Reproductive ecology of the three Nordic Pinguicula species (Lentibulariaceae)

Reproductive ecology (pollination biology, breeding systems, and reproductive effort and success) of the three Nordic species of Pinguicula, P. alpina, P. villosa , and P. vulgaris (Lentibulariaceae), was investigated in a subarctic-subalpine area at Abisko, N Swedish Lapland. Additional studies were carried out at Latnjajaure Field Station, a subarctic-alpine tundra site in the Abisko mountains ( P. alpina, P. vulgaris ), and in W Greenland ( P. vulgaris ). At Abisko the species are sympatric and large populations of all three were found within a 50 × 50 m area. The three species are reproduc-tively isolated by internal barriers by occupying different ploidy levels. Pinguicula alpina and P. vulgaris thrive in base-rich habitats, whereas P. villosa is restricted to nutrient-poor Sphagnum bogs, but habitat separation alone is probably not sufficient to prevent illegitimate pollen flow among the species. However, results showed that there are large and consistent differences in pollination biology, flowering phenology, and breeding systems, and these factors interact to create a highly efficient reproductive isolation at all levels, pre-zygotic as well as post-zygotic. Pinguicula alpina is an early-flowering outbreeder, P. vulgaris is an opportunistic late-flowering inbreeder, and P. villosa is quite intermediate between the two extremes. The phenology-based life history strategies of the Pinguicula species were in accordance with a general model developed for arctic flowering plants, predicting maximized fitness through pollen or seed in early- and late-flowering species, respectively.     

Nitrogen uptake from prey and substrate as affected by prey capture level and plant reproductive status in four carnivorous plant species

Uptake of nitrogen from prey and substrate and partitioning of prey-derived nitrogen were studied in the carnivorous plant species Pinguicula alpina, P. villosa, P. vulgaris and Drosera rotundifolia in a subarctic environment. Efficiency in nitrogen uptake from prey was evaluated by tracing ¹⁵N from ¹⁵N-enriched Drosophila flies fed to the plants. The in situ uptake efficiency differed somewhat between species and ranged from 29 to 41% of prey N. This efficiency was not affected by different feeding levels or plant reproductive status (flowering or non-flowering). A test of the amount of N absorbed from prey caught on flower stalks of Pinguicula villosa and P. vulgaris showed that both species took up little of what was available in prey (2.5% or less). The uptake efficiency found in greenhouse grown plants was higher than in plants in situ (40–50% vs. 30–40% respectively). This could probably best be explained by the absence of rain and a higher temperature in the greenhouse. The prey-derived ¹⁵N was traced to reproductive organs and winter buds. Non-flowering individuals allocated 58–97% of the N derived from prey to their winter buds. Flowering individuals allocated 17–43% of the N income from prey to reproduction, while 34–71% were allocated to buds. Root uptake of nitrogen was stimulated by increased prey capture. This increase in uptake of nitrogen from the substrate was larger than the potential direct uptake of nitrogen from captured prey.     

Sexual dimorphism and sex ratios of two Australian dioecious species of alpine pineapple grass, Astelia alpina var. novaehollandiae and Astelia psychrocharis (Asteliaceae)

Sexual dimorphism and male biased sex ratios have been predicted for dioecious plants experiencing the limited conditions for growth and reproduction found in many alpine environments. To test these predictions, the reproductive ecology of two congeneric, co-occurring, dioecious, clonal, species was examined in the subalpine and alpine zones of Kosciuszko National Park, southeastern Australia. Specifically, plant size (vegetative cover of plants in quadrats), floral display (number of flowers per inflorescence, number of inflorescences per quadrat) and sex ratios (proportion of females in quadrats with flowers) were examined in ten populations of Astelia psychrocharis (Asteliaceae) and nine populations of Astelia alpina var. novae-hollandiae (Asteliaceae). Sexual dimorphism did occur, with males having more flowers per inflorescence (106% more flowers for A.alpina males and 12% more for A.psychrocharis males compared to females) and more inflorescences per quadrat than females (78% more inflorescences for A.alpina males and 46% more inflorescences for A.psychrocharis males compared to females). Plant size did not differ between male and female quadrats of either species, nor were there male biased sex ratios. However, plant size was related to flowering status in A.psychrocharis with the 65 quadrats that did not flower having lower vegetative cover than the 175 flowering quadrats indicating that there may be a minimum size/ cover required prior to flowering in this species. For A.alpina, all but two of the 185 quadrats randomly sampled flowered. There was no effect of altitude on plant size and very little effect of altitude on floral display for either species, apart from a slight increase in the number of inflorescences per quadrat with increasing altitude for A.psychrocharis, and slight decrease in number of flowers per inflorescence with increasing altitude for A.alpina females.     

