Occurrence of vesicular-arbuscular mycorrhizae in Pseudotsuga menziesii and Tsuga heterophylla seedlings grown in Oregon Coast Range soils

 Vesicular-arbuscular mycorrhizae (VAM) were common in seedlings of Pseudotsuga menziesii and Tsuga heterophylla grown in a greenhouse soil bioassay in soils collected from the Oregon Coast Range. Although root samples were heavily colonized by ectomycorrhizal fungi (EM), VAM colonization was observed in the cortical cells of both secondary and feeder roots. Vesicles, arbuscules, and hyphae typical of VAM occurred in 48% of 61 P. menziesii and 25% of 57 T. heterophylla seedlings. The ecological significance of VAM presence in the Pinaceae, as well as interactions among VAM, EM, and the plant host, deserve future investigation. Accepted: 16 August 1995     

The population biology of Bromus tectorum in forests: effect of disturbance,grazing, and litter on seedling establishment and reproduction

Summary The effect of tree canopy, understory, herbivores, and litter depth on seedling establishment, survival, and reproduction of the alien grass, Bromus tectorum (cheatgrass), was examined in a series of experiments in four forest habitat types in western North America. Higher recruitment, survival, and reproduction on clearcuts, which would be expected if the overstory alone is limiting the distribution of cheatgrass in forests, were not observed. Removing the understory in an otherwise undisturbed Pinus ponderosa forest did, however, increase the emergence of B. tectorum, but plants in these experimentally-created openings were more vulnerable to grazing by small mammals. In contrast, removing the sparse understory in an Abies forest neither enhanced recruitment nor increased the incidence of grazing of B. tectorum seedlings. Regardless of the forest habitat, most grazed plants died before maturity; even fewer grazed plants produced seeds. Litter depth influenced both recruitment and biomass production: both the rate of germination and the size of resultant seedlings were lower on thick litter (6 cm) compared to results on thin litter (1.5 cm). In the more open Pinus ponderosa and Pseudotsuga menziesii forests, cheatgrass colonization may often occur in openings in the understory alone. Colonization in the more shady A. grandis and Thuja plicata forests is unlikely, however, unless the opening extends through both the understory and the overstory. As a result, cheatgrass is unlikely to increase in any of these forests unless the scale and incidence of disturbance increases substantially.     

Vesicular endophytes in roots of the Pinaceae

Vesicles and hyphae typical of vesicular-arbuscular mycorrhizae (VAM) were common in seedlings of Pseudotsuga menziesii, Abies lasiocarpa and Tsuga mertensiana growing in openings where herbaceous hosts of these fungi were common. Seedlings of A. lasiocarpa, Tsuga heterophylla, and T. mertensiana growing under closed forest canopies also had vesicles but at a much lower incidence than seedlings in the openings. The Pinaceae are generally assumed to be ectomycorrhizal, but Glomus-type colonizations occurred on the same seedlings as the ectomycorrhizae. The ecological significance of abundant VAM-type endophytes in otherwise ectomycorrhizal hosts deserves comprehensive study.     

Inoculation of containerized Pseudotsuga menziesii and Pinus pinaster seedlings with spores of five species of ectomycorrhizal fungi

