Contagious dispersal of seeds of synchronously fruiting species beneath invasive and native fleshy‐fruited trees

In subtropical Australia, many native and invasive plant species rely on a shared suite of frugivores, largely birds, for seed dispersal. Many native plants fruit during summer in this region, whereas most invasive plants fruit during winter, thus providing the opportunity for contagious dispersal of seeds beneath synchronously fruiting species. We sampled invasive and native seed rain beneath the canopy of a native summer‐fruiting tree Guioa semiglauca and an invasive winter‐fruiting tree Cinnamomum camphora, in three study sites over the course of a year. In July, during peak fruiting season for C. camphora and other invasive species, seed rain of invasive species was higher beneath C. camphora than G. semiglauca. This was partly due to the invasive tree Ligustrum lucidum, whose seed rain was three times higher beneath C. camphora than beneath the native tree. In February, seed rain of native species was more abundant beneath the canopy of G. semiglauca than beneath C. camphora, despite the fact that C. camphora was also fruiting at this time. This was probably due to the larger fruit crop produced by G. semiglauca at this time of year. Our study provides evidence that the presence of invasive bird‐dispersed plants may facilitate contagious seed dispersal of other invaders, and likewise native species may facilitate seed spread of other native plants.     

Intraspecific Variation of Floral Scent Chemistry in Magnolia kobus DC. (Magnoliaceae)

Magnolia kobus was examined at 32 sites in Japan (109 female-stage flowers from 52 plants) by GC-MS. Major chemical compounds (a total of 36 chemicals) emitted from the flowers were: linalool (and its oxides), limonene, cis- and trans-β-Ocimene, benzaldehyde, benzyl alcohol, benzyl cyanide, and 2-aminobenzaldehyde. Linalool and its oxides were the most abundant components of floral scents in 21 individuals. The rate at which chemical volatiles were emitted ranged from 0.002 to 0.929 μg/flower/hour (average 0.211). High quantitative and qualitative variation in floral scent chemistry among individuals was found throughout the range of M. kobus, especially in central Honshu. The high variability in floral scent chemistry may be due to the importance of visual cues in the reproductive biology of M. kobus which flowers in early spring, resulting in decreased selection for specific floral scent profiles. Alternatively, different scent compounds or chemical profiles may be equally effective in attracting pollinators. Received 25 June 2001/ Accepted in revised form 25 August 2001     

Thinning operations increase the demographic performance of the rare subtree species <Emphasis Type="Italic">Magnolia stellata</Emphasis> in a suburban forest landscape

To ensure the persistence of populations and species that maintain unique biodiversity in suburban landscapes, it is necessary to establish an efficient management system for abandoned suburban secondary forests. Thinning operations seem to be an effective management approach to inhibiting the progress of forest succession, which could lower the level of species diversity. To evaluate the effect of thinning operations on the demographic performance (growth, survival, and sexual reproduction) of the rare subtree species Magnolia stellata, we set up a field experiment and monitored the population dynamics over 9 years. The results revealed that stem survival and flower production per genet showed a decreasing trend in the nonthinned site. However, thinning operations increased the demographic performance, in particular the stem growth and survival rates as well as flower and seed production. The findings suggested that thinning operations may decrease the extinction probability of not only the targeted local population but also the metapopulation, because the thinned population serves as a better seed source. Thus, the creation of well-lit sites by thinning operations would be a useful conservation strategy for M. stellata.     

Asymmetric introgression between Magnolia stellata and M. salicifolia at a site where the two species grow sympatrically

