—also known as transitional conifers, usually with multilobed ovuliferous scales—have been hypothesized to be among the stem lineages of modern crown conifers. This article describes an exquisitely preserved voltzian seed cone from the Aptian-Albian of Mongolia that introduces new taxonomic diversity and morphological data into the complex pattern of conifer evolution.Methodology. Bulk lignite samples collected from the Tevshiin Govi locality were disaggregated in water, cleaned with hydrochloric and hydrofluoric acids, washed, and dried in air. Fossils were examined using LM, SEM, and X-ray microtomography.Pivotal results. Krassilovia mongolica gen. et sp. nov. has seed cones with hel...

Integration influences patterns of trait evolution, but the relationship between these patterns and the degree of trait integration is not well understood. To explore this further, we study a specialized pollination mechanism in conifers whose traits are linked through function but not development. This mechanism depends on interactions among three characters: pollen that is buoyant, ovules that face downward at pollination, and the production of a liquid droplet that buoyant grains float through to enter the ovule. We use a well-sampled phylogeny of conifers to test correlated evolution among these characters and specific sequences of character change. Using likelihood models of character evolution, we find that pollen morphology and ovule characters evolve in a concerted manner, where the flotation mechanism breaks down irreversibly following changes in orientation or drop production. The breakdown of this functional constraint, which may be facilitated by the lack of developmental integration among the constituent traits, is associated with increased trait variation and more diverse pollination strategies. Although this functional "release" increases diversity in some ways, the irreversible way in which the flotation mechanism is lost may eventually result in its complete disappearance from seed plant reproductive biology.

Premise of research. Exceptionally well-preserved lignified fossils from the Early Cretaceous of Mongolia include abundant conifer leafy shoots with attached pollen cones and seed cones. A whole-plant reconstruction based on these fossils enables a critical evaluation of the relationship of this extinct plant with extant conifers.Methodology. Bulk lignite samples collected from the Tugrug lignite mine were disaggregated in water, cleaned with hydrochloric and hydrofluoric acids, washed, and dried in air. Fossils were then examined using light and scanning electron microscopy.Pivotal results. Elatides zhoui sp. nov. has helically arranged leaves with two narrow lateral stomatal bands, predominantly on the adaxial leaf surface. Pollen cones are usually borne laterally on shoots in tight spirals; each microsporophyll bears three pollen sacs that produce nonsaccate pollen with a small circular aperture. Seed cones have numerous bract-scale complexes, each with a small membranous ovuliferous scale and four to ...

Summary The relationship between branch diameter and leaf size has been widely used to understand how vegetative resources are allocated in plants. Branching architecture influences reproductive allocation as well, but fewer studies have explored this relationship at broad phylogenetic or ecological scales. In this study, we tested whether pollen-producing and seed-producing cone size scales with branch diameter in conifers, a diverse and globally distributed lineage of nonflowering seed plants. Branch diameter and cone size were analyzed using multiple regression models and evolutionary models of trait evolution for a data set of 293 extant conifer species within an explicit phylogenetic framework. Branch diameter is a strong predictor of cone size across conifer species, particularly for pollen cones and dry seed cones. However, these relationships are complex in detail because leaf morphology and seed dispersal biology influence the specific ways in which they are expressed. The ubiquity and strength of these scaling relationships across conifers suggest that reproductive and vegetative morphologies are coupled in the group, and it is therefore difficult to disentangle the evolution of cone size from the evolution of branching architecture.

Island species are thought to be extinction-prone because of small population sizes, restricted geographical distribution and limited dispersal ability. However, the topographical and environmental heterogeneity, geographical isolation and stability of islands over long timescales could create refugia for taxa whose source area is threatened by environmental changes. We address this possibility by inferring the evolution of the New Caledonia (NC) and New Zealand (NZ) conifer diversity, which represents over 10% of the world’s diversity for this group. We estimate speciation and extinction rates in relation to the presence/absence on these islands, and dispersal rates between the islands and surrounding areas. We also test the Eocene submersion of NC and the Oligocene drowning of NZ by comparing the fit of biogeographical scenarios using ancestral area estimations. We find that extinction rates were significantly lower for island species, and dispersal “out of islands” was higher. A model including a diversification shift when NC emerged better explains the diversification dynamics. Biogeographical analyses corroborate that conifers experienced high continental extinctions, but survived on islands. NC and NZ have thus contributed to the world’s conifer diversity as “island refugia”, by maintaining early-diverging lineages from continents during environmental changes on continents. These ancient islands also acted as “species pumps”, providing species into adjacent areas. Our study highlights the important but neglected role of islands in promoting the evolution and conservation of biodiversity. © The Willi Hennig Society 2016.