 Container-grown Pseudotsuga menziesii and Pinus pinaster seedlings were inoculated with water suspensions of spores of five ectomycorrhizal fungi commonly found in northeastern Spain. Pseudotsuga menziesii seedlings were inoculated with basidiospores of Melanogaster ambiguus, or Rhizopogon subareolatus, or with ascospores of Tuber maculatum. Pinus pinaster seedlings were inoculated with basidiospores of Melanogaster ambiguus, Rhizopogon roseolus or Scleroderma citrinum. The spore concentrations were 10²–10⁷ spores per seedling for Melanogaster ambiguus (in Pseu dotsuga menziesii) and Rhizopogon subareolatus, 10³–10⁷ for Melanogaster ambiguus (in Pinus pinaster), Rhizopogon roseolus, and Scleroderma citrinum, and 10²–10⁴ for Tuber maculatum. Melanogaster ambiguus colonized more short roots in a larger proportion of plants at 10⁷ spores per seedling than at any other rate. The highest colonization by Rhizopogon subareolatus was obtained at 10⁴ spores per seedling and higher, and all inoculated plants became infected at 10⁶ spores per seedling and higher. Tuber maculatum colonized a high percentage of short roots at all rates tested; the proportion of infected plants was over 80% at 10³–10⁴ spores per plant, decreasing to 50% at 10² spores per plant. Rhizopogon roseolus colonized the highest number of short roots on nearly all the inoculated plants when applied at 10⁵ spores per seedling and higher. Scleroderma citrinum colonized a high percentage of short roots on all inoculated plants when applied at 10⁵ spores per seedling and higher. The abundance of sporocarps of Melanogaster ambiguus, Rhizopogon subareolatus, R hizopogon roseolus and Scleroderma citrinum and their colonization ability at relatively low rates allows these spores to be used as ectomycorrhizal inocula on a large scale. Accepted: 27 February 1996     

Arbuscular mycorrhizal fungal phylogeny-related interactions with a non-host

Arbuscular mycorrhizal fungi (AMF) form symbiotic relationships with most vascular plants including some gymnosperm species. Although species in the gymnosperm family Pinaceae normally develop ectomycorrhizal associations, AMF hyphae and vesicles, typical of members of the Suborder Glomineae, have been reported in the roots of some Pinaceae species. However, it is not known whether AMF belonging to various species and suborders are able to colonize roots of Pinaceae species and to what extent this influences the performance of Pinaceae seedlings. We tested in each of the Glomaceae, Acaulosporaceae and Gigasporineae AMF families two species for their ability to colonize and affect the growth of Pinus strobus (eastern white pine) in the presence or absence of an AMF host plant (Trifolium pretense—red clover). Glomus intraradices was the only AMF that colonized pine roots, predominantly in the presence of clover, forming intracellular hyphae and vesicles but not arbuscules. Colonization, however, did not relate to increased pine biomass and the overall presence of AMF, regardless of colonization abilities, resulted in a biomass reduction. This effect on pine seedling biomass was explained by the AMF family to which the AMF belonged, indicating that the effects of AMF on the non-host pine may be related to phylogeny. Acaulosporaceae species reduced pine biomass the most whereas, Gigasporineae species had the smallest effect on biomass. These preliminary results suggest that AMF may affect the soil microflora differently among AMF families in previously unsuspected ways with potential consequences for non-AMF host growth.     

Anaerobic induced ethanol synthesis in the stems of greenhouse-grown conifer seedlings

Ethanol synthesis was induced in stem segments from greenhouse-grown conifer seedlings by placing them in a N₂ atmosphere at 30 °C for 24 h. Stems from ponderosa pine,Pinus ponderosa Dougl. ex Laws., sugar pine,Pinus lambertiana Dougl., Pacific silver fir,Abies amabalis Dougl. ex Forbes, and lodgepole pine,Pinus contorta Dougl. ex Loud, produced the highest quantities of ethanol. This group also had the smallest and slowest growing stems. Within each of these species the amount of ethanol produced was inversely related to the stem volume. Stems from western hemlock,Tsuga heterophylla (Raf.) Sarg., grand fir, Abies grandis Dougl. ex Forbes, Douglas-fir,Pseudotsuga menziesii (Mirb.) Franco, and western redcedar,Thuja plicata Donn ex D. Don, all produced equivalent but low ethanol concentrations. These species had the largest and fastest growing stems. In this group only grand fir exhibited an inverse relationship between ethanol concentrations and stem volume. The relative amounts of ethanol synthesized by stems from Douglas-fir, western hemlock and western redcedar seedlings were not the same as subsequently observed in logs from mature trees of the same species under field conditions. Differences in the anaerobic environments for the two stem types could have affected the quantities of ethanol produced. The observed high amounts of ethanol produced by the stems from pine species were discussed in terms of their ability to handle periods of anaerobic stress or hypoxia.     

