Physiological mechanisms drive differing foliar calcium content in ferns and angiosperms

Recent evidence points to ferns containing significantly lower contents of foliar calcium and other cations than angiosperms. This is especially true of more ancient ‘non-polypod’ fern lineages, which predate the diversification of angiosperms. Calcium is an important plant nutrient, the lack of which can potentially slow plant growth and litter decomposition, and alter soil invertebrate communities. The physiological mechanisms limiting foliar calcium (Ca) content in ferns are unknown. While there is a lot we do not know about Ca uptake and transport in plants, three physiological processes are likely to be important. We measured transpiration rate, cation exchange capacity, and leaching loss to determine which process most strongly regulates foliar Ca content in a range of fern and co-occurring understory angiosperm species from a montane Hawaiian rainforest. We found higher instantaneous and lifetime (corrected for leaf lifespan) transpiration rates in angiosperms relative to ferns. Ferns preferentially incorporated Ca into leaves relative to strontium, which suggests that root or stem cation exchange capacity differs between ferns and angiosperms, potentially affecting calcium transport in plants. There were no differences in foliar Ca leaching loss between groups. Among the physiological mechanisms measured, foliar Ca was most strongly correlated with leaf-level transpiration rate and leaf lifespan. This suggests that inter-specific differences in a leaf’s lifetime transpiration may play a significant role in determining plant nutrition.     

Rapid nutrient cycling in leaf litter from invasive plants in Hawai’i

Physiological traits that contribute to the establishment and spread of invasive plant species could also have impacts on ecosystem processes. The traits prevalent in many invasive plants, such as high specific leaf areas, rapid growth rates, and elevated leaf nutrient concentrations, improve litter quality and should increase rates of decomposition and nutrient cycling. To test for these ecosystem impacts, we measured initial leaf litter properties, decomposition rates, and nutrient dynamics in 11 understory plants from the Hawaiian islands in control and nitrogen + phosphorus fertilized plots. These included five common native species, four of which were ferns, and six aggressive invasive species, including five angiosperms and one fern. We found a 50-fold variation in leaf litter decay rates, with natives decaying at rates of 0.2–2.3 year^–1 and invaders at 1.4–9.3 year^–1. This difference was driven by very low decomposition rates in native fern litter. Fertilization significantly increased the decay rates of leaf litter from two native and two invasive species. Most invasive litter types lost nitrogen and phosphorus more rapidly and in larger quantities than comparable native litter types. All litter types except three native ferns lost nitrogen after 100 days of decomposition, and all litter types except the most recalcitrant native ferns lost >50% of initial phosphorus by the end of the experiment (204–735 days). If invasive understory plants displace native species, nutrient cycling rates could increase dramatically due to rapid decomposition and nutrient release from invasive litter. Such changes are likely to cause a positive feedback to invasion in Hawaii because many invasive plants thrive on nutrient-rich soils.     

First insights into fern matK phylogeny

MatK, the only maturase gene in the land plant plastid genome, is a very popular phylogenetic marker that has been extensively applied in reconstructing angiosperm phylogeny. However, the use of matK in fern phylogeny is largely unknown, due to difficulties with amplification: ferns have lost the flanking trnK exons, typically the region used for designing stable priming sites. We developed primers that are either universal or lineage-specific that successfully amplify matK across all fern families. To evaluate whether matK is as powerful a phylogenetic marker in ferns as in angiosperms, we compared its sequence characteristics and phylogenetic performance to those of rbcL and atpA. Among these three genes, matK has the highest variability and substitution evenness, yet shows the least homoplasy. Most importantly, applying matK in fern phylogenetics better resolved relationships among families, especially within eupolypods I and II. Here we demonstrate the power of matK for fern phylogenetic reconstruction, as well as provide primers and extensive sequence data that will greatly facilitate future evolutionary studies of ferns.     

