Incorporating differences between genetic diversity of trees and herbaceous plants in conservation strategies.

Reviews that summarize the genetic diversity of plant species in relation to their life history and ecological traits show that forest trees have more genetic diversity at population and species levels than annuals or herbaceous perennials. In addition, among-population genetic differentiation is significantly lower in trees than in most herbaceous perennials and annuals. Possible reasons for these differences between trees and herbaceous perennials and annuals have not been discussed critically. Several traits, such as high rates of outcrossing, long-distance pollen and seed dispersal, large effective population sizes (Ne ), arborescent stature, low population density, longevity, overlapping generations, and occurrence in late successional communities, may make trees less sensitive to genetic bottlenecks and more resistant to habitat fragmentation or climate change. We recommend that guidelines for genetic conservation strategies be designed differently for tree species versus other types of plant species. Because most tree species fit an LH scenario (low [L] genetic differentiation and high [H] genetic diversity), tree seeds could be sourced from a few populations distributed across the species' range. For the in situ conservation of trees, translocation is a viable option to increase Ne . In contrast, rare herbaceous understory species are frequently HL (high differentiation and low diversity) species. Under the HL scenario, seeds should be taken from many populations with high genetic diversity. In situ conservation efforts for herbaceous plants should focus on protecting habitats because the typically small populations of these species are vulnerable to the loss of genetic diversity. The robust allozyme genetic diversity databases could be used to develop conservation strategies for species lacking genetic information. As a case study of reforestation with several tree species in denuded areas on the Korean Peninsula, we recommend the selection of local genotypes as suitable sources to prevent adverse effects and to insure the successful restoration in the long term.

Incorporación de diferencias de diversidad genética entre árboles y plantas herbáceas en estrategias de conservación Resumen Las revisiones que resumen la diversidad genética de las plantas en relación con sus características ecológicas y biológicas muestran que los árboles forestales tienen más diversidad genética a nivel de población y de especie que las plantas anuales o las perennes herbáceas. Sumado a esto, la diferenciación genética entre poblaciones es significativamente más baja en los árboles que en la mayoría de las perennes herbáceas y las anuales. Hasta la fecha no se han discutido críticamente las posibles explicaciones de estas diferencias entre los árboles y las perennes herbáceas y las plantas anuales. Varias características, como las tasas altas de alogamia, la dispersión a gran distancia de polen y semillas, el gran tamaño de la población efectiva (Ne ), la estatura arbórea, la baja densidad poblacional, la longevidad, el solapamiento de generaciones y la presencia dentro de comunidades sucesionales tardías, pueden generar en los árboles una menor sensibilidad a los cuellos de botella genéticos y una mayor resistencia a la fragmentación del hábitat o al cambio climático. Recomendamos que las directrices para las estrategias de conservación genética estén diseñadas de manera diferente para las especies arbóreas que para otro tipo de plantas. Ya que la mayoría de las especies arbóreas encajan dentro de un escenario LH (baja [L] diferenciación genética y alta [H] diversidad genética), las semillas de los árboles podrían tomarse de unas cuantas poblaciones dispersas a lo largo del área de distribución de la especie. Por lo anterior, para la conservación in situ de los árboles, la translocación es una opción viable para incrementar la Ne . Al contrario de esta situación, las especies herbáceas raras del sotobosque con frecuencia son especies HL (alta diferenciación y baja diversidad). En el escenario HL, las semillas deberían ser recolectadas de muchas poblaciones con diversidad genética alta y los esfuerzos de conservación in situ para las plantas herbáceas deberían enfocarse en la protección del hábitat ya que las poblaciones típicamente pequeñas de estas especies son vulnerables a la pérdida de la diversidad genética. Las robustas bases de datos de diversidad genética aloenzimática podrían usarse para desarrollar estrategias de conservación para las especies que carecen de información genética. Como caso de estudio de reforestación con varias especies arbóreas en áreas deforestadas de la Península de Corea, recomendamos la selección de genotipos locales como fuente adecuada para prevenir los efectos adversos y para asegurar la restauración exitosa a largo plazo.

An unprotected vulnerable relict subtropical conifer-Keteleeria evelyniana: Its forests, populations, growth and endangerment by invasive alien plant species in China.