The co-occurrence of ectomycorrhizal,arbuscular mycorrhizal,and dark septate fungi in seedlings of four members of the Pinaceae

Although roots of species in the Pinaceae are usually colonized by ectomycorrhizal (EM) fungi, there are increasing reports of the presence of arbuscular mycorrhizal (AM) and dark septate endophytic (DSE) fungi in these species. The objective of this study was to determine the colonization patterns in seedlings of three Pinus (pine) species (Pinus banksiana, Pinus strobus, Pinus contorta) and Picea glauca x Picea engelmannii (hybrid spruce) grown in soil collected from a disturbed forest site. Seedlings of all three pine species and hybrid spruce became colonized by EM, AM, and DSE fungi. The dominant EM morphotype belonged to the E-strain category; limited colonization by a Tuber sp. was found on roots of Pinus strobus and an unknown morphotype (cf. Suillus–Rhizopogon group) with thick, cottony white mycelium was present on short roots of all species. The three fungal categories tended to occupy different niches in a single root system. No correlation was found between the percent root colonized by EM and percent colonization by either AM or DSE, although there was a positive correlation between percent root length colonized by AM and DSE. Hyphae and vesicles were the only AM intracellular structures found in roots of all species; arbuscules were not observed in any roots.     

Occurrence and dominance of six Pacific Northwest conifer species

Questions: Can probability of occurrence and dominance be accurately estimated for six important conifer species with varying range sizes? Does range size impact the accuracy of species probability of occurrence models? Is species predicted probability of occurrence significantly related to observed dominance? Location: Pacific Northwest region, North America (60°–40°N, 140°–110°W). Methods: This study develops near range‐wide predictive distribution maps for six important conifer species (Pseudotsuga menziesii, Tsuga heterophylla, Pinus contorta, Thuja plicata, Larix occidentalis, and Picea glauca) using forest inventory data collected across the United States and Canada. Species model accuracies are compared with range size using a rank scoring system. A suite of climate and topographic predictor variables are used to investigate environmental constraints that limit species range and quantify relationships between species predicted probability of occurrence and dominance at both plot and landscape scales. Results: Evaluation statistics revealed significant and accurate probability of occurrence models were developed for all six species. Based on ranked evaluation statistics, Tsuga heterophylla had highest overall model accuracy (statistic rank score=5) and Pinus contorta the lowest (statistic rank score=17). Across species, ranked evaluation statistics also revealed a pattern of decreasing model accuracy with increasing range size. At plot level, correlations between dominance and probability of occurrence were weakly positive for all species with only half of the species having statistically significant correlations. Pseudotsuga menziesii had the highest correlation (r=0.36, P<0.001) and Thuja plicata lowest (r=0.038, P=0.799). At the 50‐km scale, correlations between dominance and probability of occurrence improved for all species except Pinus contorta. Pseudotsuga menziesii displayed the highest correlation (r=0.68, P<0.001) and Thuja plicata the lowest (r=0.07, P>0.709). Conclusions: Species probability of occurrence model accuracy decreased with increasing range size. The strength and significance of correlations between probability of occurrence and dominance varied considerably by species and across spatial scales. Apart from Pseudotsuga menziesii and L. occidentalis, the results suggest that probability of occurrence is not a consistently reliable surrogate for species dominance in Pacific Northwest forests. We demonstrate how the degree of correlation between species occurrence and dominance can be used as an indicator of how well predictions of occurrence characterize the optimal niche of a species.     