The potential for forest canopy litterfall interception by a dense fern understorey, and the consequences for litter decomposition

Most studies on litter decomposition have assumed that all falling plant litter reaches the ground where it then decomposes. In many forests a proportion of this litter may in fact be intercepted by understorey vegetation, but the ecological significance of this remains poorly understood. We performed two experiments in a temperate rainforest in southern New Zealand, in which there was a dense understorey of the crown fern Blechnum discolor. The fronds of this fern originate from a crown, and have a funnel‐like arrangement that can trap falling litter and prevent it from reaching the ground. The first experiment measured the effects of ferns on the spatial distribution of litter accumulation over one year. The ferns intercepted a substantial proportion of the total litterfall, and the fern crowns (from which the fronds originate) retained 10% of the total incoming litterfall (despite occupying only 2% of the ground area). The retained litter had a substantially higher ratio of twig to foliar litter than did the incoming litterfall. Further, much of the litter not retained on the crowns of the ferns accumulated at the base of the fern trunks. The second experiment considered litter decomposition in fern crowns versus the ground under the ferns. The litter that had accumulated in the crowns was characterized by higher microbial basal respiration and active microbial biomass than was the litter that had accumulated on the ground. The use of litterbags revealed that litter decomposition rates were significantly higher on the fern crowns than on the ground at 30 cm and 60 cm from the fern trunks. These results show that litter interception ameliorates the decomposer environment and increases the rate of litter decomposition. In total, this study provides evidence for understorey ferns greatly influencing both the spatial distribution of litterfall and the decomposition of plant litter. Although the ecological role of understorey vegetation in forested ecosystems has received little attention to date, our results point to understorey species as an important driver of forest ecosystem processes.     

Structure-function constraints of tracheid-based xylem: a comparison of conifers and ferns

The ferns comprise one of the most ancient tracheophytic plant lineages, and occupy habitats ranging from tundra to deserts and the equatorial tropics. Like their nearest relatives the conifers, modern ferns possess tracheid-based xylem but the structure-function relationships of fern xylem are poorly understood. Here, we sampled the fronds (megaphylls) of 16 species across the fern phylogeny, and examined the relationships among hydraulic transport, drought-induced cavitation resistance, the xylem anatomy of the stipe, and the gas-exchange response of the pinnae. For comparison, the results are presented alongside a similar suite of conifer data. Fern xylem is as resistant to cavitation as conifer xylem, but exhibits none of the hydraulic or structural trade-offs associated with resistance to cavitation. On a conduit diameter basis, fern xylem can exhibit greater hydraulic efficiency than conifer and angiosperm xylem. In ferns, wide and long tracheids compensate in part for the lack of secondary xylem and allow ferns to exhibit transport rates on a par with those of conifers. We suspect that it is the arrangement of the primary xylem, in addition to the intrinsic traits of the conduits themselves, that may help explain the broad range of cavitation resistance in ferns.     

Litterfall dynamics in a mixed conifer-angiosperm forest in northern New Zealand

Litterfall in a mixed conifer-angiosperm temperate forest in northern New Zealand was traced for 5 years to determine the patterns of litter production and turnover for conifer and angiosperm components of the forest. Basal area and above-ground biomass was shared approximately equally between conifer (mostly Agathis australis; New Zealand kauri) and angiosperm species (plus tree ferns). The five-year mean annual litterfall, excluding macro-litter, was 7.76± 0.39(SEM) t ha^?1 and ranged from 6.77±0.70 t ha^?1 in 1983–4 to 8.79±1.00 t ha^?1 in 1987–8. Mean monthly litterfall showed a strong seasonal pattern with low rates in winter and early spring, increasing to a peak in early autumn. There were major differences in the nature and timing of litterfall between the conifer and angiosperm fractions. Angiosperm leaf litter reached a maximum in early summer, while conifer litterfall showed highest rates for leaves, twigs and cone scales in late summer-autumn. Conifer reproductive structures (strobili and cone scales) contributed from 13 to 21% of total litterfall, a value high relative to other temperate forests. However, conifer leaf turnover was low relative to that for the angiosperms. Size of the microlitter store was 16.16±1.97 t ha^?1 prior to conifer cone fall, and 18.70±2.02 t ha^?1 following it, and conifer litter made up 76–78% of the total litter store. The estimated mean annual decomposition constant, k, was 0.39 overall, 0.33 for conifer leaf litter and 0.71 for angiosperm leaf litter, values which agree well with previously published rates for decomposition in this forest stand. Differences in the costs of biomass production and rates of turnover, as measured by litterfall and decomposition, may help to explain the functional coexistence of conifers and angiosperms in mixed forests.     