In order to understand the ongoing evolutionary relationships between species, it is important to elucidate patterns of natural hybridization. In the zone where two species are sympatrically distributed, we examined 274 individuals of Magnolia stellata, Magnolia salicifolia, and their putative hybrids by means of 16 nuclear and three chloroplast microsatellite markers. Hybrid classes of individuals were estimated by admixture analyses. Morphological traits were also investigated for 64 of the 274 individuals. Admixture analyses revealed that 66 of the 274 individuals were classified as hybrids, comprising 17 F₁ and 19 F₂ individuals, 27 backcrosses to M. salicifolia, and 3 individuals of unknown origin. Morphological data from the 64 individuals agreed well with their genetic admixture rates. Spatial locations of F₁ and F₂ hybrids at the study site were intermediate between the two purebred species, indicating that the site preferences of hybrids are intermediate. The occurrences of F₂ and backcross hybrids indicate that F₁ hybrids are fertile. The chloroplast DNA haplotypes of all F₁ hybrids corresponded to those detected in M. salicifolia, so that maternal parents of the F₁ hybrids were all M. salicifolia. Furthermore, no hybrid individuals derived from a backcross to M. stellata were detected. These results suggest that the direction of hybridization and the subsequent introgression have been quite asymmetric and that the introgression occurred from M. stellata into M. salicifolia.     

The role of ABC genes in shaping perianth phenotype in the basal angiosperm Magnolia

It is generally accepted that the genus Magnolia is characterised by an undifferentiated perianth, typically organised into three whorls of nearly identical tepals. In some species, however, we encountered interesting and significant perianth modifications. In Magnolia acuminata, M. liliiflora and M. stellata the perianth elements of the first whorl are visually different from the others. In M. stellata the additional, spirally arranged perianth elements are present above the first three whorls, which suggests that they have been formed within the domain of stamen primordia. In these three species, we analysed expression patterns of the key flower genes (AP1, AGL6, AP3, PI, AG) responsible for the identity of flower elements and correlated them with results of morphological and anatomical investigations. In all studied species the elements of the first whorl lacked the identity of petals (lack of AP3 and PI expression) but also that of leaves (presence of AGL6 expression), and this seems to prove their sepal character. The analysis of additional perianth elements of M. stellata, spirally arranged on the elongated floral axis, revealed overlapping and reduced activity of genes involved in specification of the identity of the perianth (AGL6) but also of generative parts (AG), even though no clear gradient of morphological changes could be observed. In conclusion, Magnolia genus is capable of forming, in some species, a perianth differentiated into a calyx (sepals) and corolla (petals). Spirally arranged, additional perianth elements of M. stellata, despite activity of AG falling basipetally, resemble petals.     

Reproduction of invasive Amur honeysuckle (Lonicera maackii) and the arithmetic of an extermination strategy

Ecological restorations often require removal of invasive species. The abundance of invasives has tended to catalyze research emphasizing removal, not broader understandings, of species mechanisms for persistence in the landscape (e.g. reproductive output and seed dispersal). Asiatic shrub honeysuckles (Lonicera spp.) are pernicious invaders throughout eastern North America. Heavy tree canopy cover apparently reduces growth and reproductive output in Lonicera maackii, which is widespread through the lower Midwestern United States. To help focus control efforts more effectively, we quantified the effect of tree canopy cover on vegetative growth, flowering, and fruit production under three canopy densities. Mean vegetative growth of flowering shoots was not affected by canopy cover. All aspects of sexual reproduction (flower production, fruit set, fruit number, fruit mass, seed number, and seed size) were strongly reduced by moderate shade. Although all individuals modify community and ecosystem properties, a limited number of high light individuals might also provide the greatest proportion of the seeds. Through model simulation of honeysuckle population structure in relation to canopy cover, we argue that it can sometimes be more efficient to initially target reproductive individuals in the high light edge and interior gap environments than to immediately focus on all individuals in the forest interior.     

Population genetic structure and demography of Magnolia kobus: variety borealis is not supported genetically