Relict subtropical coniferous forests in China face severe fragmentation, resulting in declining populations, and some are under significant threat from invasive alien species. Despite the crucial importance of understanding forest dynamics, knowledge gaps persist, particularly regarding the impact of invasive plants on vulnerable natives like Keteleeria evelyniana. In this study, we investigated the impact of invasive plants on the regeneration of forests dominated by K. evelyniana, a subtropical relict species in southwestern China. For this purpose, we characterized forest dynamics of 160 forest plots featuring K. evelyniana as the primary dominant species and determined whether the presence of invasive plants was correlated with regeneration of K. evelyniana. We identified four distinct forest types in which K. evelyniana was dominant. We found that radial growth of K. evelyniana trees is faster in younger age-classes today than it was for older trees at the same age. The population structure of K. evelyniana in each forest type exhibited a multimodal age-class distribution. However, three forest types lacked established saplings younger than 10 years old, a situation attributed to the dense coverage of the invasive alien Ageratina adenophora. This invasive species resulted in a reduction of understory species diversity. Additionally, our analysis uncovered a significant negative correlation in phylogenetic relatedness (net relatedness index) between native and invasive alien plant species in eastern Yunnan. This suggests closely related invasive species face heightened competition, hindering successful invasion. Taken together, our findings indicate that successful establishment and habitat restoration of K. evelyniana seedling/saplings require effective measures to control invasive plants.

Survival of a tertiary relict species, Liriodendron chinense (Magnoliaceae), in southern China, with special reference to village fengshui forests.

PREMISE OF THE STUDY: We investigate factors supporting the persistence in southern China of a rare Tertiary relict tree species, Liriodendron chinense, which has been almost eliminated by recent land use conversion. We hypothesize that cultural practices and traditional sustainable forest resource uses provide niches for the species' regeneration that will complement infrequent natural disturbances, while the species' survival on remote mountain slopes where there are no humans depends on natural disturbances alone. • METHODS: We examined and analyzed various landscape contexts, community associations, age distributions, and regeneration patterns of Liriodendron chinense. • KEY RESULTS: Forest communities containing Liriodendron chinense were of three types: (1) village fengshui forests-mature forests dominated by Tertiary relict taxa Liriodendron, Toona, and Emmenopterys, protected for their supposed spiritual value; (2) young secondary forests near villages, dominated solely by Liriodendron; and (3) old secondary forest remnants on mountain slopes far from villages, dominated by Liriodendron with other Tertiary relicts of the genera Davidia and Sassafras. The age structure of Liriodendron indicated ample recruitment in the first two forest types, where the activities of local people have provided regeneration niches for the survival of this shade-intolerant pioneer species. On the remote mountain slopes that have never been converted to agriculture, Liriodendron has survived through regeneration made possible by natural disturbances. • CONCLUSIONS: The traditional human land use, influenced by cultural values, has supplemented infrequent natural disturbances, providing regeneration niches for this and other Tertiary remnant species near villages in mountain valleys, while on uninhabited mountain slopes the species depends on natural disturbances to survive.

Population persistence of a Tertiary relict tree Tetracentron sinense on the Ailao Mountains, Yunnan, China.

The persistence of the Tertiary relict tree Tetracentron sinense Oliv. on the eastern slope of the Ailao Mountains, Yunnan, SW China, was here studied in terms of population structure (size, age) and regeneration patterns. T. sinense occurred in unstable habitats by stream banks, on steep slopes, on scree slopes, or on roadsides near streams in narrow valleys, all places subject to frequent natural disturbances, whereas none were found on stable gentle slopes free of major disturbances at similar altitudes. Further, no established saplings of T. sinense were found in forests having high bamboo (Yushania crassicollis Yi) coverage in their understory. The size and age structure of T. sinense were multimodal. The reproduction of the tree was either by means of abundant minute wind-dispersed seeds or by resprouts in unstable habitats. These populations depended on disturbance or gap regeneration to survive. T. sinense, along with other tree life-forms including evergreen broad-leaved species and conifers, dominated in the forest canopy layer, even reaching the emergent layer in places. Results of the study provide insight into the ecological characteristics and survival mechanisms of this East Asian paleoendemic tree species. The study will provide a scientific basis for recommendations for the conservation of this species and for other Tertiary relict plants having similar regeneration dynamics.

Cut to Disarm Plant Defence: A Unique Oviposition Behaviour in Rhynchites foveipennis (Coleoptera: Attelabidae).