†Background and Aims Correlated relationships between branch thickness, branch density, and twig and leaf size have been used extensively to study the evolution of plant canopy architecture, but fewer studies have explored the impact of these relationships on the allocation of reproductive resources. This study quantifies pollen cone production in conifers, which have similar basic reproductive biology but vary dramatically in branching habit, in order to test how differences in branch diameter influence pollen cone size and the density with which they are deployed in the canopy. †Methods Measurements of canopy branch density, the number of cones per branch and cone size were used to estimate the amount of pollen cone tissues produced by 16 species in three major conifer clades. The number of pollen grains produced was also estimated using direct counts from individual pollen cones. †Key Results The total amount of pollen cone tissues in the conifer canopy varied little among species and clades, although vegetative traits such as branch thickness, branch density and pollen cone size varied over several orders of magnitude. However, branching habit controls the way these tissues are deployed: taxa with small branches produce small pollen cones at a high density, while taxa with large branches produce large cones relatively sparsely. †Conclusions Conifers appear to invest similar amounts of energy in pollen production independent of branching habit. However, similar associations between branch thickness, branch density and pollen cone size are seen across conifers, including members of living and extinct groups not directly studied here. This suggests that reproductive features relating to pollen cone size are in large part a function of the evolution of vegetative morphology and branching habit.

A broad perspective of seed plant reproductive biology, including information from both living and fossil gymnosperm groups, is crucial in order to understand the evolution of angiosperm reproductive innovations. Within this context, angiosperms are unique in the degree to which they have shifted reproductive functions away from their ovules to associated tissues and organs. This is particularly true in regard to the carpel, which has assumed roles in pollination, seed protection, and seed dispersal previously restricted to the ovule itself. Perhaps as a consequence, angiosperm ovule development is more efficient than in nearly all other groups of living and extinct seed plants, where reproductive functions often constrain both patterns of development and the allocation of resources. For example, many living gymnosperm species are as efficient as angiosperms in minimizing ovule development prior to pollination, but they require a megagametophyte for embryo provisioning that constrains their efficiency in minimizing investment prior to fertilization. By decoupling ovules from pollination, the carpel also fundamentally altered the way in which pollination occurs in angiosperms relative to other seed plants. In particular, this change gave angiosperms greater control over the spatial positioning of their reproductive organs and may have therefore expanded the types of reproductive structures angiosperms could develop and the specificity with which these structures could interact with pollinators.

Angiosperms represent more than three-quarters of all land plant species despite existing for only a quarter of land plant history, inspiring much interest in understanding what aspects of angiosperm biology may have enabled their unparalleled success. However, comparative study of extant plants cannot fully address the basis of angiosperm dominance because most living nonflowering plants are members of recently radiating lineages as young or younger than the flowering plants and, thus, are a highly biased sampling of the full 400 Myr history of vascular plant diversity. Here, a survey of the fossil record demonstrates that most anatomical traits that are now unique to the angiosperms were more broadly distributed among extinct lineages. However, the growth capacities of angiosperms are likely to have been uniquely high for all of vascular plant history and may have enabled a greater range of architectural possibilities, including the fast-growing annual herbs that represent a large proportion of angiospe...

Conifers are an excellent group in which to explore how changing ecological interactions may have influenced the allocation of reproductive tissues in seed plants over long time scales, because of their extensive fossil record and their important role in terrestrial ecosystems since the Palaeozoic. Measurements of individual conifer pollen-producing and seed-producing cones from the Pennsylvanian to the Recent show that the relative amount of tissue invested in pollen cones has remained constant through time, while seed cones show a sharp increase in proportional tissue investment in the Jurassic that has continued to intensify to the present day. Since seed size in conifers has remained similar through time, this increase reflects greater investment in protective cone tissues such as robust, tightly packed scales. This shift in morphology and tissue allocation is broadly concurrent with the appearance of new vertebrate groups capable of browsing in tree canopies, as well as a diversification of insect-feeding strategies, suggesting that an important change in plant–animal interactions occurred over the Mesozoic that favoured an increase in seed cone protective tissues.

