The trade-off between vegetative and generative reproduction among angiosperms influences regional hydrochorous propagule pressure

Aim Local communities are subject to spatiotemporal contingencies of landscape processes; community assembly is thus often considered to be unpredictable and idiosyncratic. However, evolved trade‐offs of species’ life histories may set distinct constraints on the assembly of species communities. In plants, the recruitment and invasion success of species into communities depend primarily on the number of propagules available and on their generative or vegetative character. Life‐history trade‐offs prevent individual plants from producing large numbers of both generative and vegetative propagules, but it is not clear whether this constrains their availability at the landscape scale. We thus tested whether: (1) the observed relationship between generative and vegetative propagules deviates from the null expectation stating that species contributing the bulk of generative propagules to the propagule rain should also contribute the bulk of vegetative propagules; and (2) whether vegetative and generative propagule pressures are negatively correlated once species abundance in the regional pool is accounted for. Location A large riparian landscape in the Netherlands. Methods Analyses were based on an extensive trapping of floating propagules (214,049 propagules of 47 species), and a rough proxy of species abundance across the entire pool. We used both species and phylogenetically independent contrasts as data points, and accounted for variation in size of generative propagules. Results Both hypotheses were confirmed. Numbers of generative and vegetative propagules trapped per species were significantly negatively correlated (r = ?0.33; t₄₅ = ?2.61, P = 0.006) and thus strongly deviated from the null expectation. This was confirmed by analyses accounting for variation in species abundance across the species pool, and in the size of generative propagules. Main conclusions The results indicate that plant recruitment and community assembly across streams may be influenced by the way individual plants allocate their resources between competing life‐history functions. Life‐history evolution across angiosperms might thus have constrained the present‐day assembly of local communities.     

Evolution and patterning of the ovule in seed plants

The ovule and its developmental successor, the seed, together represent a highly characteristic feature of seed plants that has strongly enhanced the reproductive and dispersal potential of this diverse group of taxa. Ovules encompass multiple tissues that perform various roles within a highly constrained space, requiring a complex cascade of genes that generate localized cell proliferation and programmed cell death during different developmental stages. Many heritable morphological differences among lineages reflect relative displacement of these tissues, but others, such as the second (outer) integuments of angiosperms and Gnetales, represent novel and apparently profound and independent innovations. Recent studies, mostly on model taxa, have considerably enhanced our understanding of gene expression in the ovule. However, understanding its evolutionary history requires a comparative and phylogenetic approach that is problematic when comparing extant angiosperms not only with phylogenetically distant extant gymnosperms but also with taxa known only from fossils. This paper reviews ovule characters across a phylogenetically broad range of seed plants in a dynamic developmental context. It discusses both well-established and recent theories of ovule and seed evolution and highlights potential gaps in comparative data that will usefully enhance our understanding of evolutionary transitions and developmental mechanisms.     

Yield assessment of integument-led seed growth following targeted repair of auxin response factor 2

SUMMARY: It is becoming increasingly vital to improve the yield of seed crops to feed an expanding population and, more recently, for biofuel production. One strategy to increase the yield is to increase the seed size, provided that there is not a concomitant decrease in seed number. In a previous study, we described a mutant in the auxin response factor 2 (ARF2) gene which produced extra cells in the seed coat and, subsequently, enlarged seeds. However, arf2 mutant plants also show severely reduced self-fertility caused, in part, by over-elongated sepals that prevent flower opening. As a low seed set increases individual seed size, a meaningful comparison of the yield in arf2 and wild-type plants could not be conducted. In this study, we show that targeted expression of wild-type ARF2 in the sepals and petals of arf2-9 mutant flowers restores flower opening and dramatically increases seed set. The restored plants retain both enlarged integuments and increased seed size, reinforcing previous evidence that arf2 mutations increase seed weight through their effect on integuments and not only via reduced fertility. We also show that the measurement of the harvest index in Arabidopsis is useful in assessing the impact of introduced traits on the yield.     