Species delimitations by morphological and by genetic markers are not always congruent. Magnolia kobus consists of two morphologically different varieties, kobus and borealis. The latter variety is characterized by larger leaves than the former. For the conservation of M. kobus genetic resources in natural forests, the relationships between morphological and genetic variation should be clarified. We investigated variations in nuclear microsatellites, chloroplast DNA (cpDNA) sequences and leaf morphological traits in 23 populations of M. kobus over the range of species. Two genetically divergent lineages, northern and southern were detected and their geographical boundary was estimated to be at 39°N. The northern lineage consisted of two genetic clusters and a single cpDNA haplotype, while the southern one had multiple genetic clusters and cpDNA haplotypes. The northern lineage showed significantly lower genetic diversity than the southern. Approximate Bayesian computation indicated that the northern and southern lineages had experienced, respectively, population expansion and long-term stable population size. The divergence time between the two lineages was estimated to be 565,000 years ago and no signature of migration between the two lineages after divergence was detected. Ecological niche modeling showed that the potential distribution area in northern Japan at the last glacial maximum was very small. It is thus considered that the two lineages have experienced different population histories over several glacial-inter-glacial cycles. Individuals of populations in the central to northern part of Honshu on the Sea of Japan side and in Hokkaido had large leaf width and area. These leaf characteristics corresponded with those of variety borealis. However, the delimitation of the northern and southern lineages detected by genetic markers (39°N) was not congruent with that detected by leaf morphologies (36°N). It is therefore suggested that variety borealis is not supported genetically and the northern and southern lineages should be considered separately when identifying conservation units based not on morphology but on genetic markers.

     

Genetic Structure in Remnant Populations of an Endangered Andean Magnolia

By genetically characterizing adult individuals, we documented the presumed prefragmentation population structure of an endangered Andean tree, Magnolia espinalii. Our sample included all known surviving individuals for this species. Genetic diversity was high, but genetic distance was not related to geographic distance between sites. Magnolia espinalii apparently was originally distributed in numerous semi‐isolated demes. The lack of evidence of recruitment around most adults in this species suggests a fragmentation‐related disruption of their reproductive ecology. Artificial propagation projects should obviously attempt to involve all 23 known surviving trees.     

Isolation and Structure Elucidation of Growth Inhibitors on Silkworm Larvae from Magnolia kobus DC

Two lignans were isolated from leaves of Magnolia kobus DC. as growth inhibitors on silkworm larvae and structurally elucidated as sesamin (I) and kobusin (II) which has been hitherto unknown.     

Spatial distribution and performance of native and invasive <Emphasis Type="Italic">Ardisia</Emphasis> (Myrsinaceae) species in Puerto Rico: the anatomy of an invasion

Comparisons between native and invasive congeners are potentially useful approaches for identifying characteristics that promote invasiveness. Those traits for which an invasive exhibits superior ecological performance are likely to contribute to its invasiveness. We tested the hypothesis that invasive tree species have better ecological performance in early life cycle stages than native species in forests where they coexist. We studied locally sympatric populations of the invasive Ardisia elliptica and the native A. obovata (Myrsinaceae) in Puerto Rico. We compared spatial distribution, herbivory and growth in seedlings, seed germination in the field and under controlled conditions, and fruit production. We found the distribution of each species was aggregated in the three categories of size (seedlings, juveniles and adults) and the populations partially overlapped. The invasive species was the most abundant species in every category of size. The two species did not differ in percentage of leaf area consumed and seedlings of both species had the same relative growth rate (RGR) in the forest. However, the invasive species had higher germination success in the field, faster mean germination time in the lab and higher fruit production. It appears that the success of A. elliptica is not through escape from pathogens or herbivores, but by a better performance in fruiting and seed germination in the forest.     

An invasive tree alters the structure of seed dispersal networks between birds and plants in French Polynesia

Aim We studied how the abundance of the highly invasive fruit‐bearing tree Miconia calvescens DC. influences seed dispersal networks and the foraging patterns of three avian frugivores. Location Tahiti and Moorea, French Polynesia. Methods Our study was conducted at six sites which vary in the abundance of M. calvescens. We used dietary data from three frugivores (two introduced, one endemic) to determine whether patterns of fruit consumption are related to invasive tree abundance. We constructed seed dispersal networks for each island to evaluate how patterns of interaction between frugivores and plants shift at highly invaded sites. Results Two frugivores increased consumption of M. calvescens fruit at highly invaded sites and decreased consumption of other dietary items. The endemic fruit dove, Ptilinopus purpuratus, consumed more native fruit than either of the two introduced frugivores (the red‐vented bulbul, Pycnonotus cafer, and the silvereye, Zosterops lateralis), and introduced frugivores showed a low potential to act as dispersers of native plants. Network patterns on the highly invaded island of Tahiti were dominated by introduced plants and birds, which were responsible for the majority of plant–frugivore interactions. Main conclusions Shifts in the diet of introduced birds, coupled with reduced populations of endemic frugivores, caused differences in properties of the seed dispersal network on the island of Tahiti compared to the less invaded island of Moorea. These results demonstrate that the presence of invasive fruit‐bearing plants and introduced frugivores can alter seed dispersal networks, and that the patterns of alteration depend both on the frugivore community and on the relative abundance of available fruit.     