Recurrent genome duplication events likely contributed to both the ancient and recent rise of ferns

Ferns, the second largest group of vascular plants, originated ～400 mil ion years ago(Mya). They became dominant in the ancient Earth landscape before the angiosperms and are stil important in current ecosystems.Many ferns have exceptional y high chromosome numbers,possibly resulting from whole-genome duplications(WGDs).However, WGDs have not been investigated molecularly across fern diversity. Here we detected and dated fern WGDs using a phylogenomic approach and by calculating synonymous substitution rates(Ks). We also investigated a possible correlation between proposed WGDs and shifts in species diversification rates. We identified 19 WGDs: three ancient events along the fern phylogenetic backbone that are shared by 66%–97% of extant ferns, with additional lineage-specific WGDs for eight orders, providing strongevidence for recurring genome duplications across fern evolutionary history. We also observed similar Ks peak values for more than half of these WGDs, with multiple WGDs occurring close to the Cretaceous(～145–66 Mya). Despite the repeated WGD events, the biodiversity of ferns declined during the Cretaceous, implying that other factors probably contributed to the floristic turnover from ferns to angiosperms. This study provides molecular evidence for recurring WGDs in ferns and offers important clues to the genomic evolutionary history of ferns.     

Leaf evolution in Southern Hemisphere conifers tracks the angiosperm ecological radiation

The angiosperm radiation has been linked to sharp declines in gymnosperm diversity and the virtual elimination of conifers from the tropics. The conifer family Podocarpaceae stands as an exception with highest species diversity in wet equatorial forests. It has been hypothesized that efficient light harvesting by the highly flattened leaves of several podocarp genera facilitates persistence with canopy-forming angiosperms, and the angiosperm ecological radiation may have preferentially favoured the diversification of these lineages. To test these ideas, we develop a molecular phylogeny for Podocarpaceae using Bayesian-relaxed clock methods incorporating fossil time constraints. We find several independent origins of flattened foliage types, and that these lineages have diversified predominantly through the Cenozoic and therefore among canopy-forming angiosperms. The onset of sustained foliage flattening podocarp diversification is coincident with a declining diversification rate of scale/needle-leaved lineages and also with ecological and climatic transformations linked to angiosperm foliar evolution. We demonstrate that climatic range evolution is contingent on the underlying state for leaf morphology. Taken together, our findings imply that as angiosperms came to dominate most terrestrial ecosystems, competitive interactions at the foliar level have profoundly shaped podocarp geography and as a consequence, rates of lineage diversification.     

The angiosperm radiation revisited, an ecological explanation for Darwin's 'abominable mystery'

One of the greatest terrestrial radiations is the diversification of the flowering plants (Angiospermae) in the Cretaceous period. Early angiosperms appear to have been limited to disturbed, aquatic or extremely dry sites, suggesting that they were suppressed in most other places by the gymnosperms that still dominated the plant world. However, fossil evidence suggests that by the end of the Cretaceous the angiosperms had spectacularly taken over the dominant position from the gymnosperms around the globe. Here, we suggest an ecological explanation for their escape from their subordinate position relative to gymnosperms and ferns. We propose that angiosperms due to their higher growth rates profit more rapidly from increased nutrient supply than gymnosperms, whereas at the same time angiosperms promote soil nutrient release by producing litter that is more easily decomposed. This positive feedback may have resulted in a runaway process once angiosperms had reached a certain abundance. Evidence for the possibility of such a critical transition to angiosperm dominance comes from recent work on large scale vegetation shifts, linking long-term field observations, large scale experiments and the use of simulation models.     

Structure and function at the root apex - phylogenetic and ontogenetic perspectives on apical cells and quiescent centres

The summit of roots of various plant species may be occupied by a single, rapidly proliferating tetrahedral apical cell (as in ferns), or by a multicellular and slowly proliferating quiescent centre (as in angiosperms), or by intermediate types of cellular organizations. The present paper attempts to deduce the phylogeny of these various types of cellular patterning at the root apex.     

Early stages of root and leaf decomposition in Hawaiian forests: effects of nutrient availability

We examined the effects of soil nutrient availability and tissue chemistry on decomposition of both fine roots (<2 mm diameter) and leaves in three sites along a forest chronosequence in the Hawaiian Islands. These sites form a natural fertility gradient, with the youngest and oldest sites having lower nutrient availability than the intermediate-aged site. Nitrogen (N) limits aboveground net primary productivity (ANPP) in the youngest site, while phosphorus (P) limits ANPP in the oldest site. Both root and leaf litter decomposed most slowly in the 4.1-Myear-old site. We also investigated root decomposition in fertilized plots at the youngest and oldest sites; when roots were produced and decomposed in fertilized plots, root decomposition rates increased with N and P additions at the 4.1-Myear-old site. At the 300-year-old site, however, root decomposition rates did not respond to N or P additions. Roots decomposed faster than leaves at the more infertile sites, in part because of lower lignin-to-nitrogen ratios in roots than in leaf litter. Decomposing roots immobilized more nutrients than did decomposing leaves, and may serve an important role in retaining nutrients in these forests. Received: 30 November 1998 / Accepted: 12 August 1999     

