Density Effect on the Fruit-set, Seed Crop Viability and Seedling Vigour of Abies pinsapo

Variation in fruit-set, cone crop, seed viability and seedlingvigour ofAbies pinsapoBoiss. were recorded at two differentstand densities during four consecutive years (1990–1993).In this period only two cone crops were observed (1991 and 1992).Extreme differences were found in response to density. In thelow density stand trees had a greater seed-cone production andfruit-set in each of the two crop years. Also, significant differenceswere recorded in the seed crop viability: in the low densityzone average seed viability was only 21% whereas in the highdensity one it was 82%. Other important differences such ascone size, number of seeds per cone and seedling vigour wererecorded at different densities. The results suggest that bothpollen availability and pollen origin in low density standsinfluence the frequency and quality of filled seeds. The resultshave practical implications for the management ofA. pinsapoforseed production and subsequent reforestation. Abies pinsapo; inbreeding; mating system; plant density; cone crop; seed viability; seedling vigour; Mediterranean fir     

Dendroecological and genetic insights for future management of an old-planted forest of the endangered Mediterranean fir Abies pinsapo

Pinsapo fir (Abies pinsapo Boiss.) is an endangered Mediterranean conifer that has raised strong conservation interest as a paradigmatic example of species characterized by small and fragmented populations. We studied an old reforestation stand composed of A. pinsapo, Pinus nigra and Pinus sylvestris established in the 1910s in central-eastern Spain (about 500 km north of the species native distribution range), with the aim of evaluating the stand’s suitability as an ex situ conservation unit for the fir. To this end, we investigated the long-term performance of the stand and assessed genetic diversity of A. pinsapo. Tree-ring width (TRW) and carbon isotope discrimination (Δ¹³C) were used to characterise growth dynamics and intrinsic water-use efficiency (WUE_i), respectively. Furthermore, 42 pinsapo firs were genotyped at five microsatellite loci to compare their genetic variation with published data on natural populations. A. pinsapo showed ca. two-fold higher radial growth than pines in the last 80 years; however, a growth decrease was observed for all species from the 1990s onwards. Indexed TRW was positively associated with Δ¹³C at the species level, denoting inter-annual growth dependence on water availability. Overall, Δ¹³C was higher for A. pinsapo compared to pines, indicating lower WUE_i, but Δ¹³C significantly decreased over the last 50 years for all species, likely as the result of tighter stomatal regulation of water loss, resulting in WUE_i increases of about 25 %. Recently, however, A. pinsapo showed reduced WUE_i increase in concord with growth slowdown, suggesting a threshold response for stomatal regulation. Although genetic diversity of A. pinsapo was about half of natural populations, the old-planted stand could be important for the conservation of this endemic species considering its good long-term growth and physiology. The latest decrease in performance of A. pinsapo, however, asks for urgent management measures aimed at reducing the competition for water and promoting growth and natural regeneration. This study illustrates the potential of combining tree-ring-based long-term physiological information with genetic data to ascertain the prospects of artificial stands for conservation purposes.     

Interacting effects of changes in climate and forest cover on mortality and growth of the southernmost European fir forests

Aims The combined effects of changes in climate and land use on tree mortality and growth patterns have rarely been addressed. Relict tree species from the Mediterranean Basin serve as appropriate models to investigate these effects, since they grow in climatically stressed areas which have undergone intense cover changes. The aim is to use climate, aerial photographs, stand structure and radial‐growth data to explain the mortality and historical patterns of growth of Abies pinsapo in the area where this relict species was first protected. Location Sierra de las Nieves, West Baetic Range, southern Spain. Methods We assessed variations of tree cover in A. pinsapo forests through image analyses of aerial photographs spanning the last 50 years. We sampled 31 stands to assess current altitudinal patterns of forest structure and mortality. We evaluated the relationships between radial growth and regional climate using linear models in three sites at different elevations. Results Regional warming and a decrease in precipitation were detected. Forest tree cover increased at all elevations from 1957 until 1991, but it afterwards decreased below 1100 m. Currently, the likelihood of tree mortality increases downwards and is associated with dense, closed stands with a low living basal area. In contrast to previous droughts, a sharp synchronized reduction in tree growth, not fully accounted for in linear climate–growth models, occurred at low elevations in 1994–95, but not upwards. It was preceded by a weakening of the negative association between low‐elevation growth and water deficit since the late 1970s. Conclusions The intense densification of A. pinsapo forests following strict protection measures in the late 1950s enhanced the vulnerability of climate‐sensitive A. pinsapo forests to recent drier conditions. Such abrupt land‐use changes help to explain recent patterns of mortality and growth decline in low‐elevation A. pinsapo forests.     