The Effect of Timing of Rehabilitation Procedures on the Establishment of a Jarrah Forest After Bauxite Mining

The restoration of the high botanical diversity of the premining jarrah (Eucalyptus marginata) forest is a major priority of rehabilitation following bauxite mining in southwestern Australia. This study investigated the effects of different ripping, seeding, and scarifying dates on the establishment of plants from propagules stored in the topsoil and from applied seed on areas being rehabilitated after mining. Seed stored in the topsoil, rather than applied seed, was the major contributor to plant diversity. Ripping late (April) or scarifying in June significantly reduced the number of species and numbers of individual plants that established from propagules in the topsoil. Species originating from broadcast seed were most numerous when the seed was broadcast in April or after scarifying in June. Scarifying before seeding, particularly in June, increased the establishment of species from the broadcast seed. To make best use of the applied seed, without jeopardizing the establishment of species from the topsoil, pits should be ripped and sown by April. We list a number of strategies that can help maximize plant numbers and botanical diversity on rehabilitated bauxite mines, which may also be of more general application for restoring the original native vegetation on disturbed sites.     

The role of Hydropteris pinnata gen. et sp. nov. in reconstructing the cladistics of heterosporous ferns

Large segments of intact plants that represent a heterosporous fern have been discovered within an aquatic plant community from the Late Cretaceous St. Mary River Formation near Cardston in southern Alberta, Canada. Branching rhizomes of Hydropteris pinnata gen. et sp. nov. are 1–2 mm wide. They produce fronds at intervals of 2–12 mm and bear numerous elongated roots. Fronds, up to approximately 6 cm long, are pinnate with subopposite to alternate pinnae that exhibit anastomosing venation. Large, multisoral sporocarps occur at the junctures of the rhizome and frond rachides. Both microsporangiate massulae and megaspore complexes occur within each sporocarp. Megaspore complexes are assignable to the sporae dispersae genus Parazolla Hall. Microspores are trilete, smooth-walled, and are embedded in episporal material of the massulae. A numerical cladistic analysis indicates that the heterosporous aquatic ferns are monophyletic, and not as closely related to either schizaeaceous or hymenophyllaceous ferns as they are to some other filicaleans. Systematic revisions are proposed to reflect newly recognized cladistic relationships within the heterosporous clade, and character originations in the evolution of heterosporous aquatic ferns are evaluated. Hydropteridaceae fam. nov. is proposed, and included with Salviniaceae and Azollaceae in the Hydropteridineae subord. nov., and the Hydropteridales Willdenow.     

Many alien invasive plants disperse against the direction of stream flow in riparian areas

Propagule pressure plays an important role in the invasion of alien plants into riparian areas. In this study, we focused on propagule pressure from both neighboring riparian areas and anthropogenic land-use areas because propagules are likely to originate from both sources. We tested the effects of whether neighboring units contained the alien plant species, focusing on the direction of invasion by alien plant species into the focal unit, and how much anthropogenic land was contained within the unit, focusing on both farmland and urbanized areas, on alien plant occurrences in Hyogo Prefecture, Japan. We modeled the occurrence of 10 alien plants using generalized linear models to evaluate species invasions by both propagules from both neighboring units and anthropogenic land within a unit. We also investigated the biological and ecological plant attributes that are likely related to invasion success, such as seed dispersal methods, seed size, and clonality, and tested the relationships between the model results and each species’ attributes. Results showed that the occurrence of an affected neighboring unit was positively associated with the occurrence of all 10 alien plants. Note that two alien invasive species were influenced by upstream flow direction, six species by downstream flow direction, and in two species, propagule supply was not distinguished by direction. In short, the dominant direction of dispersal was against the stream current, while dispersal in the downstream direction was less common. Species attributes were associated with these directions of dispersal. In addition, anthropogenic land was positively associated with the occurrence of most alien plants, although this effect was weaker than the neighbor unit effects. These results indicate that alien plants spreading into riparian areas do not always follow the natural flow regime; rather, they spread against the flow regime in some cases. We discuss an ecological explanation for these results and provide perspectives for future river management of alien plants that invade the riparian zone.     