Amazonian Dark Earth Shapes the Understory Plant Community in a Bolivian Forest

Amazonian Dark Earths (ADE) are the result of human modification of the Amazonian landscape since pre‐Columbian times. ADE are characterized by increased soil fertility compared to natural soils. In the Amazonian forest, soil fertility influences understory herb and fern species composition. However, little research has been done to evaluate the effect of ADE on the composition of the understory community. We evaluated the effects of ADE and soil in 36 plots (150 m × 4 m) established in a Bolivian moist forest (La Chonta). For each plot, we determined soil nutrients, and the composition, richness, and abundance of terrestrial fern, angiosperm herb, and understory palm species. We found that the presence of ADE created a gradient in soil nutrients and pH that affected the understory species composition especially of ferns and palms. Additionally, the higher nutrient concentration and more neutral soil pH on ADE soils caused a decrease of ferns species richness. We therefore conclude that the current composition of the understory community in this particular Bolivian forest is a reflection of past human modifications of the soil.     

An invasive tree fern alters soil and plant nutrient dynamics in Hawaii

Invasive species that alter ecosystems are often successful competitors due to their effects on nutrient cycling. Sphaeropteris cooperi (Cyatheaceae; Australian tree fern) has been invading intact Hawaiian rainforests for decades and displacing the dominant native tree fern Cibotium glaucum (Cibotiaceae). S. cooperi produces more leaves that grow faster, contain more N and P, and decompose faster than C. glaucum leaves. Our experiment tested the effects of additions of leaf litter from native and non-native tree ferns on the growth and nutrient content of four native angiosperm species in forest (N-rich) and landslide (P-rich) soils. Both litter treatments inhibited growth initially in all species, but subsequent responses were species-specific. Compared to control treatments, the increase in biomass was highest in the fast-growing Carex wahuensis and Hibiscus arnottianus with S. cooperi litter in landslide soil. Leaf N in C. wahuensis was higher with S. cooperi litter and in forest soil, as expected, but other leaf nutrient responses showed some evidence of nutrient immobilization from litter addition. Several growth measures were higher with S. cooperi than C. glaucum litter and in forest than landslide soil, suggesting that N availability is the strongest driver of growth. Our results show that S. cooperi can alter nutrient cycling in Hawaiian plants, sometimes with positive effects on growth. However, under natural conditions, native plants must compete for these additional nutrients with S. cooperi and other invasive species. This study contributes to invasion biology as the first to examine the impact of leaf litter of an invasive fern on native species.     

Highly conserved low-copy nuclear genes as effective markers for phylogenetic analyses in angiosperms

? Organismal phylogeny provides a crucial evolutionary framework for many studies and the angiosperm phylogeny has been greatly improved recently, largely using organellar and rDNA genes. However, low-copy protein-coding nuclear genes have not been widely used on a large scale in spite of the advantages of their biparental inheritance and vast number of choices. ? Here, we identified 1083 highly conserved low-copy nuclear genes by genome comparison. Furthermore, we demonstrated the use of five nuclear genes in 91 angiosperms representing 46 orders (73% of orders) and three gymnosperms as outgroups for a highly resolved phylogeny. ? These nuclear genes are easy to clone and align, and more phylogenetically informative than widely used organellar genes. The angiosperm phylogeny reconstructed using these genes was largely congruent with previous ones mainly inferred from organellar genes. Intriguingly, several new placements were uncovered for some groups, including those among the rosids, the asterids, and between the eudicots and several basal angiosperm groups. ? These conserved universal nuclear genes have several inherent qualities enabling them to be good markers for reconstructing angiosperm phylogeny, even eukaryotic relationships, further providing new insights into the evolutionary history of angiosperms.     

Positive feedbacks between decomposition and soil nitrogen availability along fertility gradients

Background and aims

We determined the relationship between site N supply and decomposition rates with respect to controls exerted by environment, litter chemistry, and fungal colonization.