Flowering and Fruiting Phenology of Magnolia hypoleuca

Abstract The floral biology of Magnolia hypoleuca , a tall tree species widely distributed in Japan, was studied in a deciduous broad-leaved forest in Hokkaido near the northern limit of distribution of the species. The flower is large, protogynous and nectarless. Its petals close after the female stage and again after the male stage. The mean duration of flowering of an individual flower is 3–4 days, while the flowering period of an individual tree lasts up to 40 days. The main pollinators of the flowers were beetles. Success of pollination is assumed to be affected by weather conditions at anthesis. Average fruit sets (ratios of fruits to flowers) were 13–25%. Hand-pollination of 44 flowers produced a significantly higher fruit set (28 fruits) than control flowers (33/133). These results suggest that the fruit set is limited by pollination. The frequency of infertile ovules was higher in controls than in hand-pollinated fruits, indicating that the seed set was also limited by pollination. We propose a mimicry hypothesis in which the non-rewarding female-stage flower mimics the rewarding male-stage flower to explain the short life-span of an individual flower, the long flowering period of an individual tree, and the coexistence of flowers of various stages on a single tree.     

Clonal growth and its effects on male and female reproductive success in <Emphasis Type="Italic">Prunus ssiori</Emphasis> (Rosaceae)

Clonal growth can increase not only floral display but also geitonogamy and may affect sexual reproduction both positively and negatively. A clonal woody species, Prunus ssiori, was partially self-incompatible according to a pollination experiment. Its main pollinators, bumble bees, were often observed to consecutively visit inflorescences within a tree. Clone identification revealed that its genets formed mutually exclusive patches. These features suggest frequent geitonogamous pollination. In a 6.24-ha plot, 212 trees belonged to 59 genets, and 42 genets consisted of a single tree, whereas the rest contained two or more clonal trees. The largest genet had 65 trees and occupied 0.4 of a hectare. Fruit set was measured in 127 inflorescences sampled from nine maternal trees at the center of the plot. Paternal genets of 107 of their 300 seeds were assigned in the plot using microsatellites. There were no selfed seeds. Male reproductive success (the probability that individual trees of each genet sired a seed) increased as tree size increased, as the distance between the trees and maternal trees decreased, when the genet did not contain the maternal trees, and when the genet consisted of a single tree. Female reproductive success (fruit set in individual inflorescences of each maternal tree) increased as the within-tree geitonogamy index, which reflected the frequency of pollination within the maternal tree, decreased. These results suggest that clonal growth reduces male reproductive success, at least, in P. ssiori, because of pollen discounting.     

Increased relative abundance of an invasive competitor for pollination, Lythrum salicaria, reduces seed number in Mimulus ringens

When exotic plant species share pollinators with native species, competition for pollination may lower the reproductive success of natives by reducing the frequency and/or quality of visits they receive. Exotic species often become numerically dominant in plant communities, and the relative abundance of these potential competitors for pollination may be an important determinant of their effects on the pollination and reproductive success of co-occurring native species. Our study experimentally tests whether the presence and abundance of an invasive exotic, Lythrum salicaria L. (Lythraceae), influences reproductive success of a co-flowering native species, Mimulus ringens L. (Phrymaceae). We also examine the mechanisms of competition for pollination and how they may be altered by changes in competitor abundance. We found that the presence of Lythrum salicaria lowered mean seed number in Mimulus ringens fruits. This effect was most pronounced when the invasive competitor was highly abundant, decreasing the number of seeds per fruit by 40% in 2006 and 33% in 2007. Reductions in the number of seeds per fruit were likely due to reduced visit quality resulting from Mimulus pollen loss when bees foraged on neighboring Lythrum plants. This study suggests that visit quality to natives may be influenced by the presence and abundance of invasive flowering plants.     