Female weevils of the family Attelabidae (Coleoptera: Curculionoidea) possess a unique behaviour of partially cutting the branches connecting egg-bearing organs of their host plants during oviposition. However, the consequence of such behaviour remains unclear. Using Rhynchites foveipennis and its host pear (Pyrus pyrifolia), the present study tested the hypothesis that the oviposition behaviour could disarm the host plants' defence. We compared the survival rates, growth rates, and performance of eggs and larvae under two conditions: (1) the fruit stems were naturally damaged by the females before and after oviposition, and (2) the fruit stems were artificially protected from the females. When fruit stems were protected from female damage, the survival rates of eggs and larvae were only 21.3-32.6%, respectively; and the larval weight was 3.2-4.1 mg 30 days after laying eggs. When the fruit stems were damaged, the survival rates of eggs and larvae reached 86.1-94.0%, respectively; and the larval weight reached 73.0-74.9 mg 30 days after laying eggs. The contents of tannin and flavonoids in the pears did not change significantly along with the oviposition and larval feeding, but weevil eggs were crushed and killed by the callus in the pears. Once the stunted larvae in branch-growing pears were moved into the picked-off ones, the growth and development recovered. The findings indicate that the oviposition behaviour can significantly increase the survival of the offspring. Our study suggested that the oviposition behaviour of attelabid weevils is a strategy to overcome plant defence.

Forest characteristics, population structure and growth trends of threatened relict Pseudotsuga forrestii in China.

Pseudotsuga forrestii is a relict evergreen coniferous tree species in Pinaceae endemic to China. P. forrestii tree numbers have greatly decreased due to deforestation, over-utilization and habitat degradation. Here we clarify P. forrestii community types, structure, species diversity, seedling recruitments and growth trends. We identified four P. forrestii community types: (1) Pseudotsuga forrestii - Quercus guyavifolia - Acer davidii evergreen coniferous and broad-leaved mixed forest; (2) Pseudotsuga forrestii - Pinus yunnanensis - Quercus guyavifolia evergreen coniferous and broad-leaved mixed forest; (3) Pseudotsuga forrestii evergreen coniferous forest; (4) Pseudotsuga forrestii - Abies georgei var. smithii evergreen coniferous forest. P. forrestii forests are characterized by both warm temperate and temperate affinities. Simpson diversity, Pielou evenness, Shannon-Wiener diversity indices ranged from 0.75 to 0.76, 0.74-0.81, and 1.62-1.93, respectively, with no significant differences among the four forest types. The forest stratification was multilayered. The canopy layer was generally 10-25 m tall, with the emergent layer reached 25-42 m. DBH and age structures of P. forrestii showed multimodal distributions. Its maximum age P. forrestii was 570 years with a DBH of 143 cm. The growth of annual ring width of P. forrestii was slow, and generally decreased with age, whereas the basal area at the breast height increased with age. Established seedlings/saplings were mainly found in unstable micro-habitats. Regeneration of P. forrestii depends on moderate natural disturbances. Finally, we provide recommendations for P. forrestii conservation.

Evidence for the persistence of wild Ginkgo biloba (Ginkgoaceae) populations in the Dalou Mountains, southwestern China.

PREMISE OF THE STUDY: The possible persistence of wild Ginkgo biloba populations in China has long been debated but never scientifically confirmed. We test our hypothesis that the extant Ginkgo populations in the Dalou Mountains (SW China) represent fragments of the original natural Ginkgo range and offer a range of pertinent perspectives on the living fossil Ginkgo's history, prehistory, ecology, and place in human culture-all important aspects of this highly valued species. METHODS: We analyzed the vegetation of the study area, determined the population age structure of Ginkgo, and compared it to existing fossil records. For supporting material, we also examined records of the lack of human presence before the mid-17th century in the area, the local people's beliefs regarding preservation of the forests and existing genetic studies. KEY RESULTS: Current species composition of Ginkgo forests in the Dalou Mountains agrees closely with floristic assemblages from fossil records bearing G. biloba. Current populations are found in habitats similar to those of fossil Ginkgo, which, as today, favored rock crevices. Female to male ratios are 3:2. Estimated ages for many of the trees show that Ginkgo was present in this area prior to human settlement and indigenous peoples of this area are unlikely to have planted Ginkgo because of traditional beliefs. Our results agree with existing genetic studies that show that these mountains were glacial refugia for G. biloba. CONCLUSIONS: The corroborative evidence confirms the finding that these populations represent fragments of the original natural Ginkgo in the valley and lower mountain slopes of the Dalou Mountains.