Methods

Two reciprocal transplant decomposition experiments were established, one in each of two long-term experiments in oak woodlands in Minnesota, USA: a fire frequency/vegetation gradient, along which soil N availability varies markedly, and a long-term N fertilization experiment. Both experiments used native Quercus ellipsoidalis E.J. Hill and Andropogon gerardii Vitman leaf litter and either root litter or wooden dowels.

Results

Leaf litter decay rates generally increased with soil N availability in both experiments while belowground litter decayed more slowly with increasing soil N. Litter chemistry differed among litter types, and these differences had significant effects on belowground (but not aboveground) decay rates and on aboveground litter N dynamics during decomposition. Fungal colonization of detritus was positively correlated with soil fertility and decay rates.

Conclusions

Higher soil fertility associated with low fire frequency was associated with greater leaf litter production, higher rates of fungal colonization of detritus, more rapid leaf litter decomposition rates, and greater N release in the root litter, all of which likely enhance soil fertility. During decomposition, both greater mass loss and litter N release provide mechanisms through which the plant and decomposer communities provide positive feedbacks to soil fertility as ultimately driven by decreasing fire frequency in N-limited soils and vice versa.     

Species control variation in litter decomposition in a pine forest exposed to elevated CO2

Net primary production and the flux of dry matter and nutrients from vegetation to soils has increased following four years of exposure to elevated CO₂ in a southern pine forest in NC, USA. This has increased the demand for nutrients to support enhanced rates of NPP and altered the conditions for litter decomposition on the forest floor. We quantified the chemistry and decomposition dynamics of leaf litter produced by five of the most abundant tree species in this ecosystem during the third and fourth growing seasons under elevated CO₂. The objectives of this study were to determine (i) if there were systemic or species‐specific changes in leaf litter chemistry associated with a sustained enhancement of plant growth under elevated CO₂; and (ii) whether the process of litter decomposition was altered by increased inputs of energy and nutrients to the forest floor in the plots under elevated CO₂. Leaf litter chemistry, including various C fractions and N concentration, was virtually unchanged by elevated CO₂. With few exceptions, plant litter produced under elevated CO₂ lost mass or N at the same relative rate as that produced under ambient CO₂. The relationship between initial litter chemistry and decomposition was not altered by elevated CO₂. The greater forest floor mass and nutrient content in the plots under elevated CO₂ had no consistent or long‐term effect on litter decomposition. Thus, we found no evidence that plant and microbial processes under elevated CO₂ resulted in systemic changes in mass loss or N dynamics during decomposition. In contrast to the limited effects of elevated CO₂ on litter chemistry and decomposition, there were large differences among species in initial litter chemistry, mass loss and N dynamics during decomposition. If the species composition of this forest community is altered by elevated CO₂, the indirect effect of a change in species composition will exert greater control over the long‐term rate of nutrient cycling than the direct effect of elevated CO₂ on litter chemistry and decomposition dynamics alone.     

Leaf energy balance modelling as a tool to infer habitat preference in the early angiosperms

Despite more than a century of research, some key aspects of habitat preference and ecology of the earliest angiosperms remain poorly constrained. Proposed growth ecology has varied from opportunistic weedy species growing in full sun to slow-growing species limited to the shaded understorey of gymnosperm forests. Evidence suggests that the earliest angiosperms possessed low transpiration rates: gas exchange rates for extant basal angiosperms are low, as are the reconstructed gas exchange rates for the oldest known angiosperm leaf fossils. Leaves with low transpirational capacity are vulnerable to overheating in full sun, favouring the hypothesis that early angiosperms were limited to the shaded understorey. Here, modelled leaf temperatures are used to examine the thermal tolerance of some of the earliest angiosperms. Our results indicate that small leaf size could have mitigated the low transpirational cooling capacity of many early angiosperms, enabling many species to survive in full sun. We propose that during the earliest phases of the angiosperm leaf record, angiosperms may not have been limited to the understorey, and that some species were able to compete with ferns and gymnosperms in both shaded and sunny habitats, especially in the absence of competition from more rapidly growing and transpiring advanced lineages of angiosperms.     