Mycorrhiza-like interaction by Morchella with species of the Pinaceae in pure culture synthesis

 Isolates from two species of Morchella were tested for ability to form mycorrhizae in pure culture synthesis with Arbutus menziesii, Larix occidentalis, Pinus contorta, Pinus ponderosa, andPseudotsuga menziesii. Ectomycorrhizal structures (mantle and Hartig net) formed with the four species of the Pinaceae but not with A. menziesii. Results are compared to previous studies on morel mycorrhizae and discussed in an ecological context. Accepted: 23 October 1999     

Ability of native ectomycorrhizal fungi from northern Spain to colonize Douglas-fir and other introduced conifers

 Thirty-six isolates from 27 species of native ectomycorrhizal fungi collected in northern Spain were tested for ectomycorrhiza formation with Pseudotsuga menziesii seedlings in pure culture syntheses. Thirteen of those species were also tested for ectomycorrhiza formation with six other species of conifers (two native and four introduced) to compare their colonization potential. Twenty-three fungal isolates from 18 species formed ectomycorrhizas with Pseudotsuga menziesii. The colonization level of the root system varied markedly among the different fungal species. Eight fungi colonized over 50% of the short roots. Nine fungi did not form ectomycorrhizas even though some of them were collected in pure stands of Pseudotsuga menziesii. Laccaria laccata, Lyophyllum decastes, Pisolithus tinctorius, and Scleroderma citrinum formed abundant ectomycorrhizas on all the conifers tested. Lactarius deliciosus, Rhizopogon spp., and Suillus luteus showed the greatest host specificity. The success in the introduction of some exotic conifers for reforestation in northern Spain is discussed in relation to their compatibility with native ectomycorrhizal fungi. Accepted: 28 August 1995     

Inoculation and field testing of Sitka spruce and Douglas fir with ectomycorrhizal fungi in the United Kingdom

Picea sitchensis and Pseudotsuga menziesii seedlings were grown in containers, inoculated with ectomycorrhizal fungi, and planted in British forestry sites. Root samples taken during the year after planting were assessed for mycorrhiza formation. Survival and shoot height were assessed at the end of each year. Observations were made each autumn on the occurrence of sporophores of ectomycorrhizal fungi. Pot experiments were used to assess the colonization potential of soils from the experimental locations. Assessment of mycorrhiza formation by the inoculant fungi both before planting and the following year showed much variation among the fungi used. Similar variation was found among field sites. Inoculation with Laccaria isolates was most successful. Height measurements are reported for the first 2 years after planting, at which time there were few significant effects on growth of Picea sitchensis or Pseudotsuga menziesii seedlings. Experimental assessment of colonization potential was of little value in this work for predicting events in the forest.     

A chalcone synthase/stilbene synthase DNA probe for conifers

A probe for chalcone synthase (CHS) was generated by PCR using chalcone synthase conserved sequences. The cloned PCR product has high similarity to both chalcone synthase and stilbene synthase sequences. The probe was used to examine the organization of chalcone synthase and stilbene synthase genes in Abies procera, Pinus lambertiana, P. monticola, Picea glauca, P. sitchensis, Pseudostuga menziesii, Taxus brevifolia, and Thuja plicata. A large number of hybridizing bands were found in all species except T. plicata which did not cross hybridize. The hybridization patterns are highly polymorphic between the species and are also polymorphic within several of them.     

Anatomical Structures of the VA Mycorrhiza in the Apocynaceae (Gentianales)

Roots of 19 Apocynaceae species were studied anatomically with respect to their symbiosis with VAM-fungi. In plants collected from the field, VAM-fungi were established in the root cortex. Also, inoculations with different Glomus species on the cultured plants are very successful in the infection and colonization of the root cortex. After penetration of the rhizodermis, the special exodermal short cells become colonized by winding hyphae. Then, in the root cortex of many Apocynaceae species, the VAM-fungi produce intercellular running hyphae which leads to extensive colonization of the root. Arbuscules develop on intracellular running hyphae, whereas vesicles develop mainly on intercellular hyphae. Except for some special details, this is the most common type of colonization of VAM fungi in flowering plants. But in Amsonia tabernaemontana, Nerium oleander, and Thevetia peruviana, another type of colonization could be observed. In these species, the colonization of the hyphae within the root cortex is only possible by intracellular growth. Intercellular running hyphae in the root are lacking. Therefore, after penetration the colonization in the cortex is cluster-like and strictly limited. Only by additional penetrations from hyphae in the soil, will roots show heavy infestations. This type of growth of the VAM fungi in the root is well known from the Gentianaceae and was explained as a structural incompatibility. In Catharanthus roseus, Pachypodium lamerei, and Trachelospermum jasminoides, intermediate stages of both types of colonization could be described. The results are discussed in the search for possible stimulants for structural incompatibility.     