Between-population variation in size-dependent reproduction and reproductive allocation in Pinguiculavulgaris (Lentibulariaceae) and its environmental correlates

Several important fitness components in herbaceous perennial plants are commonly related to plant size: flowering probability, reproductive allocation and fecundity. However, evidence for such size-dependence of fitness components is mostly anecdotal and unconnected to other life history traits. Here we report size-dependence for flowering probability and reproductive allocation in 11 populations of Pinguicula vulgaris and relate it to environmental factors. Flowering probability was size-dependent in all populations of P. vulgaris , and indicated the existence of a threshold size for reproduction. Populations at low altitudes and in wet soils showed a significantly higher threshold size for reproduction. Reproductive mass was also size-dependent in all populations. We found considerable between-population differences in the slope and the intercept of the regression between plant size and reproductive mass. This variation was weakly related to the environmental factors measured. In general, relationships between different size-dependent fitness components were low. Instead of showing a covariation of traits, in line with interpretations in terms of life history "tactics", P. vulgaris seemed to independently vary each size-dependent fitness component in each locality. In particular, no significant relationship was found between threshold size for reproduction and the slope of size-dependent reproductive allocation, as predicted by previous authors. Neither we found a significant influence of somatic cost of reproduction on size-dependent fitness components.     

Phylogenetic analysis of Pinguicula (Lentibulariaceae): chloroplast DNA sequences and morphology support several geographically distinct radiations

The genus Pinguicula is one of the three genera of the carnivorous Lentibulariaceae, comprising approximately 80 species. Phylogeny inference using nucleotide sequences of the chloroplast gene matK and the trnK group II intron, as well as a set of 32 morphological characters revealed five well-supported, major lineages within the genus. These lineages largely reflect radiations in clearly defined geographic regions, whereas most previously recognized sections of the genus are shown to be para- or polyphyletic. A species-rich Mexican-Central American-Caribbean clade has the Eurasian P. alpina and an East Asian clade as successive sisters. All three are characterized by a production of flower buds on winter-resting plants, a specific corolla hair structure and a very large corolla lower central lobe. Another diverse clade is composed of species with primarily European distribution including the widespread type species P. vulgaris. For this clade, vegetative reproduction during dormancy is synapomorphic. Species native to SE North America and the South American Andes and a group of Mediterranean and NE Atlantic coast species together appear in a fifth well-supported clade, that is characterized by a tropical-type growth habit. It is the only clade that has reached temperate zones of the southern hemisphere.     

Microsatellite diversity of the agriculturally important alpine grass Poa alpina in relation to land use and natural environment

BACKGROUND AND AIMS: The Alpine Meadow Grass Poa alpina is common in subalpine and alpine natural sites and agriculturally used land, where it is an important fodder grass. Natural factors and human land use are supposed to have been shaping its genetic diversity for hundreds of years. The species comprises sexually and vegetatively reproducing plants. The aim of this study was to investigate the effects of agricultural land use, environmental factors and the mode of reproduction on the distribution of its microsatellite diversity within and among populations and to analyse whether its genetic diversity is correlated with plant species diversity in grassland parcels. METHODS: Genetic diversity of P. alpina was assessed with five microsatellite markers for 569 plants originating from 20 natural sites and from 54 grassland parcels of different cultural tradition, land use and altitude in the Swiss Alps. Due to polyploidy and frequent aneuploidy of the species, data analyses were based on the presence of microsatellite bands. KEY RESULTS: A low but significant differentiation was found in microsatellite bands among natural sites and agriculturally used parcels, while their microsatellite band diversity within populations did not differ. An increased differentiation was found in microsatellite bands with increasing geographic distance among parcels, and a differentiation among grazed and mown parcels, and among sexually and vegetatively reproducing populations. Band richness of sampled plants per village was higher for villages where parcels represented more different land-use types. Within populations, microsatellite band diversity was higher in grazed than in mown parcels. CONCLUSIONS: The diversity of human land use in the Alps was associated with genetic diversity of P. alpina. Therefore, the ongoing socio-economically motivated land-use changes, which reduce the number of different land-use types, will affect the genetic diversity of P. alpina negatively.     