Larval feeding behaviour affects the impact of staminate flower production on the suitability of balsam fir trees for spruce budworm

Laboratory rearing of spruce budworm, Choristoneura fumiferana, in conjunction with field rearing indicated that the feeding behaviour of the larvae, which is affected by the insect population density, significantly influenced the impact of balsam fir, Abies balsamea, staminate flowering on spruce budworm biology. At low budworm density, the production of pollen in the midcrown of host trees reduced the insect development time by 5 days without affecting pupal weight, fecundity and survival. However, at high budworm density, the small amount of current-year foliage produced by flowering branches forced old larvae (sixth instar) either to feed on 1-year-old foliage (backfeeding) or to move from the midcrown to the lower crown section where staminate flowers are absent and more current-year foliage is available. When old larvae fed on old foliage, they exhibited reduced pupal weight and fecundity without losing the advange in development time that they obtained from feeding on pollen during their early stages of development. On the other hand, when old larvae moved to the lower crown section, they avoided the negative effects of backfeeding but lost the advantage in development time that was gained from feeding on pollen. Results from this study indicated that the production of staminate flowers by balsam fir trees could have opposite effects on spruce budworm population dynamics depending upon the insect population density when flowering occurs.     

Specific features of meiosis in the Siberian Fir (<Emphasis Type="Italic">Abies sibirica</Emphasis>) in the forest Arboretum of the V. N. Sukachev Institute,Russia

Investigating the tolerance of plant reproductive systems to environmental changes has become a research priority under current climate change scenarios. Successful plant conservation requires knowledge of plant reproductive biology, particularly the meiotic characteristics of planted species. Meiosis, as part of microsporogenesis, is a critical plant developmental stage controlling future pollen quality. Meiosis in a Siberian fir (Abies sibirica) plantation, established in the Forest Arboretum of the Sukachev Institute, Russia, was studied from 2002 to 2004. The microsporogenesis pattern found for the Siberian fir appeared to be largely similar to that exhibited by other conifer species. Meiosis in the Siberian fir has the following characteristics: asynchrony, rapid progression of telophases I and II, and parallel and linear spindle arrangements at different meiosis II stages. General and specific meiosis irregularities were recorded at each stage. Some specific features of meiosis and the specific development of some irregularities were revealed. Pollen development analysis showed that irregular pollen grains made up less than 1% of all grains. The specific features of meiosis identified in fir trees growing in the Arboretum indicated low resistance of male reproductive structures to climatic changes and might account for high fir pollen sterility in this new environment.     

Changes in gender expression in korean populations of<Emphasis Type="Italic">Pinus densiflora</Emphasis> over a five-year period

Male and female cone crop patterns and gender expression inPinus densiflora were examined in three adjacent Korean populations over four to five years. Three parameters — cone ratio, sexual system, and standardized phenotypic gender — were assessed according to cone production. Both male and female cone crops and cone ratios varied by year and by site. After controlling for plant size, population differences in cone crops and cone ratios became stronger. At all three sites, the proportion of trees with differing sexual systems tended to fluctuate among years. Overall, 37% of all tagged trees changed their sexual system at least once, mostly ranging between monoecy and male. These particular trees were relatively small and exhibited slow radial growth compared with the non-sex-changing trees, suggesting a trade-off between vegetative and reproductive growth. Despite annual variations in their standardized phenotypic genders, the rankings of trees remained concordant between successive years at all sites. These results indicate that, inP. densiflora, male and female cone crops and, consequently, gender expression, may be affected by diverse factors that include plant size, trade-offs between vegetative and reproductive growth, and genetic components.     