What happens to the mycorrhizal communities of native and exotic seedlings when Pseudotsuga menziesii invades Nothofagaceae forests in Patagonia,Argentina?

Pseudotsuga menziesii is one of the most widely planted conifers in the Patagonian Andes of Argentina, having invading characteristics that are broadly reported. We studied the mycorrhizal status of seedlings along six Nothofagaceae + P. menziesii invasion matrices to investigate their role in the invasive process, according to these hypothesis: a) The abundance and richness of EM will be higher in seedlings grown in their own soil; b) In the presence of native EM inoculum, the invasive plant will be associated with generalist mycorrhizae (EM and/or AM), c) AM associations will be more abundant in P. menziesii seedlings grown in Interface or native forest soils, d) Mycorrhizal community differences between treatments will alter host fitness (growth and nutritional parameters). Seedlings from Nothofagus dombeyi, N. antarctica, Lophozonia alpina, L. obliqua and Pseudotsuga menziesii were set up in a soil-bioassay that included soils from non-invaded Nothofagaceae forests, pure P. menziesii plantations, and the interface between both. Pseudotsuga menziesii seedlings showed a decreasing, although never null, ectomycorrhizal (EM) colonization pattern from plantations to non-invaded forests, mainly with exotic EM species. Hebeloma mesophaeum and Wilcoxina sp. 1, two EM species with cosmopolitan distribution, were found to be shared by both tree species. Hebeloma hiemale and Wilcoxina sp. 1, common mycorrhizal partners of P. menziesii in Patagonia although not registered from Nothofagaceae forest, were found to be associated with N. antarctica, being the first report for both fungal species. Pseudotsuga menziesii seedlings showed the ability to form different arbuscular mycorrhiza (AM) colonization types (Paris-, Arum-, Both- and Intermediate-types) depending on the treatments, with significantly higher presence of Intermediate-type in the Interface treatment, where colonization was low. The shared EM species and the presence of different AM colonization types imply enhanced possibilities for invasive P. menziesii seedlings establishment and development. Seedling features and EM colonization rates evidenced that P. menziesii invasion could produce maladaptation (defined as a relative decline in host fitness due to altered mycorrhizal communities from native settings) of mycorrhizal communities, seriously injuring native ecosystem.     

Growth and nutrient uptake of sorghum cultivated with vesicular-arbuscular mycorrhiza isolates at varying pH

This study was conducted to determine the effects of different pH regimes on root colonization with four vesicular-arbuscular mycorrhiza (VAM) isolates, and VAM effects on host plant growth and nutrient uptake. Sorghum [Sorghum bicolor (L.) Moench] was grown at pH 4.0, 5.0, 6.0 and 7.0 (±0.1) in hydroponic sand culture with the VAM isolates Glomus etunicatum UT316 (isolate E), G. intraradices UT143 (isolate I), G. intraradices UT126 (isolate B), and an unknown Glomus isolate with no INVAM number (isolate A). Colonization of roots with the different VAM isolates varied differentially with pH. As pH increased, root colonization increased with isolates B and E, remained unchanged with isolate I, and was low at pH 4.0 and high at pH 5.0, 6.0, and 7.0 with isolate A. Isolates E and I were more effective than isolates A and B in promoting plant growth irrespective of pH. Root colonization with VAM appeared to be independent of dry matter yields or dry matter yield responsiveness (dry matter produced by VAM compared to nonmycorrhizal plants). Dry matter yield responsiveness values were higher in plants whose roots were colonized with isolates E and I than with isolates A and B. Shoot P concentrations were lower in plants colonized with isolates E and I than with isolates A and B or nonmycorrhizal plants. This was probably due to the dilution effect of the higher dry matter yields. Neither the VAM isolate nor pH had an effect on shoot Ca, Mg, Zn, Cu, and Mn concentrations, while the VAM isolate affected not only P but also S, K, and Fe concentrations. The pH x VAM interaction was significant for shoot K, Mg, and Cu concentrations.     