Changes in reproductive investment with altitude in an alpine plant

Aims In perennial species, the allocation of resources to reproduction results in a reduction of allocation to vegetative growth and, therefore, impacts future reproductive success. As a consequence, variation in this trade-off is among the most important driving forces in the life-history evolution of perennial plants and can lead to locally adapted genotypes. In addition to genetic variation, phenotypic plasticity might also contribute to local adaptation of plants to local conditions by mediating changes in reproductive allocation. Knowledge on the importance of genetic and environmental effects on the trade-off between reproduction and vegetative growth is therefore essential to understand how plants may respond to environmental changes.Methods We conducted a transplant experiment along an altitudinal gradient from 425 to 1?921 m in the front range of the Western Alps of Switzerland to assess the influence of both altitudinal origin of populations and altitude of growing site on growth, reproductive investment and local adaptation in Poa alpina .Important findings In our study, the investment in reproduction increased with plant size. Plant growth and the relative importance of reproductive investment decreased in populations originating from higher altitudes compared to populations originating from lower altitudes. The changes in reproductive investment were mainly explained by differences in plant size. In contrast to genetic effects, phenotypic plasticity of all traits measured was low and not related to altitude. As a result, the population from the lowest altitude of origin performed best at all sites. Our results indicate that in P. alpina genetic differences in growth and reproductive investment are related to local conditions affecting growth, i.e. interspecific competition and soil moisture content.     

The realized food niches of three species of stream-dwelling triclads (Turbellaria)

SUMMARY. 1. The diet of Phagocata vitta, Crenobia alpina and Polycelis felina in a Welsh stream was examined, using a serologicat technique, at monthly intervals over 18 months.
2. For each triclad species, dietary composition changed little with time or increasing body size, and overall feeding intensity was greatest in summer and least in winter.
3. Alt species fed most heavily on oligochaetes, chironomids and Gammarus. Phagocata vitta fed lightly on Plecoptera, Trichoptera and Ephemeroptera whilst the other two triclad species fed substantially on the first two of these prey categories.
4. Food niche breadth indices indicated a wide diet for all species with some specialization in some months. Electivity indices showed positive electivity for oligochaetes and chironomids by all species and for Plecoptera by C. alpina and P. felina .
5. Diets overlapped and none had a food refuge: therefore the potential for interspecific competition for food exists.     

Tolerance to atmospheric ozone in transgenic tobacco over‐expressing glutathione synthetase in plastids

A cross between transgenic tobacco ( Nicotiana tabacum L.) plants which over‐expressed either γ‐glutamylcysteine synthetase (cpGSHI) or glutathione synthetase (cpGSHII) in their chloroplasts was used to compare the consequences of over‐expression of components of the glutathione synthetic pathway on tolerance to atmospheric O₃ or paraquat. A high proportion (50%) of those progeny which carried the cpGSHII transgene alone showed tolerance to atmospheric O₃ but not to paraquat. Progeny of an additional two, independent, self‐pollinated primary transgenic lines, which segregated in a Mendelian fashion for the presence of the cpGSHII transgene and therefore included both transformed and non‐transformed (recessive, wild‐type) plants, were also challenged by fumigation with O₃. Again, in both cases, about 50% of the plants expressing the epGSHII transgene were found to be O₃‐tolerant on the basis of reduced ethylene emissions and increased or unchanged total pigment concentrations. However, this tolerance was not due to specific changes in stomatal densities.     

Seedling growth response to added nutrients depends on seed size in three woody genera

1 We tested whether seedlings of small‐seeded species were more reliant on soil nutrients than large‐seeded ones by growing 21 species from three woody genera ( Eucalyptus, Hakea and Banksia ) along a gradient of nutrient availability.
2 At very low nutrient availability, larger seeds produced larger seedlings. This was seen especially among the eucalypts, but the difference was eliminated at optimal soil nutrient levels. Hakea species with large seed mass, and all Banksia species, appeared unable to exploit additional soil nutrients for growth, whatever the level supplied.
3 Larger seeds tended to have proportionately higher contents of N, P and K and, under nutrient‐poor conditions, supplied more of these to their seedlings, although at a diminishing rate.
4 We suggest that large‐seededness could be an adaptation to the high‐light, nutrient‐impoverished habitats in which these species occur by providing the seedling with the mineral nutrients, rather than carbon‐based metabolites, needed for maximizing initial root growth. Reaching reliable moisture before summer (drought avoidance) is an alternative strategy to physiological tolerance of drought.     