Stand-structural effects on Heterobasidion abietinum-related mortality following drought events in Abies pinsapo

Climate change may affect tree–pathogen interactions. This possibility has important implications for drought-prone forests, where stand dynamics and disease pathogenicity are especially sensitive to climatic stress. In addition, stand structural attributes including density-dependent tree-to-tree competition may modulate the stands’ resistance to drought events and pathogen outbreaks. To assess the effects of stand structure on root-rot-related mortality after severe droughts, we focused on Heterobasidion abietinum mortality in relict Spanish stands of Abies pinsapo, a drought-sensitive fir. We compared stand attributes and tree spatial patterns in three plots with H. abietinum root-rot disease and three plots without root-rot. Point-pattern analyses were used to investigate the scale and extent of mortality patterns and to test hypotheses related to the spread of the disease. Dendrochronology was used to date the year of death and to assess the association between droughts and growth decline. We applied a structural equation modelling approach to test if tree mortality occurs more rapidly than predicted by a simple distance model when trees are subjected to high tree-to-tree competition and following drought events. Contrary to expectations of drought mortality, the effect of precipitation on the year of death was strong and negative, indicating that a period of high precipitation induced an earlier tree death. Competition intensity, related to the size and density of neighbour trees, also induced an earlier tree death. The effect of distance to the disease focus was negligible except in combination with intensive competition. Our results indicate that infected trees have decreased ability to withstand drought stress, and demonstrate that tree-to-tree competition and fungal infection act as predisposing factors of forest decline and mortality.     

Environmental conditions affect sex expression in monoecious, but not in male and female plants of Urtica dioica

Urtica dioica is a sub-dioecious plant species, i.e. males and females coexist with monoecious individuals. Under standard conditions, seed sex ratio (SSR, fraction of males) was found to vary significantly among seed samples collected from female plants originating from the same population (0.05–0.76). As a first step, we investigated the extent to which SSR and sex expression of male, female, and monoecious individuals is influenced by external factors. We performed experiments to analyse: (1) whether the environment of a parental plant affects the sex ratio (SR) of its offspring, (2) whether SSR can be affected by environmental conditions before flowering, and (3) whether sex expression of male, female and monoecious plants that have already flowered can be modified by environmental conditions or by application of phyto-hormones. Within the range of our experimental design, SSR was not influenced by external factors, and gender in male and female plants was stable. However, sex expression in monoecious plants was found to be labile: flower sex ratio (FSR, fraction of male flowers) differed considerably between clones from the same individual within treatments, and increased toward 100% maleness under benign conditions. These results provide strong evidence that monoecious individuals are inconstant males, which alter FSR according to environmental circumstances. In contrast, we consider sex expression in male and female individuals to be solely genetically based. The observed variation in SSR between maternal parents cannot be explained by sex-by-environment interactions.     

Reproductive biology of Abies cephalonica Loudon in Mount Aenos National Park, Cephalonia, Greece

The reproductive biology of the endemic to Greece Abies cephalonica Loudon, including the phenology of the reproductive life cycle, cone production in relation to plant age and the required seed germination conditions, was studied. All individuals growing within 20 permanent plots of 100 m² each established in 11 different locations covering the entire strictly protected area of Mount Aenos National Park (Cephalonia) were monitored over a period of 4 years. The cones are formed on the upper part of the previous year’s branches, mostly at the upper crown half of the reproductive individuals (over 53 years old). Female flowers are pollinated in spring and cone maturation lasts until the beginning of autumn, when seed dispersal occurs. The annual production of cones per individual varied significantly, revealing masting behavior for this tree. This behavior was also expressed in the number of trees producing cones, as well as the percentage of sound seeds per cone. A significant difference in the mass and the length of the cones between the years of low and high cone production was observed, being higher in the years of massive cone production. Similar germination percentages were observed in full darkness or under “canopy light” and “sun light”, provided that the seeds were previously stratified for 6 weeks. It is concluded that A. cephalonica exhibits plasticity expressed in its reproductive behavior for alternating years of high to low cone production and in its seed germination for an array of habitat light conditions.     