Arbuscular mycorrhizal development in three crucifers

To determine the developmental patterns of arbuscular mycorrhizae (AM) in three crucifers (Brassicaceae) of differing life histories, we inoculated seedlings of the annual Capsella bursa-pastoris, biennial Hesperis matronalis, and the perennial Matthiola incana with Glomus intraradices. The plants were grown either alone or in a matrix of living roots of the mycotrophic grass Sorghum sudanense. The percent root length colonized was greatest in C. bursa-pastoris and least in H. matronalis. Colonization was greater in plants grown in the grass matrix than in plants grown alone, and colonization in grass matrix-grown plants continued to increase over the 90-day growth period, whereas colonization leveled off or decreased near the end of the growth period in crucifers grown alone. No arbuscules were observed in crucifer roots at any time, which suggests that AM in these crucifers is nonfunctional. Furthermore, the increase in colonization only in pots with both crucifers and active mycotrophic roots suggests that AM development in crucifer roots is primarily the consequence of progressive root senescence in the crucifer and continued inoculum spread from the mycotrophic plant.     

Comparative effects of four mycorrhizal fungi on loblolly pine seedlings growing in a greenhouse in a Piedmont soil

Summary A greenhouse study was conducted to evaluate the effect of ectomycorrhizae on loblolly pine (Pinus taeda L.) growing in a Piedmont soil. Pine seedlings were inoculated with one of four species of fungi (Scleroderma aurantium, Pisolithus tinctorius, Thelophora terrestris, andRhizopogon roseolus). The seedlings were grown in pots containing a Cecil sandy clay loam amended to create a gradient of extractable P ranging from 5.9 to 52.5 g/g. After ten months, all colonized seedlings were significantly larger than control seedlings. However, of the four fungi,Scleroderma aurantium mediated a far superior shoot growth response to increasing levels of soil P; the seedlings were significantly larger than those colonized by any other fungus and also had the largest root systems and greatest degree of mycorrhizal colonization.     

Influence of phosphorus and formononetin on isozyme expression in the Zea mays‐Glomus intraradices symbiosis

Maize (Zea mays L. cv. Great Lakes 586) plants were either inoculated with the vesicular-arbuscular mycorrhizal (VAM) fungus Glomus intraradices Schenck and Smith, or grown in the presence of the isoflavone formononetin or were provided with both G. intraradices and formononetin. All plants were grown in soil containing one of five levels of inorganic P (between 8 and 110 µg g^?1 soil). By 3 weeks there were significant differences in a number of enzyme activities and in the pattern of isoenzymes in roots colonized by the VAM fungus or treated with formononetin. One NAD-malate dehydrogenase (MDH) isozyme was expressed only in mycorrhizal roots, whether treated or not with formononetin. Despite differences in the soil P level, the expression of this isozyme was not observed in non-mycorrhizal roots, indicating specific expression in the mycorrhizae. We suggest that MDH isozyme could serve as a specific, early indicator of the Zea-Glomus symbiosis. Differences in the esterase (EST) isozyme pattern were not detectable between VAM and non-VAM roots, suggesting that this enzyme system is not a good parameter for the evaluation of mycorrhizal colonization. As available P in the soil increased, total EST activity appeared to increase as well. Interestingly, total peroxidase (POX) activity increased along with P suggesting that as plant P nutrition improved, both cell wall ramification and the quantity of defense peroxidases increased as well. Total POX activity from mycorrhizal roots was inversely correlated with root colonization, indicating that there was suppression of POX activity by the host under low soil P. Most interestingly, formononetin further decreased POX activity regardless of the level of P or mycorrhizal status. This may suggest one mechanism by which formononetin enhances root VAM colonization. The presence of this isoflavone suppressed POX activity in mycorrhizal roots allowing a rapid penetration and spread of the fungus in the root cortex. The interplay between host root, soil P levels, secondary metabolites and endogenous host enzyme activities and a particular VAM fungus has a profound effect on the efficiency, duration and functioning of an endomycorrhizal symbiosis.     