Maternal control of seed size by EOD3/CYP78A6 in Arabidopsis thaliana

Seed size in higher plants is coordinately determined by the growth of the embryo, endosperm and maternal tissue, but relatively little is known about the genetic and molecular mechanisms that set final seed size. We have previously demonstrated that Arabidopsis DA1 acts maternally to control seed size, with the da1-1 mutant producing larger seeds than the wild type. Through an activation tagging screen for modifiers of da1-1, we have identified an enhancer of da1-1 (eod3-1D) in seed size. EOD3 encodes the Arabidopsis cytochrome P450/CYP78A6 and is expressed in most plant organs. Overexpression of EOD3 dramatically increases the seed size of wild-type plants, whereas eod3-ko loss-of-function mutants form small seeds. The disruption of CYP78A9, the most closely related family member, synergistically enhances the seed size phenotype of eod3-ko mutants, indicating that EOD3 functions redundantly with CYP78A9 to affect seed growth. Reciprocal cross experiments show that EOD3 acts maternally to promote seed growth. eod3-ko cyp78a9-ko double mutants have smaller cells in the maternal integuments of developing seeds, whereas eod3-1D forms more and larger cells in the integuments. Genetic analyses suggest that EOD3 functions independently of maternal factors DA1 and TTG2 to influence seed growth. Collectively, our findings identify EOD3 as a factor of seed size control, and give insight into how plants control their seed size.     

Optimal resource allocation to seeds and vegetative propagules under density-dependent regulation in Syneilesis palmata (Compositae)

Syneilesis palmata reproduces by both seeds and vegetative propagules (short rhizomes). The latter result in the production of new plants that are larger in size and hence have a higher survival probability and a higher growth rate than seeds. A previous study predicted that the optimal reproductive strategy, in terms of maximizing population growth rate (a fitness measure under no density regulations), was pure vegetative reproduction. However, high resource investment to vegetative propagules can cause local crowding resulting in reduced demographic performances of the plants, because the vegetative propagules of Syneilesis are produced close to one another. We examined, in this situation, the impact of allocating a certain proportion of reproductive resource to seeds with relatively greater capacity for dispersal. We simulated dynamics of hypothetical Syneilesis populations with various reproductive resource allocation balances (from pure seed to pure vegetative reproduction), using a density-dependent matrix model. In the model, it was assumed that plants from vegetative propagules experienced density-dependent reduction in their survival probabilities, but this was not the case for plants originating from seeds. Each allocation strategy was evaluated based on an equilibrium population density, a fitness measure under density-dependent regulations. The optimal reproductive strategy predicted was pure vegetative reproduction. Unrealistic conditions were required for seed reproduction to be favoured, such as the production of seeds one hundred times the normal number per unit resource investment. However, the conditions were fairly relaxed compared with those required in the model where no density effects were incorporated. This indicates that escape from local crowding is likely to be one of the roles of seed production in Syneilesis.     

Direct and indirect effects of pollinators and seed predators to selection on plant and floral traits

Although flowering traits are often assumed to be under strong selection by pollinators, significant variation in such traits remains the norm for most plant species. Thus, it is likely that the interactions among plants, mutualists, and other selective agents, such as antagonists, ultimately shape the evolution of floral and flowering traits. We examined the importance of pollination vs pre-dispersal seed predation to selection on plant and floral characters via female plant-reproductive success in Castilleja linariaefolia (Scrophulariaceae). C. linariaefolia is pollinated by hummingbirds and experiences high levels of pre-dispersal seed predation by plume moth and fly larvae in the Rocky Mountains of Colorado, USA, where this work was conducted. We first examined whether female reproduction in C. linariaefolia was limited by pollination. Supplemental pollination only marginally increased components of female reproduction, likely because seed predation masked, in part, the beneficial effects of pollen addition. In unmanipulated populations, we measured calyx length, flower production, and plant height and used path analysis combined with structural equation modeling to quantify their importance to relative seed set through pathways involving pollination vs seed predation. We found that the strength of selection on calyx length, flower production, and plant height was greater for seed predation pathways than for pollination pathways, and one character, calyx length, experienced opposing selection via pollination vs seed predation. These results suggest that the remarkable intraspecific variation in plant and floral characters exhibited by some flowering plants is likely the result of selection driven, at least in part, by pollinators in concert with antagonists, such as pre-dispersal seed predators. This work highlights the subtle but complex interactions that shape floral and vegetative design in natural ecosystems.     