Competition and drought limit the response of water-use efficiency to rising atmospheric carbon dioxide in the Mediterranean fir <Emphasis Type="Italic">Abies pinsapo</Emphasis>

The gas-exchange and radial growth responses of conifer forests to climatic warming and increasing atmospheric CO₂ have been widely studied. However, the modulating effects of variables related to stand structure (e.g., tree-to-tree competition) on those responses are poorly explored. The basal-area increment (BAI) and C isotope discrimination (C stable isotope ratio; δ¹³C) in the Mediterranean fir Abies pinsapo were investigated to elucidate the influences of stand competition, atmospheric CO₂ concentrations and climate on intrinsic water-use efficiency (WUEi). We assessed the variation in δ¹³C of tree-rings from dominant or co-dominant trees subjected to different degrees of competition. A high- (H) and a low-elevation (L) population with contrasting climatic constraints were studied in southern Spain. Both populations showed an increase in long-term WUEi. However, this increase occurred more slowly at the L site, where a decline of BAI was also observed. Local warming and severe droughts have occurred in the study area over the past 30 years, which have reduced water availability more at lower elevations. Contrastingly, trees from the H site were able to maintain high BAI values at a lower cost in terms of water consumption. In each population, trees subjected to a higher degree of competition by neighboring trees showed lower BAI and WUEi than those subjected to less competition, although the slopes of the temporal trends in WUEi were independent of the competitive micro-environment experienced by the trees. The results are consistent with an increasing drought-induced limitation of BAI and a decreasing rate of WUEi improvement in low-elevation A. pinsapo forests. This relict species might not be able to mitigate the negative effects of a decrease in water availability through a reduction in stomatal conductance, thus leading to a growth decline in the more xeric sites. An intense and poorly asymmetric competitive environment at the stand level may also act as an important constraint on the adaptive capacity of these drought-sensitive forests to climatic warming. Electronic supplementary material  The online version of this article (doi:) contains supplementary material, which is available to authorized users.     

Impact of balsam fir flowering on pollen and foliage biochemistry in relation to spruce budworm growth, development and food utilization

The impact of balsam fir (Abies balsamea (L.) Miller) flowering on nutritional and allelochemical quality of pollen, current-year and one-year-old foliage is studied in relation to spruce budworm (Choristoneura fumiferana Clem.) (Lepidoptera: Tortricidae) growth, development and utilization of food and nitrogen. In the laboratory, using fresh food from the field, we simulated conditions of low larval population density, in which there is no current-year foliage depletion during the spruce budworm feeding period. Similarly, we simulated conditions of high larval population density when current-year foliage depletion occurs.Because of the high nutritive value of pollen (high amounts of amino acids and minerals, especially nitrogen; low monoterpene content), insects from flowering trees reached the fifth instar five days earlier than those from non-flowering trees, and had heavier dry- and nitrogen-weights at the beginning of the fifth instar. At budbreak, switching from pollen to current-year foliage negatively affected conversion efficiencies and digestibilities of food and nitrogen (AD; ADN; ECDN; ECI; ECIN). The switch from pollen to new foliage had a detrimental impact on fifth-instar survival and on newly-moulted sixth-instar dry- and nitrogen-weights. Moreover, during the fifth instar, balsam fir flowering reduced the nutritive value of current-year foliage, which in turn, might have contributed to the reduced larval growth. Nevertheless, during the sixth instar, balsam fir flowering affected the biochemistry of current-year foliage in ways that enabled larvae to compensate for their low fifth-instar biological performance; larvae also managed to reach pupal dry weight similar to larvae reared on non-flowering trees. Current-year foliage from flowering trees contained less nitrogen, total soluble sugars and total monoterpenes. Those foliar characteristics enabled larvae to increase food and nitrogen consumption rates (RCR; RNCR), because of lower repellency and/or post-ingestional feedback from monoterpenes.As for current-year foliage, balsam fir flowering reduced nitrogen, total soluble sugar and total monoterpene contents in one-year-old foliage during the sixth-instar feeding period. These characteristics enabled sixth-instar larvae, fed on old foliage from flowering trees, to have high relative food and nitrogen consumption rates (RCR; RNCR). Larvae were also able to reach higher relative growth rates (RGR) and relative nitrogen accumulation rates (RNAR) compared to larvae reared on one-year-old foliage from non-flowering trees. Finally, larvae on flowering trees had pupal dry weight similar to those from non-flowering trees, but reached the adult stage nine days earlier.Regardless the foliage type consumed by spruce budworm larvae during the sixth instar, pollen consumption during early larval stages reduced total development time, and thus exposure time to natural enemies. This phenomenon might increase larval survival. Balsam fir flowering changed the biochemistry of one-year-old and current-year foliages, but did not affect pupal dry weights of larvae reared on flowering trees compared to those reared on non-flowering trees. Thus, at low population density, spruce budworm populations in balsam fir flowering stands might be favoured over those in balsam fir non-flowering stands. In addition, when larvae consumed one-year-old foliage during the entire sixth instar, those on flowering trees are probably favoured over those on non-flowering trees. However, because flowering trees produce less new foliage than non-flowering trees, current-year foliage depletion may occur earlier on flowering trees than on non-flowering trees. Thus, at similar larval population density, larvae on flowering trees might have to feed on one-year-old foliage earlier than those on non-flowering trees. In that case, spruce budworm populations on non-flowering stands would be favoured over those on flowering stands.     