Climate Stability Index maps, a global high resolution cartography of climate stability from Pliocene to 2100.

Climate changes are top biodiversity shapers, both during the past and future. Mapping the most climatic stable and unstable zones on Earth could improve our understanding of biodiversity distribution and evolution. Here, we present a set of maps based on a global scale, high resolution (ca. 5 km) new Climate Stability Index (CSI). The CSI considers bioclimatic variables for two different time ranges: (1) from Pliocene (3.3 Ma) to the present (CSI-past map set), using 12 time periods of PaleoClim representing warm and cold cycles; and (2) from present to the year 2100 (CSI-future), using nine general circulation models of climate change of four periods available from WorldClim. We calculated standard deviation of the variables and selected an uncorrelated set for summing, normalizing and obtaining the CSI maps. Our approach is useful for fields such as biogeography, earth sciences, agriculture, or sociology. However, CSI is an index that can be re-calculated according to particular criteria and objectives (e.g. temperature variables); maps are, therefore, customizable to every user.

Regeneration pattern of primary forest species across forest-field gradients in the subtropical mountains of Southwestern China.

Evergreen broad-leaved forest is now gradually degraded and fragmented, and there is an increase in the amount of habitat edges as a result of long-term human activity. However, the role of edges in the regeneration of primary forest species is poorly understood. After 20 years of the edge creation, we analyzed primary forest species distribution and abundance, and changes in floristic composition, vegetation structure across forest-field gradients in Ailao Mountain, SW China. Our results revealed that there was a higher abundance and richness of primary species, late secondary species and thorny lianas at the distances 0-50 m than at the distances more than 50 m from the edge into the forest exterior. At the distances >50 m, no individuals of dominant canopy trees Lithocarpus xylocarpus, Castanopsis wattii, and L. jingdongensis were found, whereas the abundance of early pioneer shrub species and herbaceous cover was significantly greater. The richness of primary species showed a decrease with increasing distances from the forest edge to the exterior, particularly of medium-seeded primary species showing a drastic decrease. Moreover, no large-seeded primary species occurred at the distances >60 m. This study indicates that the forest edge as a buffer zone may be in favor of primary species regeneration. A dense shrub and herb layer, and seed dispersal may be the major factors limiting the forest regeneration farther from the forest edge. Therefore, to facilitate forest recovery processes, management should give priority to the protection of buffer zones of this forest edge.

Ecology of subtropical evergreen broad-leaved forests of Yunnan, southwestern China as compared to those of southwestern Japan.

Subtropical evergreen broad-leaved forests of Yunnan, China, including mid-montane moist, monsoon, and semi-humid categories, were studied in terms of ecological attributes of pertinent species, diversity, structure, dynamics and succession, as compared with the subtropical/warm-temperate evergreen broad-leaved rain forest of southwestern (SW) Japan. The genera and species of the forests of Yunnan and SW Japan are East Asian elements and indicate a very close relationship between the respective floras, though different assemblages of species occur in the two regions. Diversity indices and numerical data on taxon richness at family, genus and species level were similar in both regions. Four types of gap-regeneration behaviors among the major tree species were recognized in the two areas. In both, as a result of long-term human activity, the plant communities ranged from pioneer deciduous broad-leaved and/or pioneer coniferous stands to late-successional evergreen broad-leaved stands. Succession in the two regions followed parallel paths, beginning with pioneer Alnus in Yunnan and Alnus, Mallotus and Styrax in Japan, accompanied by coniferous Pinus in the two areas, and with late-successional evergreen broad-leaved Cyclobalanopsis and Castanopsis as their final stage.

Forest characteristics and population structure of Glyptostrobus pensilis, a globally endangered relict species of southeastern China.