SPATIAL AND TEMPORAL VARIATION IN CHASMOGAMY AND CLEISTOGAMY IN OXALIS MONTANA (OXALIDACEAE)

Chasmogamous (CH) and cleistogamous (CL) flower production was investigated in natural populations of the perennial herb Oxalis montana in southern Quebec, Canada. Every 10–12 days during two flowering seasons, we recorded the reproductive output of approximately 2,000 plants distributed among five forest sites. The percentage of plants flowering, proportion of flowering plants producing CH and CL flowers, CH and CL flower number per flowering plant, and the proportion of flowers that are CL differed significantly among sites and corresponded to site differences in forest type and habitat quality. Unlike patterns in most CL species, however, reproduction by cleistogamy increased in sites with habitat conditions favorable to plant growth and sexual reproduction, and decreased in less productive sites. Chasmogamous and CL flower production increased with increasing plant size but plant size explained a significantly greater proportion of the variation in CL flower numbers. The percentage of flowering plants producing CH flowers decreased between years while the proportion of CL flowers increased at all sites during the second flowering season. The somewhat unusual aspects of reproduction in Oxalis may stem from its perennial habit which allows use of stored resources in adjusting the balance of cleistogamy versus chasmogamy in different environmental regimes.     

Carbon source for reproduction in a spring ephemeral herb, Corydalis ambigua (Papaveraceae)

1.  The carbon source for reproduction in plants may differ between flowering and fruiting stages. To clarify how spring ephemerals use current photosynthetic products for reproduction, the allocation patterns of photosynthate at flowering and fruiting and the effects of resource limitation on reproductive performance in Corydalis ambigua were assessed.
2.  A ¹³C tracing experiment revealed that about 20% of the current photosynthetic carbon was used for reproduction at both flowering and fruiting. The proportion of ¹³C allocated to fruits was constant irrespective of the light level. In contrast, ¹³C translocation to tubers increased at fruiting, and this trend was accelerated when plants were shaded.
3.  Defoliation treatment significantly reduced nectar production and tuber mass, while seed production was not affected. Therefore, when carbon assimilation was limited, carbon was preferentially allocated to current reproduction (seeds) rather than to pollinator attraction (nectar) or storage (tuber).
4.  If seed production is partly supported by carbohydrate reserved in the old tissue of tubers, nectar and seed production may not compete strongly for carbon sources. In contrast to the ability of high seed production, the susceptibility of nectar production to current photosynthesis indicates that seed production of this species is basically limited by pollen capture.
5.  Therefore, temporal separation of resource pool for reproduction may mitigate the joint limitation of seed production between pollinator attraction and resource availability. Temporal variation of the sink–source balance of storage organ is crucial to understand the cost of reproduction in perennial plants.     

Consequences of preformation for dynamic resource allocation by a carnivorous herb,Pinguicula vulgaris (Lentibulariaceae)

When resource availability changes frequently and unpredictably, natural selection favors flexible resource allocation; however, such versatility may be compromised in perennial plants that differentiate leaves or flowers a year in advance of their development (preformation). We investigated resource allocation by the carnivorous perennial Pinguicula vulgaris to determine whether increased resource availability changes within-season allocation to growth, vegetative propagation, and reproduction. In response to resource supplementation (feeding with fruit flies), plants attained a mass 60% greater than that of unfed plants after a single growing season. Feeding also enhanced vegetative propagation, which is closely associated with growth, without modifying relations between these two vegetative functions. In contrast, feeding did not alter the size of vegetative rosettes or the within-season incidence of either flowering or fruiting. This lack of immediate responses occurred because floral and leaf primordia differentiated up to 10 mo before resource supplementation and flower development. However, enhanced resource status likely affected future reproduction indirectly through resource effects on plant size. Large plants produced more floral primordia and between-year changes in fruiting status corresponded to changes in plant size. These results illustrate that preformation can delay responses to enhanced resources by perennial plants.     