Spore wall ultrastructure in the early lycopsid Leclercqia (Protolepidodendrales) from the Lower Devonian of North America: Evidence for a fundamental division in the lycopsids

Documenting the morphology and ultrastructure of spores from known Silurian-Devonian plants clarifies organization and probable affinities of dispersed spores and contributes to analyses of evolutionary changes and phylogenetic relationships in early plants. In this study of fossil in situ spores from the early protolepidodendralean lycopsid Leclercqia, we identified new characters including an additional synapomorphy of the ligulate lycopsid clade. A detailed light (LM), scanning electron (SEM), and transmission electron microscope (TEM) analysis of spores from two species of Leclercqia from the Lower Devonian (Emsian) of New Brunswick, eastern Canada, L. andrewsii and L. complexa, shows both are homosporous, yielding spores belonging to the dispersed spore form taxon Acinosporites lindlarensis. Important features of wall ultrastructure include the presence of a paraexospore, peculiar exospore-derived, peg-like structures located in the gap between the outer exospore/inner paraexospore, and multilamellate regions in the interradial areas of the proximal surface. Similar interradial multilamellate regions occur in other ligulate lycopsids (fossil and extant). This character is probably a further synapomorphy for the ligulate lycopsid clade, within which heterosporous lycopods form a monophyletic group. These data suggest the ligule and interradial multilamellate region appeared prior to heterospory.     

植物功能性状与外来植物入侵

揭示影响外来植物入侵性的功能性状及其生态机制是入侵植物生态学的核心任务之一。本文综述了植物功能性状与外来植物入侵性的研究进展, 通过分析植物功能性状对外来植物入侵的贡献以及外来植物的不同入侵阶段对其功能性状的需求, 探讨植物功能性状与外来植物入侵的相关性及其入侵机理。迄今研究较多的影响外来植物入侵性的功能性状主要包括形态性状、生长性状、生理性状、繁殖性状、种子性状、克隆性状、表型可塑性和遗传变异等。这些功能性状对外来植物入侵的贡献随着入侵阶段的不同而变化。在传播到达阶段, 种子性状对入侵具有重要影响; 在定居建群阶段, 与植物抗逆性和适应性相关的生理性状和繁殖性状发挥主要作用; 在扩散入侵阶段, 克隆性状和影响植物竞争能力的生理性状对植物成功入侵具有重要贡献。由于植物入侵性是其功能性状和环境因素互作的结果, 且功能性状的作用随环境因素和入侵阶段不同而异, 因此, 结合外来植物入侵阶段, 并考虑功能性状与环境因子的互作, 是入侵生物学中植物功能性状研究的发展趋势。     

Adaptive introductions: How multiple experiments and comparisons to wild populations provide insights into requirements for long-term introduction success of an endangered shrub

Recovery of an imperiled plant species may require augmentation of existing populations or creation of new ones. Hundreds of such projects have been conducted over the last few decades, but there is a bias in the literature favoring successes over failures. In this paper, we evaluate a series of introductions that experimentally manipulated microhabitat and fire in an adaptive introduction framework. Between 2002 and 2012, we (and our collaborators) carried out ten introductions and augmentations of Florida ziziphus Pseudoziziphus (Condalia, Ziziphus) celata, a clonal shrub limited to very small populations and narrowly endemic to pyrogenic central Florida sandhills. Six of the introductions were designed as experiments to test hypotheses about how demographic performance was affected by microhabitat, fire, and propagule type. Introduced transplants had high survival (<90% annually), inconsistent growth, and little transition to reproduction, while introduced seeds had low germination and survival. Transplants were more efficient than seeds as translocation propagules. Shaded (vs. open) sites supported generally higher transplant and seedling survival and seed germination percentages, but growth responses varied among experiments. Supplemental irrigation increased transplant survival and seed germination, but otherwise seedling and plant survival and growth were not significantly affected. Contrary to expectations based on wild populations, introduced propagules have not been more successful in unshaded sites, suggesting that Florida ziziphus has broader microhabitat preferences than hypothesized. Compared to wild plants, introduced plants had similar survival and responses to fire, slower growth, and more delayed flowering. Introduced plants had no clonal spread. While no introduced population has demonstrated a capacity for long-term viability, one augmented population has flowered and produced viable fruits. Given that Florida ziziphus genets are long-lived, low levels of sexual reproduction may be adequate for the establishment of viable populations. Thus, after many translocations over more than a decade, it is premature to characterize any single translocation as a success or a failure, underscoring the need for a long view of translocation success.     