The Chinese water pine Glyptostrobus pensilis is the sole surviving species of the genus Glyptostrobus. It is endemic to southern China, central Vietnam, and eastern Laos, and today it is nearly extinct in the wild. Forest community characteristics and population structure of G. pensilis in China have remained unknown up to now. We investigated six swamp forest stands and analyzed their forest community characteristics (i.e. vertical stratification, species composition, and diversity) and population structure, including the frequency distribution of DBH (diameter at breast height) and age-classes as found in Fujian Province, southeastern China. The vertical stratifications of all the forest stands were rather simple. The remaining wild specimens ranged from roughly 15 to some 357 years for an average of ca. 85 years, with only a few individuals less than 20 years old. Compared with the stands and populations of G. pensilis in Vietnam, the taxonomic compositions of the stands in the two regions were different, except for the dominant species-G. pensilis. The Shannon-Wiener index showed the overstory of each stand had much lower diversity (0.26 on average) in Fujian Province than that (1.97 on average) in Vietnam, whereas the diversity indices were about the same (around 2.41) for the understories in the two regions. Furthermore, we discovered 18 G. pensilis seedlings at the study sites in Fujian Province. This discovery demonstrates that G. pensilis regeneration is extremely poor and its populations are declining, although these populations are relatively healthier than those in Vietnam.

Author Correction: Identifying long-term stable refugia for relict plant species in East Asia.

In the original version of this Article, an incorrect sample size was provided for the number of relict species (443 instead of 442) and the number of relict forests (423 instead of 422). These errors have been corrected in both the PDF and HTML versions of the Article.

Identifying long-term stable refugia for relict plant species in East Asia.

Today East Asia harbors many "relict" plant species whose ranges were much larger during the Paleogene-Neogene and earlier. The ecological and climatic conditions suitable for these relict species have not been identified. Here, we map the abundance and distribution patterns of relict species, showing high abundance in the humid subtropical/warm-temperate forest regions. We further use Ecological Niche Modeling to show that these patterns align with maps of climate refugia, and we predict species' chances of persistence given the future climatic changes expected for East Asia. By 2070, potentially suitable areas with high richness of relict species will decrease, although the areas as a whole will probably expand. We identify areas in southwestern China and northern Vietnam as long-term climatically stable refugia likely to preserve ancient lineages, highlighting areas that could be prioritized for conservation of such species.

Potential effects of climate change on geographic distribution of the Tertiary relict tree species Davidia involucrata in China.

This study, using species distribution modeling (involving a new approach that allows for uncertainty), predicts the distribution of climatically suitable areas prevailing during the mid-Holocene, the Last Glacial Maximum (LGM), and at present, and estimates the potential formation of new habitats in 2070 of the endangered and rare Tertiary relict tree Davidia involucrata Baill. The results regarding the mid-Holocene and the LGM demonstrate that south-central and southwestern China have been long-term stable refugia, and that the current distribution is limited to the prehistoric refugia. Given future distribution under six possible climate scenarios, only some parts of the current range of D. involucrata in the mid-high mountains of south-central and southwestern China would be maintained, while some shift west into higher mountains would occur. Our results show that the predicted suitable area offering high probability (0.5‒1) accounts for an average of only 29.2% among the models predicted for the future (2070), making D. involucrata highly vulnerable. We assess and propose priority protected areas in light of climate change. The information provided will also be relevant in planning conservation of other paleoendemic species having ecological traits and distribution ranges comparable to those of D. involucrata.

Community Structure and Survival of Tertiary Relict Thuja sutchuenensis (Cupressaceae) in the Subtropical Daba Mountains, Southwestern China.

A rare coniferous Tertiary relict tree species, Thuja sutchuenensis Franch, has survived in the Daba Mountains of southwestern China. It was almost eliminated by logging during the past century. We measured size and age structures and interpreted regeneration dynamics of stands of the species in a variety of topographic contexts and community associations. Forest communities containing T. sutchuenensis were of three types: (1) the Thuja community dominated by T. sutchuenensis, growing on cliffs; (2) the Thuja-Quercus-Cyclobalanopsis community dominated by T. sutchuenensis, Quercus engleriana and Cyclobalanopsis oxyodon, along with Fagus engleriana and Carpinus fargesiana, on steep slopes; (3) the Thuja-Tsuga-Quercus community dominated by T. sutchuenensis, Tsuga chinensis, and Quercus spinosa, on crest ridges. The established seedlings/saplings were found in limestone crevices, on scarred cliff-faces, cliff-edges, fallen logs, canopy gaps and forest margins. The radial growth rate was 0.5-1.1 mm per year. Its growth forms were distorted. It had strong sprouting ability after disturbances. The T. sutchuenensis population thrives on cliffs where there is little competition from other species because of harsh conditions and rockslide disturbances. It is shade-intolerant but stress-tolerant. Its regeneration has depended on natural disturbances.