The Importance of Large-Diameter Trees to Forest Structural Heterogeneity

Large-diameter trees dominate the structure, dynamics and function of many temperate and tropical forests. However, their attendant contributions to forest heterogeneity are rarely addressed. We established the Wind River Forest Dynamics Plot, a 25.6 ha permanent plot within which we tagged and mapped all 30,973 woody stems ≥1 cm dbh, all 1,966 snags ≥10 cm dbh, and all shrub patches ≥2 m². Basal area of the 26 woody species was 62.18 m²/ha, of which 61.60 m²/ha was trees and 0.58 m²/ha was tall shrubs. Large-diameter trees (≥100 cm dbh) comprised 1.5% of stems, 31.8% of basal area, and 17.6% of the heterogeneity of basal area, with basal area dominated by Tsuga heterophylla and Pseudotsuga menziesii. Small-diameter subpopulations of Pseudotsuga menziesii, Tsuga heterophylla and Thuja plicata, as well as all tree species combined, exhibited significant aggregation relative to the null model of complete spatial randomness (CSR) up to 9 m (P≤0.001). Patterns of large-diameter trees were either not different from CSR (Tsuga heterophylla), or exhibited slight aggregation (Pseudotsuga menziesii and Thuja plicata). Significant spatial repulsion between large-diameter and small-diameter Tsuga heterophylla suggests that large-diameter Tsuga heterophylla function as organizers of tree demography over decadal timescales through competitive interactions. Comparison among two forest dynamics plots suggests that forest structural diversity responds to intermediate-scale environmental heterogeneity and disturbances, similar to hypotheses about patterns of species richness, and richness- ecosystem function. Large mapped plots with detailed within-plot environmental spatial covariates will be required to test these hypotheses.     

Late quaternary dynamics of pollen influx at mineral lake,Washington

Pollen influx analysis at Mineral Lake, Washington, indicates that immediately south of the Puget Lobe of the Fraser Glaciation, tundra was a characteristic vegetation until 16,300 years ago. Invasion ofPinus contorta began 17,500 years B.P., and boreal climax conifers (Abies, Picea andTsuga mertensiana), 16,300, but was temporarily interrupted by the Vashon advance (14,500–14,000 yr B.P.).Pseudotsuga menziesii began to grow in population 10,750 years ago, and woodland was established within a time span of 1,000 years. Modern lowland coniferous forests began to form 7,000 years ago. Logistic analysis of pollen abundance changes show that the intrinsic growth rate,r (yr⁻¹), of pioneer species (e.g. 0.024–0.026 inPteridium aquilinum) is higher than that of climax species (e.g. 0.003 inThuja plicata).P. menziesii, a subclimax species, shows an intermediater value (0.013) between these two ecologically different taxa. The absoluter value ofP. contorta (−0.011) is only slightly lower than that ofP. menziesii, although their replacement began almost simultaneously. Thus competition between these species was intense before the inflection point ofPinus curve 10,100 years ago. At this time, forest gaps became abundantly available forPseudotsuga, as indicated by a peak of the diagnostic factor (the reciprocal of the pollen influx).