OVULE AND SEED DEVELOPMENT IN CERTAIN CACTI

Engleman , E. Mark . (U. of California, Davis.) Ovule and seed development in certain cacti. Amer. Jour. Bot. 47(6) : 460–467. Illus. 1960.—Ovules of Astrophytum myriostigma, Thelocactus bicolor, and Toumeya papyracantha are crassinucellate, possess a nucellar cap, and have a micropyle formed only by the inner of 2 integuments. Survival of the chalazal megaspore results in a monosporic gametophyte. A single layer of endosperm cells forms a cap over the radicle in a mature seed, as occurs in some of the Centrospermae. The characteristic hilum cup consists of the hilum and variable amounts of thickened cells of the funiculus which border a cuplike cavity. The hilum cup is associated with a funicular hump near the junction of the funiculus and the integuments. The funicular hump is similar in position to a swelling on the funiculus of some members of the Centrospermae.     

Small-scale spatial structure within patterns of seed dispersal

A large proportion of dispersing propagules land near their maternal plant, even in species that have evolved structures which enhance dispersal. For these propagules, their post-dispersal spatial pattern is likely to reflect the overall shape and scale of the parental plant canopy and, especially in poorly dispersing species, aggregation of propagules on the plant prior to dispersal. Localised patterns within seed shadows are also likely to be affected by secondary movement after dispersal, leading to either more or less small-scale aggregation, depending on the mechanism. Our general aim was to study the small-scale spatial structure within patterns of seed dispersal of Raphanus raphanistrum L. to generate hypotheses about the sequence of processes and events leading to the spatial pattern of dispersal in this species. More specifically, we determined the sizes of small-scale structures within the seed shadows on the ground after dispersal and the extent to which these match the sizes of pre-dispersal aggregations within the parental canopy. Variation in plant size and shape was provided by four levels of inter-specific competition resulting from differing wheat crop densities. Positions of propagules were determined using a three-dimensional digitizer, and the data for each plant were analysed using point pattern analysis. Not surprisingly, larger plants, growing at lower plant density, had larger seed shadows, showing an overall influence of maternal plant size. The pattern of propagules exhibited significant small-scale aggregates, with similar sizes on the plant and on the ground. There was no evidence that aggregation size was greater on the ground or increased with time, but the strength of the aggregation increased with length of time on the ground.     

动物对孢子植物的传播模式及进化意义

孢子植物物种多样性丰富, 是自然生态系统的重要组成部分。孢子植物的传播通常被认为主要依靠风、水、弹力等非生物媒介, 而动物的作用往往被忽略。本文主要概述了: (1)孢子植物对动物传播的适应: 一方面孢子植物可为动物提供食物、庇护所、繁殖场所等, 另一方面孢子植物也可产生视觉、嗅觉等方面的线索来吸引动物, 从而促进动物传播其繁殖体。(2)动物对孢子植物的传播模式: 包括体内传播(消化道和组织寄生)和体外传播两种, 这些模式都能对孢子植物繁殖体进行有效传播。由于动物间形态或生活习性的不同, 以致传播距离存在差异, 最短距离为0.1 cm, 最长距离可从北半球至南半球。(3)动物对孢子植物传播的生态与进化意义; 由于某些孢子植物繁殖体的结构特点或萌发的需求, 以致其繁殖体只能通过动物的传播才能得以定殖, 因此动物与孢子植物之间存在密不可分的关系。目前, 动物对孢子植物的传播研究主要是描述性的内容以及研究单方面的传播途径, 建议在今后的研究中考虑动物对孢子植物传播的有效性以及多途径同时传播对孢子植物定殖的影响, 同时应更加关注孢子植物和动物互惠关系的形成、维持机制及将来的进化趋势。     

Climate drives differences in the germination niche of a globally distributed invasive grass

外来入侵物种在入侵地经常面临很强的选择压力,这常常导致种内变异。在生活史事件发生的时间上,种群分化对气候梯度的响应被认为是促进许多入侵物种扩张的重要机制。对于产生种子的植物来说,种子萌发的时间决定了萌芽所经历的第一个环境条件,这对入侵植物的成功定植、种群建立和传播具有重要意义——尽管对于萌发在植物入侵成功中所起的作用仍然知之甚少。在本研究中,我们评估了入侵植物石茅(Sorghum halepense)在北美分布的10个种群的种子萌发在温度和降水梯度上的动态变化,以及这种变化是否与本地气候有关。种子被放置于一个较宽的温度范围(11-48˚C)以及两个水分梯度处理中。我们发现,石茅的种子可以在很宽的温度范围下萌发,但在不同的种群中,种子萌发的比例随温度和水分的不同而有较大的变化。有证据表明,当石茅的生长范围从温暖的气候扩展到寒冷的气候时,种子萌发温度的生态位也同时发生了向低温的转变。我们的研究结果表明,石茅种子的萌发已经适应了当地的气候,使其在整个生长范围内的萌发最大化,这可能是其入侵新环境的一个重要贡献因素。     