Reproductive output of invasive versus native plants

Aim Propagule size and output are critical for the ability of a plant species to colonize new environments. If invasive species have a greater reproductive output than native species (via more and/or larger seeds), then they will have a greater dispersal and establishment ability. Previous comparisons within plant genera, families or environments have conflicted over the differences in reproductive traits between native and invasive species. We went beyond a genus‐, family‐ or habitat‐specific approach and analysed data for plant reproductive traits from the global literature, to investigate whether: (1) seed mass and production differ between the original and introduced ranges of invasive species; (2) seed mass and production differ between invasives and natives; and (3) invasives produce more seeds per unit seed mass than natives. Location Global. Methods We combined an existing data set of native plant reproductive data with a new data compilation for invasive species. We used t‐tests to compare original and introduced range populations, two‐way ANOVAs to compare natives and invasives, and an ANCOVA to examine the relationship between seed mass and production for natives and invasives. The ANCOVA was performed again incorporating phylogenetically independent contrasts to overcome any phylogenetic bias in the data sets. Results Neither seed mass nor seed production of invasive species differed between their introduced and original ranges. We found no significant difference in seed mass between invasives and natives after growth form had been accounted for. Seed production was greater for invasive species overall and within herb and woody growth forms. For a given seed mass, invasive species produced 6.7‐fold (all species), 6.9‐fold (herbs only) and 26.1‐fold (woody species only) more seeds per individual per year than native species. The phylogenetic ANCOVA verified that this trend did not appear to be influenced by phylogenetic bias within either data set. Main conclusions This study provides the first global examination of both seed mass and production traits in native and invasive species. Invasive species express a strategy of greater seed production both overall and per unit seed mass compared with natives. The consequent increased likelihood of establishment from long‐distance seed dispersal may significantly contribute to the invasiveness of many exotic species.     

Magnolia tiepii sp. nov. from Vietnam

A new species of Magnolia sect. Gwillimia DC., Magnoliaceae from Vietnam is described and illustrated. Magnolia tiepii is recognized from southern Vietnam, where it occurs in Khanh Vinh slope, Khanh Vinh District, Khanh Hoa Province. It is distinguished from the closely related M. henryi by having narrowly cuneate leaf blade, short petiole, stipular scar at the base of the leaf blade, short peduncle and ellipsoid fruit.     

Effects of invasive Green Iguanas (<Emphasis Type="Italic">Iguana iguana</Emphasis>) on seed germination and seed dispersal potential in southeastern Puerto Rico

Green Iguanas (Iguana iguana) are invasive in Puerto Rico due to a variety of negative economic effects, yet we know very little about their ecological impacts. Because they are herbivorous, defecate intact seeds, move through the forest, and have long gut-passage times, Green Iguanas may affect seed germination and seed dispersal. In summer 2013, a total of 258 Green Iguana scat samples were collected at the Humacao Natural Reserve in southeastern Puerto Rico. Seeds extracted from scat and collected from fruit were planted under common garden conditions using experimental treatments designed to tease apart the effects of feces, fruit, and ingestion on seed germination. Green Iguanas decreased the time for seeds to germinate in Ficus spp. by removing fruit pulp, but had no effect on germination of native Annona glabra seeds. For non-native P. pterocarpus and Pterocarpus spp., Green Iguanas produced conflicting results, decreasing the percentage of seeds germinating, but at the same time, reducing the time for seeds to germinate. Green Iguanas likely disperse most seeds beyond the canopies of parental tree at our site. Government and economic resources are being used to eradicate Green Iguana populations in Puerto Rico, but the lack of consistent effects of Green Iguanas on seed germination for the plant species consumed at our site complicates generalizing about their ecological effects and developing management plans that minimize negative effects for native plant communities. We recommend additional studies that target both species of particular concern, such as threatened native or invasive species, as well as studies of sensitive habitats in Puerto Rico.     