Gender specialization and factors affecting fruit set of the wind-pollinated heterodichogamous Juglans regia

Gender expression, flowering phenology, reproductive performance and factors affecting fruit set (i.e., flowering synchrony, size and distance to the nearest pollen donor) were investigated in a cultivated population of a wind-pollinated self-compatible heterodichogamous Juglans regia (Juglandaceae). Four flowering morphs, (i.e., protandrous, protogynous, male and female) were observed. The sexual functions of the protandrous and protogynous morphs were almost synchronous; however, they were not reciprocal, and the separation of male and female flowering within most monoecious individuals was not complete. Thus, within-morph mating and geitonogamous pollination may be common. The ratio of protandrous versus protogynous morphs was biased towards the protandrous morph, but the fruit set did not differ between the morphs, suggesting that the fruit set of the protandrous morph could be partly compensated by within-morph pollination. The ratio of the female flower number or fruit number to the total male catkin length was higher in the protogynous morph than in the protandrous morph and did not vary with plant size, suggesting that gender variation was not size dependent and that the sexual function of protandrous morphs was more male biased. Fruit set depended on plant size only for protandrous morphs. The fruit set of individual plants decreased with increasing distance to the nearest pollen donor regardless of morph, possibly because of pollen limitation. The fruit set of individual plants increased with flowering synchrony, indicating that flowering synchrony could affect reproductive success.     

Reproductive biology of the endangered cypress Calocedrus macrolepis

We studied the reproductive biology of the endangered species Calocedrus macrolepis. We examined flower phenology and morphology, spatial distribution of male and female cones, pollen viability, variation in pollen density around the crown, and the pollination process. Calocedrus macrolepis is monoecious with unisexual flowers on the same branches. Pollen‐shed from male cones occurred from early September to mid‐February, whereas the female cones were receptive from mid‐September to mid‐March. The inconsistency of flowering periods between male and female cones within populations and among individuals extended the pollination period. There was a layered distribution pattern of male and female cones in the crown, with more male cones in the lower layer and more female cones in the upper layer. This resulted in different selfing and outcrossing rates at various locations. The pollen of C. macrolepis lack airbags, and consequently, it was dispersed within only 10 m of the mother tree. The short pollen dispersal distance could restrict gene flow among different geographical populations. Calocedrus macrolepis is endangered because of its high degree of selfing and inbreeding resulting from its monoecious characteristics and the short distance of pollen spread. These factors may result in a population decline.     

Relationships between tree size, crown shape, gender segregation and sex allocation in Pinus halepensis, a Mediterranean pine tree

Background and Aims

Sex allocation has been studied mainly in small herbaceous plants but much less in monoecious wind-pollinated trees. The aim of this study was to explore changes in gender segregation and sex allocation by Pinus halepensis, a Mediterranean lowland pine tree, within tree crowns and between trees differing in their size or crown shape.