Exploring patterns in the evolution of seed plant reproductive morphology within a functional context offers a framework in which to identify and evaluate factors that potentially drive reproductive evolution. Conifers are a particularly useful group for studies of this kind because they have a long geologic history and their reproductive organs are borne on separate structures with discrete functions. Multivariate analysis of morphological data collected from pollen-producing and seed-producing cones of Paleozoic, Mesozoic, and extant conifer species shows that seed cones underwent a significant expansion of morphological diversity that began during the Early-Middle Jurassic and has continued into the present day. In contrast, pollen cones show significantly lower levels of morphological diversity and exhibit similar basic morphologies throughout conifer evolutionary history. The increase in seed cone diversity through time is primarily the result of two novel structural and organizational features that evolved independently in different conifer families during the Mesozoic: robust, tightly packed cones in members of Araucariaceae, Cupressaceae sensu lato, and Pinaceae, and highly reduced, fleshy cones or solitary seeds in Podocarpaceae, Taxaceae, and some members of Cupressaceae sensu stricto. In extant conifers, these cone morphologies are associated with species that have strong interactions with vertebrate seed predators, seed dispersers, or a combination of both. This suggests that increases in the strength and complexity of biotic interactions in the Jurassic and Cretaceous were a primary driver of conifer reproductive evolution, and that patterns of character evolution relate to the increasing importance of cone tissue in seed protection and seed dispersal through time.

Fossil floras from three exposures of the Emma Fiord Formation from Ellesmere Island, Axel Heiberg Island, and Devon Island in the Canadian Arctic Archipelago fill a gap in our knowledge of the regional distribution of Visean age plant assemblages. Plant fossils occur as compressions in two lithologies, calcareous shale and black shale, representing two different lacustrine depositional environments within a broader system of rift lakes that developed prior to the beginning of widespread Sverdrup Basin sedimentation. Calcareous shales from the Grinnell Peninsula of Devon Island preserve an in situ forest dominated by articulated arborescent lycopsid compression fossils similar to Lepidodendron veltheimii , while black shales from the Kleybolte Peninsula of Ellesmere Island and the Svartevaeg Cliffs of Axel Heiberg Island preserve a more diverse assemblage consisting of fragmentary lycopsid compressions and disarticulated foliage taxa similar to species of Rhodea , Aneimites , and Fryopsis . The Emma Fiord macroflora is comparable to those found in similar tectonic and sedimentary settings in Alaska, Greenland, and Spitsbergen. These assemblages are not as diverse as contemporaneous or roughly contemporaneous tropical lowland floras from North America and Europe.

Upatoia barnardii gen. et sp. nov., a conifer pollen cone from the Late Cretaceous (Santonian) Eutaw Formation of Upatoi Creek, Georgia, USA, is known from lignified and fusainised mesofossils that preserve its three-dimensional structure. The cone consists of numerous helically arranged microsporophylls, each composed of a thin stalk and distal lamina. Three elongate pollen sacs are attached to the base of the lamina. Pollen grains isolated from the pollen sacs are relatively large (52 – 75 μm), spheroidal to ellipsoidal in outline, lack sacci, and have a thickened equatorial exine that is often strongly folded. Pollen of Upatoia barnardii indicates a close relationship to extant Araucariaceae. Microsporophylls of U. barnardii confirm suggestions from previous studies of fossil material that some Mesozoic Araucariaceae had only three pollen sacs per microsporophyll, in contrast to extant species that often have more than ten pollen sacs per microsporophyll.

Saccate pollen is present in many ancient seed plants and in two families of extant conifers. In living conifers, saccate pollen occurs in taxa that combine erect ovulate cones at the time of pollination, inverted ovules, and pollination drops. Sacci function primarily to float pollen grains through a liquid pollination drop and concentrate them near the nucellus of the ovule. The morphology and anatomy of sacci are crucial to the proper operation of this pollination mechanism, and similar size proportions in fossil and modern saccate grains suggest that ancient pollen may have functioned in the same way as modern forms. Multivariate analysis of discrete pollen characters performed on data from in situ grains of several groups of fossil seed plants indicates that modern saccus morphology and structure appear as early as the Upper Carboniferous in some nonconiferous seed plants but later in the evolutionary history of the conifer lineage. Combined evidence from pollen morphology suggests that sacci may hav...

Integration influences patterns of trait evolution, but the relationship between these patterns and the degree of trait integration is not well understood. To explore this further, we study a specialized pollination mechanism in conifers whose traits are linked through function but not development. This mechanism depends on interactions among three characters: pollen that is buoyant, ovules that face downward at pollination, and the production of a liquid droplet that buoyant grains float through to enter the ovule. We use a well-sampled phylogeny of conifers to test correlated evolution among these characters and specific sequences of character change. Using likelihood models of character evolution, we find that pollen morphology and ovule characters evolve in a concerted manner, where the flotation mechanism breaks down irreversibly following changes in orientation or drop production. The breakdown of this functional constraint, which may be facilitated by the lack of developmental integration among the constituent traits, is associated with increased trait variation and more diverse pollination strategies. Although this functional "release" increases diversity in some ways, the irreversible way in which the flotation mechanism is lost may eventually result in its complete disappearance from seed plant reproductive biology.