A multi-gene phylogeny of Cephalopoda supports convergent morphological evolution in association with multiple habitat shifts in the marine environment

ABSTRACT: BACKGROUND: The marine environment is comprised of numerous divergent organisms living under similar selective pressures, often resulting in the evolution of convergent structures such as the fusiform body shape of pelagic squids, fishes, and some marine mammals. However, little is known about the frequency of, and circumstances leading to, convergent evolution in the open ocean. Here, we present a comparative study of the molluscan class Cephalopoda, a marine group known to occupy habitats from the intertidal to the deep sea. Several lineages bear features that may coincide with a benthic or pelagic existence, making this a valuable group for testing hypotheses of correlated evolution. To test for convergence and correlation, we generate the most taxonomically comprehensive multi-gene phylogeny of cephalopods to date. We then create a character matrix of habitat type and morphological characters, which we use to infer ancestral character states and test for correlation between habitat and morphology. RESULTS: Our study utilizes a taxonomically well-sampled phylogeny to show convergent evolution in all six morphological characters we analyzed. Three of these characters also correlate with habitat. The presence of an autogenic photophore is correlated with a pelagic habitat, while the cornea and accessory nidamental gland correlate with a benthic lifestyle. Here, we present the first statistical tests for correlation between convergent traits and habitat in cephalopods to better understand the evolutionary history of characters that are adaptive in benthic or pelagic environments, respectively. DISCUSSION: Our study supports the hypothesis that habitat has influenced convergent evolution in the marine environment: benthic organisms tend to exhibit similar characteristics that confer protection from invasion by other benthic taxa, while pelagic organisms possess features that facilitate crypsis and communication in an environment lacking physical refuges. Features that have originated multiple times in distantly related lineages are likely adaptive for the organisms inhabiting a particular environment: studying the frequency and evolutionary history of such convergent characters can increase understanding of the underlying forces driving ecological and evolutionary transitions in the marine environment.     

Rescuing Robert Brown—The Origins of Angio-Ovuly in Seed Cones of Conifers

Robert Brown (1827) recognized the distinction between the two major classes of seed plants on the basis, respectively, of exposed ovules, which receive pollen directly, and enclosed ovules, which do not, at the time of pollination. The two groups, unfortunately, became known as “Gymnosperms” and “Angiosperms”, a distinction not made by Brown. The names are, at best, useful only as identifying labels. As is established here, gymno-ovuly and angio-ovuly would have been better words and should serve as the most consistent distinguishing character among groups of seed plants. Although conifers can hardly be claimed to be members of any lineage that led to the Angiosperms, the gymno-ovulate method of reproduction must have occurred in the direct ancestors of flowering plants. Interpolating development and functional evidence serves to resurrect Robert Brown’s important but neglected observation and correct misstatements implied in generally accepted systematic terminology. The result of this approach is to discover that the conifers in their reproductive diversity show some trends indicative of those characters that define the flowering plants. Although non-homologous these character manifestations provide useful models which can lead to a better understanding of the origin of “Angiosperms” from “Gymnosperms.” In the complex and extended process of reproductive development in both major groups regular transpositions occur, depending on whether “gymno-”and “angio-”are applied to ovules or seeds. Gymno-ovuly and angio-spermy are the most common conditions in conifers but result in intrinsically opposed processes of pollen reception and subsequent seed protection, which are resolved by successive changes in seed-cone structure. The modern conifers illustrate considerable diversity of ovulate and seed cone structure in relation to several biological functions, including traits that anticipate the angio-ovulate condition, in cone inception, ovule enclosure, pollen and embryo development and seed dispersal. From the paleobotanical record we do have a good grasp of the structural changes in the progressive modification of the coniferous seed cone in conifers, starting with the antecedent Carboniferous Cordaitales. These changes have involved the progressive condensation of a lax, branched strobilus and end with the highly condensed uniovulate “cone” as in many Podocarpus species in which the equivalent of a bitegmic ovule exists. The progressive changes can be perceived typologically among existing conifers but need to be understood as the result of biological processes that suggest increased reproductive efficiency, and can be added to our present quite robust understanding of conifer phylogeny.