Evaluation of a field experiment for the conservation of a <Emphasis Type="Italic">Magnolia stellata</Emphasis> stand using clear-cutting

Magnolia stellata is a rare subcanopy tree species that grows in secondary forests in warm temperate zones. It is now endangered due to habitat degradation by vegetation succession. In an attempt to improve the habitat, a 30 m?×?10 m plot (0.03 ha) was set up with all vegetation including M. stellata being clear-cut in January 2012. The number of sprouts increased for 1–2 years after clear-cutting and then gradually decreased or remained constant. Five years after clear-cutting, the numbers of individuals and stems, and the total basal area (BA), were 87.0, 165.5 and 3.2%, respectively, of the values before clear-cutting. BA was highest for Ilex pedunculosa, followed by M. stellata and Hydrangea paniculata. Some sprouted individuals of M. stellata produced flower buds in the second year after clear-cutting, and flowered and fruited in the spring and summer of the third year, respectively. The densities of potential canopy species were 18,533 ha^?1 (height >?0.5 m) and 7,267 ha^?1 (height >?1.2 m), vastly exceeding the value of the criterion for successful natural regeneration after clear-cutting of warm temperate forests in the region (3,000 ha^?1). Based on this criterion, it is thus considered that the natural regeneration has reached completion. However, 45.1% (height >?0.5 m) and 95.5% (height >?1.2 m) of M. stellata individuals were regenerated by sprouting. Further research is needed into how individuals, regenerated from seedlings, develop and reach sexual maturity, and how successive generations change.     

Prolific fruit output by the invasive tree Bellucia pentamera Naudin (Melastomataceae) is enhanced by selective logging disturbance

Selective logging in tropical rain forests may promote population growth of invasive plants. The ability of invaders to respond, specifically in reproductive traits, to increase in resource abundance may allow them to increase their presence in the seed rain of recipient communities. The invasive pioneer tree Bellucia pentamera (Melastomataceae) is currently spreading within Gunung Palung National Park in West Kalimantan, Indonesia. The park has also experienced periods of illegal, small‐scale, selective logging that seem to have facilitated population growth and spread of this invader. We first used fruiting frequency as a proxy for fruit output to make comparisons between B. pentamera and the native tree community of over 200 genera. We then constructed two Generalized Linear Autoregressive Moving Average (GLARMA) models using 13 months of phenology data to predict both fruiting frequency and crop size of B. pentamera under selective logging versus natural treefall disturbance regimes. Bellucia pentamera fruited at considerably higher frequency than all 200 native genera considered. This invader also responded positively to selective logging with both odds of being in fruit and crop sizes at least doubling in logged plots. Prolific seed output of B. pentamera is especially problematic within the lottery competition of tropical rain forest gap tree communities, in which prominence in the seed rain is particularly important. Furthermore, the reproductive response of B. pentamera to selective logging suggests that this harvesting practice may have a considerable role in facilitating this invasion in Southeast Asia.     

Comparative demographic analysis in contrasting environments of Magnolia dealbata: an endangered species from Mexico

The study of population regulation and demography in natural habitats is critical for the conservation of rare and endangered species. We address the impact of cattle exclusion on the demographics of Magnolia dealbata (Magnoliaceae), an endangered species according to IUCN and Federal Mexican laws. Sixteen permanent plots were established, eight of which were enclosed to livestock, in the mountain cloud forest in Coyopolan, Mexico, which has the largest population of M. deadlbata. Censuses of the plots were undertaken annually during three annual cycles to record seed number, recruitment, mortality, and growth (defined as length and diameter at breast height). The effects of two treatments (with and without livestock exclusion) on the dynamics of M. dealbata were investigated using transition matrix models and life table response experiment (LTRE) analysis. Contrary to expectations, there was no significant effect of cattle exclusion on population growth rates (λ). Furthermore, the λ, estimated from the mean transition matrix for both treatments was greater than one. The transitions with the highest elasticity values were similar between the two treatments, while reproductive stage contributed more to differences in population growth rate and were less variable than the non-reproductive stage. LTRE analysis showed that treatment differences had little effect on λ. Livestock exclusion appears to lead more to differences in the arrangement of the values of the transition matrix than to the rate of population growth for M. dealbata.