Are functional traits and litter decomposability coordinated across leaves,twigs and wood? A test using temperate rainforest tree species

Synthesis This study compared the decomposability of leaf, twig and wood litter from 27 co‐occurring temperate rainforest tree species in New Zealand. We found that interspecific variation in decomposition was not coordinated across the three litter types. Analysis of the relationships between functional traits and decomposition revealed that traits predictive of wood decomposition varied among the species independently from traits predictive of the decomposition of leaf and twig litter. We conclude that efforts to understand how tree species influence C, N and P dynamics in forested ecosystems through the decomposition pathway need to consider the functional traits of multiple plant structures. Plant functional traits are increasingly used to evaluate changes in ecological and ecosystem processes. However our understanding of how functional traits coordinate across different plant structures, and the implications for trait‐driven processes such as litter decomposition, remains limited. We compared the functional traits of green leaves and leaf, twig and wood litter among 27 co‐occurring tree species from New Zealand, and quantified the loss of mass, N and P from the three litter types during decomposition. We hypothesised that: a) the functional traits of green leaves, and leaf, twig and wood litter are co‐ordinated so that species which produce high quality leaves and leaf litter will also produce high quality twig and wood litter, and b) the decomposability of leaf, twig and wood litter is coordinated because breakdown of all three litter types is driven by similar combinations of traits. Trait variation across species was co‐ordinated between leaves, twigs and wood when angiosperm and gymnosperm species were considered in combination, or when angiosperms were considered separately, but trait coordination was poor for gymnosperms. There was little coordination among the three litter types in their decomposability, especially when angiosperms and gymnosperms were considered separately; this was caused by the decomposability of each of the three litter types, at least partially, being driven by different functional traits or trait combinations. Our findings indicate that although interspecific variation in the functional traits of trees can be coordinated among leaves, twigs and wood, different or unrelated traits predict the decomposition of these different structures. Furthermore, leaf‐level analyses of functional traits are not satisfactory proxies for function of whole trees and related ecological processes. As such, efforts to understand how tree species influence C, N and P dynamics in forested ecosystems through the decomposition pathway need to consider functional traits of other plant structures.     

Does initial litter chemistry explain litter mixture effects on decomposition?

Abstract We tested the hypothesis that interactions in litter mixtures (expressed as the difference between observed and expected decomposition rates) are greater when the component species differ more in their initial litter chemistry. Thereto, we collected freshly senesced leaf litter from a wide range of species from an old field and woodland vegetation, and a fen ecosystem in The Netherlands. Litterbags with either mono-specific litter (20 and 15 species), or litter mixtures (50 and 42 species pairs) of randomly drawn combinations of two representatives from different plant functional types were incubated for 20, 35 and 54 weeks in a purpose-built decomposition bed (woodland/old field) or in the field (fen). Species showed a wide range of decomposition rates. For the woodland/old field species, initial litter C and P concentrations were significantly correlated with litter decomposition rate. For the fen species, litter phenolics concentration was correlated with decomposition rate. If the Sphagnum species were left out of the analyses, initial litter P and phenolics concentration both showed a significant relationship, albeit only with the remaining mass after 1 year. Differences between observed and expected decomposition were often considerable in individual litter mixtures. Regression analysis showed that such differences were not related to the differences in litter chemistry of the component species. Furthermore, litter mixtures containing species with very different initial litter chemistry did not show stronger interaction when tested against litter mixtures containing chemically similar litter types. From these observations we conclude that the difference in initial single litter chemistry parameters of the component is not a useful concept to explain interactions in litter mixtures.Due to an error in the citation line, this revised PDF (published in December 2003) deviates from the printed version, and is the correct and authoritative version of the paper.     

Early angiosperm ecology: evidence from the Albian-Cenomanian of Europe

BACKGROUND AND AIMS: The mid-Cretaceous is a period of sudden turnover from gymnosperm to angiosperm-dominated floras. The aim was to investigate the fossil plant ecology in order to follow the spread of angiosperm taxa. METHODS: Floristic lists and localities from the latest Albian-Cenomanian of Europe are analysed with Wagner's Parsimony Method, a clustering method currently used in phylogeny (cladistics). KEY RESULTS: Wagner's Parsimony Method points out that (a) gymnosperms dominated brackish water-related environments while angiosperms dominated freshwater-related environments (e.g. swamps, floodplains, levees, channels), (b) angiosperms showed the highest diversity in stable, freshwater-related environments, (c) a single angiosperm, 'Diospyros' cretacea, is restricted to brackish water-related environments and (d) the families Lauraceae and Platanaceae were exclusive to disturbed, braided river environments, implying a opportunist strategy for early tree angiosperms. CONCLUSIONS: During the Mid-Cretaceous, European floras were characterized by (a) coastal gymnosperms, (b) highly diversified fluvial angiosperms and (c) the first European brackish water-related angiosperm.