Nutrient deficiency promotes male-biased apparent sex ratios at the ramet level in the dioecious plant <Emphasis Type="Italic">Myrica gale</Emphasis> var. <Emphasis Type="Italic">tomentosa</Emphasis> in oligotrophic environments in bogs

In populations of dioecious plants, the differences in the cost of reproduction between male and female plants can promote a male-biased sex ratio. In this study, we examine the macronutrient levels in tissues of the dioecious wetland shrub Myrica gale to identify the cost of reproduction for male and female plants and to examine the effect of nutrients on the apparent sex ratio at the ramet level. We examined plants across 12 populations of M. gale inhabiting bogs and fens in Japan. For each population, we used line transects to estimate the apparent sex ratio and measured the concentrations of nitrogen (N), phosphorus (P), and potassium (K) in the leaves sampled from male and female plants and in the fruits from female plants. For five of the populations, we calculated the flowering frequency, mortality, and the recruitment rate (as the rate of clonal propagation). We found that the proportion of females was positively affected, and the male bias of sex ratios reduced, by increases in P concentration in leaves sampled from female plants. Neither mortality nor recruitment was affected by sex or by the nutrient concentration (P, K). The flowering frequency was not affected by sex or by K concentration, but decreased with decreases in the P concentration measured in leaves. This study confirmed that reproduction in M. gale is P-limited. We found no distinct differences in the flowering frequency, mortality, or recruitment rate between the male and female plants.     

Host shifting by Operophtera brumata into novel environments leads to population differentiation in life-history traits

Abstract.  1. Operophtera brumata L. (Lepidoptera: Geometridae), a polyphagous herbivore usually associated with deciduous trees such as oak Quercus robur L . , has expanded its host range to include the evergreen species heather Calluna vulgaris (L.) Hull and, most recently, Sitka spruce Picea sitchensis (Bong.) Carrière.
2. Phenology, morphology, and survival of O. brumata were measured at several life-history stages in populations from the three different host plant communities sampled from a range of geographical locations. The data were used to test for population differences, reflecting the marked differences in host-plant secondary chemistry, growth form, and site factors such as climate. The hypothesis that spruce-feeding populations originated from populations feeding on moorland, commonly sites of coniferous afforestation, was also tested.
3. Altitude, not host plant species, was the major influence on the timing of adult emergence. An effect of insect population independent of altitude was found, implying that additional unidentified factors contribute to this phenological variation. Larval survival and adult size varied between populations reared on different host plant species. Survival of larvae was affected negatively when reared on the novel host plant, Sitka spruce, versus the natal plant (oak or heather) but oak and heather-sourced insects did not differ in survivorship on Sitka spruce.
4. Host range extension into novel environments has resulted in population differentiation to the local climate, demonstrating that host shifts pose challenges to the herbivore population greater than those offered by the host plant alone. The hypothesis that Sitka spruce feeding populations have arisen predominantly from moorland feeding populations was not supported.     

Non-structural carbohydrate accumulation and use in an understorey rain-forest shrub and relevance for the impact of leaf herbivory

1. Piper arieianum, an evergreen, understorey shrub of lowland moist forests of Central and South America, exhibits marked seasonal variation in reproductive activity even though climatic variation is low at the study site. Despite a lack of climatic seasonality, previous experimental leaf removal suggested that carbohydrate accumulation is seasonal, occurring prior to flowering.
2. We first tested the hypothesis that carbohydrates necessary for reproduction are accumulated prior to flowering, rather than during or after. By measuring non-structural carbohydrate production in the form of glucose and starch we found that the concentration of these reserves is greatest 1–3months before flowering, decreasing by 50% during peak fruit maturation.
3. The hypothesis that reproduction was the cause of this decrease in carbohydrate reserves was then tested by comparing reserves in plants that were prevented from flowering with those that flowered and produced fruit naturally. As predicted, reserves declined more in flowering than in non-flowering plants. A smaller decline in reserves of non-flowering plants was accompanied by greater stem and leaf production, suggesting that stored carbohydrates are also required for growth.
4. Because concentrations of non-structural carbohydrates were similar in roots, stems and leaves, and because the greatest amount of plant biomass is in stems for plants of a range of sizes, stems appear to be the main storage site of carbohydrate reserves in this plant species.
5. These results, together with previous studies, demonstrate that the impact of leaf herbivory on seed production in P. arieianum depends on the timing of that herbivory relative to the accumulation and use of non-structural carbohydrates.     