Methods

The production of new male and female cones and sex allocation of biomass, nitrogen and phosphorus were studied. The relationship between branch location, its reproductive status and proxies of branch vigour was also studied.

Key Results

Small trees produced only female cones, but, as trees grew, they produced both male and female cones. Female cones were produced mainly in the upper part of the crown, and male cones in its middle and lower parts. Lateral branch density was correlated with the number of male but not female cones; lateral branches were more dense in large than in small trees and even denser in hemispherical trees. Apical branches grew faster, were thicker and their phosphorus concentration was higher than in lateral shoots. Nitrogen concentration was higher in cone-bearing apical branches than in apical vegetative branches and in lateral branches with or without cones. Allocation to male relative to female function increased with tree size as predicted by sex allocation theory.

Conclusions

The adaptive values of sex allocation and gender segregation patterns in P. halepensis, in relation to its unique life history, are demonstrated and discussed. Small trees produce only female cones that have a higher probability of being pollinated than the probability of male cones pollinating; the female-first strategy enhances population spread. Hemispherical old trees are loaded with serotinous cones that supply enough seeds for post-fire germination; thus, allocation to males is more beneficial than to females.     

Testing masting mechanisms of boreal forest species at different stand densities

Mast seeding is considered a reproductive trait resulting from several potential evolutionary forces. Although the mechanisms driving reproduction have been thoroughly investigated and discussed, their relative importance and possible coexistence remain an open question. Seed rain abundance and viability of balsam fir Abies balsamea, white spruce Picea glauca, and white birch Betula papyrifera were monitored during 1994–2007 along a chronosequence of developmental stages at different densities represented by four stands in the boreal forest of Quebec, Canada. The aim of the study was to verify whether seeding temporal dynamics and abundance were affected by stand density, and to test the causal relationships between the hypotheses of pollination efficiency and predator satiation. Seed rain abundance varied substantially among years and species, with the highest values being observed in 1994 and 1996, and attaining up to 28.2 × 10³ seeds m^?2 year^?1. However, the annual dynamics of seed production was similar in the three sites with the higher densities. The greater proportions of germinating and dead seeds were observed in white birch, while balsam fir showed an average of 11.2% of damaged seeds, which were infected by larvae. The conifer species had the higher proportions of empty seeds. The causal models tested on balsam fir demonstrated that seed rain abundance influenced the amount of both viable and larval‐infected seeds, although no causal relationship existed between these last two variables. The comparable and synchronous seed rain abundances observed among sites demonstrated that stand density can significantly affect the reproduction of trees, probably by limiting the access to resources, but not the annual dynamics of seeding. Pollination efficiency and predator satiation are important forces in the reproductive effort of trees and could be not mutually exclusive because both these mechanisms of natural selection synergistically coexist in driving the seeding dynamics of balsam fir.     

Changes in Siberian Fir Crown Architecture under Impaired Homeostasis

The crown of the Siberian fir (Abies sibirica) is characterized by a specific organization, namely, a narrow-pyramidal form, apical dominance, tiering by types of sexualization, and whorled branching. The gravitational vector is clearly manifested at all levels of the organization of the crown, that is, in the architecture (diageotropism of branches), morphostructure (dorsoventrality of tissues and organs of shoots, as well as the location of reproductive organs in the tree crown and on the shoot), and physiological processes. It was shown that the internal state of the tree is a determining factor in graviperception. When Siberian fir trees weaken, the diageotropism of branches is lost, which can serve as a universal biomarker of impairment of homeostasis.

     

Morphological diversity and structure of West Mediterranean Abies species

The aim of this study was the biometrical comparison of Abies pinsapo Boiss., Abies maroccana Trab. and Abies tazaotana Côzar ex Villar from Spain and Morocco. The morphological and anatomical analysis of the needle characters of these firs revealed significant taxonomic differences. The results showed considerable distances between regions and very small differences among the Moroccan fir taxa. The examination of the needles and earlier genetic analysis did not provide evidence for distinguishing A. tazaotana at the species level.     

Nutritional status of Abies pinsapo forests along a nitrogen deposition gradient: do C/N/P stoichiometric shifts modify photosynthetic nutrient use efficiency?