 Premise of the study: Seed cone scales assigned to the genus Schizolepidopsis are widespread in Late Triassic to Cretaceous Eurasian deposits. They have been linked to the conifer family Pinaceae based on associated vegetative remains, but their exact affi nities are uncertain. Recently discovered material from the Early Cretaceous of Mongolia reveals important new information concerning Schizolepidopsis cone scales and seeds, and provides support for a relationship between the genus and extant Pinaceae.  Methods: Specimens were collected from Early Cretaceous (probable Aptian-Albian) lignite deposits in central Mongolia. Lignite samples were disaggregated, cleaned in hydrofl uoric acid, and washed in water. Specimens were selected for further study using light and electron microscopy.  Key results: Schizolepidopsis canicularis seed cones consist of loosely arranged, bilobed ovulate scales subtended by a small bract. A single inverted seed with an elongate micropyle is borne on each lobe of the ovulate scale. Each seed has a wing formed by the separation of the adaxial surface of the ovulate scale.  Conclusions: Schizolepidopsis canicularis produced winged seeds that formed in a manner that is unique to Pinaceae among extant conifers. We do not defi nitively place this species in Pinaceae pending more complete information concerning its pollen cones and vegetative remains. Nevertheless, this material suggests that Schizolepidopsis may be important for understanding the early evolution of Pinaceae, and may potentially help reconcile the appearance of the family in the fossil record with results based on phylogenetic analyses of molecular data .

The evolution of plants exhibiting different sexes, or dioecy, is correlated with a number of ecological and life-history traits such as woody growth form and animal-dispersed seeds, but the underlying causes of these associations are unclear. Previous work in seed plants has suggested that the evolution of fleshy cones or seeds may favour dioecy. In this study, we use a well-sampled molecular phylogeny of conifers to show that although dioecy and fleshiness strongly co-occur at the species level, this relationship has not resulted from numerous separate origins of this trait combination or from differential rates of diversification. Instead, we suggest that two character combinations—the ancestral dry-monoecious condition and the derived fleshy-dioecious condition—have persisted in conifers longer than other combinations over evolutionary time. The persistence of these trait combinations appears to reflect differences in the rate of successful transition into and out of these character states over time, as well as the geographical restriction of species with rare combinations and their consequent vulnerability to extinction. In general, we argue that such persistence explanations should be considered alongside ‘key innovation’ hypotheses in explaining the phylogenetic distribution of traits.

Fundamental differences in the distribution of oceans and landmasses in the Northern and Southern Hemispheres potentially impact patterns of biological diversity in the two areas. The evolutionary history of conifers provides an opportunity to explore these dynamics, because the majority of extant conifer species belong to lineages that have been broadly confined to the Northern or Southern Hemisphere during the Cenozoic. Incorporating genetic information with a critical review of fossil evidence, we developed an age-calibrated phylogeny sampling ∼80% of living conifer species. Most extant conifer species diverged recently during the Neogene within clades that generally were established during the later Mesozoic, but lineages that diversified mainly in the Southern Hemisphere show a significantly older distribution of divergence ages than their counterparts in the Northern Hemisphere. Our tree topology and divergence times also are best fit by diversification models in which Northern Hemisphere conifer lineages have higher rates of species turnover than Southern Hemisphere lineages. The abundance of recent divergences in northern clades may reflect complex patterns of migration and range shifts during climatic cycles over the later Neogene leading to elevated rates of speciation and extinction, whereas the scattered persistence of mild, wetter habitats in the Southern Hemisphere may have favored the survival of older lineages.

Summary •Among many species of living conifers the presence of pollen with air bladders (saccate pollen) is strongly associated with downward-facing ovules and the production of pollination drops. This combination of features enables saccate pollen grains captured in the pollination drop to float upwards into the ovule. Despite the importance of this mechanism in understanding reproduction in living conifers and in extinct seed plants with similar morphologies, experiments designed to test its effectiveness have yielded equivocal results. •In vitro and in vivo pollination experiments using saccate and nonsaccate pollen were performed using modeled ovules and two Pinus species during their natural pollination period. •Buoyant saccate pollen readily floated through aqueous droplets, separating these grains from nonbuoyant pollen and spores. Ovules that received saccate pollen, nonsaccate pollen or a mixture of both all showed larger amounts and higher proportions of saccate pollen inside ovules after drop secretion. •These results demonstrate that flotation is an effective mechanism of pollen capture and transport in gymnosperms, and suggest that the prevalence of saccate grains and downward-facing ovules in the evolutionary history of seed plants is a result of the widespread use of this mechanism.