Air‐breathing behaviour of the African anabantoid fish Ctenopoma muriei

Frequency of aerial respiration in Ctenopoma muriei increased as dissolved oxygen (DO) decreased; breathing behaviour was spatially synchronous, and there was no difference in air‐breathing frequency between populations from a high‐ and low‐DO site. Although C. muriei survived without access to the surface, its loss of buoyancy and equilibrium under these conditions suggested that it is a functionally obligatory air breather.     

Environmental gradients and the horizontal distribution of microcrustaceans in lakes

1. The assemblage of suspension‐feeding microcrustaceans in lakes changes along a habitat gradient from nearshore to offshore. This gradient of microcrustaceans was explored in relation to differences in macrophytes and the associated changes in water chemistry, food resources and types of predators.
2. Some microcrustacean species were littoral or limnetic specialists, while others changed their distribution along this horizontal habitat gradient on a diel or seasonal basis. Distribution patterns were similar in a lake and a pond which differed in extent of macrophyte habitat.
3. There was a large shift in the composition of sestonic food, indicating heterotrophic seston nearshore and more autotrophic seston offshore. Sit‐and‐wait predators of microcrustaceans (e.g. Enallagma spp.) dominated nearshore and cruising predators (e.g. Leptodora kindtii Focke) were more common offshore.
4. A reciprocal transplant experiment revealed that littoral specialists could survive equally well when fed littoral or limnetic seston, while limnetic specialists performed poorly when fed littoral seston. Food resources may be important in determining where some microcrustacean species live along this horizontal habitat gradient.     

Population structure of the freshwater flatworm Crenobia alpina (Dana): old lineages and low gene flow

Aim  The freshwater flatworm Crenobia alpina lives almost exclusively in headwaters of mountainous areas and is supposed to be a glacial relict. We examined genetic diversity within and between populations of C. alpina in order to determine the taxonomic status of purported subspecies and to understand large-scale biogeographical patterns of glacial relicts.
Location  Central Europe.
Methods  We analysed mitochondrial DNA sequences and polymorphic allozyme loci of C. alpina populations across its range in central Europe. Sequences were compared using parsimony, minimum evolution and maximum likelihood. Allozymes were analysed using traditional as well as Bayesian estimates of F statistics.
Results  We found considerable divergence between haplotypes. For each of the two lineages occurring throughout central Europe, allozymes showed considerable differentiation between populations and a strong isolation by distance effect. Hence populations are effectively isolated even across rather small spatial scales.
Main conclusions  There is strong evidence that C. alpina is a complex of distinct lineages or cryptic species that date back to the late Miocene. The separation of lineages may be associated with the formation of deep valleys at the end of the Messinian Crisis.     

Colonising aliens: caterpillars (Lepidoptera) feeding on Piper aduncum and P. umbellatum in rainforests of Papua New Guinea

Abstract.  1. Caterpillar assemblages feeding on two alien plants, Piper aduncum and P. umbellatum , were studied in lowland rainforest in Papua New Guinea and compared with assemblages from 69 species of native woody hosts, including congeneric P. macropiper .
2. Species richness of caterpillars feeding on P. aduncum (29 species per 1500 m² of foliage) and P. umbellatum (36 species) was higher than the median richness for the 69 native hosts (23 species).
3. The probability that a caterpillar species colonised alien Piper increased with its host range from 3% for the species feeding on a single plant family to 92% for the species with host range >10 plant families.
4. The assemblage on P. aduncum was dominated by a single species ( Herpetogramma sp. near licarsisalis , Crambidae), which represented 48% of individuals, and also had a high proportion (34%) of rare species, collected as single individuals. This community structure was indistinguishable from that of a typical native host. In contrast, the P. umbellatum assemblage was unusual as no species represented >10% of individuals.
5. The aggressive invasion by P. aduncum of early successional vegetation is not explained by a competitive advantage due to low herbivore load, as the abundance of caterpillars feeding on it was comparable to that of native pioneer plants.
6. The caterpillar assemblage on P. aduncum demonstrated that an assemblage indistinguishable from native assemblages in density, species richness, and dominance structure (but not in host specificity) can originate from the existing species pool in lowland rainforests on a recently established tree species in <50 years.