Chronic atmospheric N deposition has modified relative N availability, altering the biogeochemical cycles of forests and the stoichiometry of nutrients in trees, inducing P limitation, and modifying the N:P ratios of plant biomass. This study examines how the variation in the foliar stoichiometry of Abies pinsapo across an N deposition gradient affects foliar traits and photosynthetic rate. We measured the maximum net assimilation rates (A _max) and the foliar nitrogen (N) and phosphorus (P) concentrations in A. pinsapo needles of five age classes. The leaf mass per area and photosynthetic N and P use efficiencies (PNUE and PPUE, respectively) were also estimated. The results from the N-saturated stand (Sierra Bermeja, B) differed from the comparable N-limited stands under investigation (Yunquera, Y, and Sierra Real, SR). The trees from Y and SR exhibited a reduction in the N content in older needles, whereas the foliar N concentration at the B site increased with needle age. N and P were positively correlated at Y and SR, but not at B, suggesting that the overload of N in the trees at site B has exceeded the homeostatic regulation capacity of the N-saturated stand in terms of foliar stoichiometry. A _max and PNUE were correlated positively with P and negatively with the N/P ratio at the three study sites. The foliar N concentration was positively correlated with A _max at Y and SR. However, this relationship was negative for the B site. These findings suggest that the nutritional imbalance caused by increased chronic deposition of N and an insufficient supply of P counteracts the potential increase in net photosynthesis induced by the accumulation of foliar N.     

SIZE-DEPENDENT GENDER CHANGE IN GREEN DRAGON (ARISAEMA DRACONTIUM; ARACEAE)

Green dragon (Arisaema dracontium; Araceae) is a perennial woodland herb capable of switching gender from year to year. Small flowering plants produce only male flowers but when larger they produce male and female flowers simultaneously. Distinct male and monoecious phenotypes (referred to hereafter as plants) share a single underlying cosexual genotype. Four populations in southern Louisiana were sampled to determine frequencies and size distributions of male and monoecious plants, and to determine the relationship of plant size with male and female flower production in monoecious plants. Male plants were significantly smaller than monoecious plants and made up 34%–78% of flowering plants within populations. Flower number (average = 120) was weakly positively correlated with size. Monoecious plants produced an average of 169 flowers (90 female) and had 100% fruit set, with individual berries containing an average of 2.5 ovules and 1.3 filled seeds. Male flower number was negatively correlated, and female flower number positively correlated, with basal stem diameter. Extrapolation of regression slopes suggested that green dragon should become completely female at a size 20% larger than the largest plant observed in this study. A simple model of inflorescence development is presented to illustrate how the reproductive system of green dragon is related to that of jack-in-the-pulpit (A. tnphyllum), which exhibits a more distinct switch between male and female phenotypes.     

Soil Phosphorus Availability and Transformation Rates in Relictic Pinsapo Fir Forests from Southern Spain

Abies pinsapo fir forests are remnant of temperate-like coniferous forests currently subjected to the typical seasonal constraints of Mediterranean-type climates. We have examined for the first time, P availability and cycling in A. pinsapo forests from southern Spain by using both, measurements of concentrations of soil P fractions (resin extractable P, bicarbonate extractable P and NaOH extractable P) and rates of P supply to (P solubilization, P mineralization and phosphomonoesterases (PMEase activity) and P immobilization from the soil solution. Soils from A. pinsapo stands differing in lithology (serpentinitic and calcareous substrates) and successional status (from young-agradative to old-growth stages) were chosen for this study. Labile organic P fractions, PMEase activity and rates of P gross mineralization and immobilization were significantly higher in agradative stands on serpentines than in successionally comparable calcareous stands. This suggests an important role of the organic P subcycle in the A. pinsapo stand on serpentine. On calcareous lithology, PMEase activity and all soil P transformation rates significantly increased throughout the successional series. Similarly, all organic and inorganic P fractions measured in old-growth forests showed the maximum values of the series. These trends fit the predictions of standard patterns of P cycling changes along with succession, in which P supply to plants greatly depends on solubilization from mineral forms at early-to-mid successional stages, whereas the importance of processes related to the organic P subcycle increases as succession progresses.