Morphometric analysis of the genusHemerocallis L. (Liliaceae) in Korea

To better understand the patterns of variability and distributions ofHemerocallis in Korea, 53 locations were visited and measurements of 19 morphological and phenological characters were taken on plants directly from their natural habitats. For morphometric analysis, 10 plants from each of 34 populations and five herbarium specimens ofH. middendorffii were used and the data from 12 quantitative characters was analyzed using univariate analysis. Except the littoral populations of Cheju, Hong, Taehuksan, and Sohuksan Islands (H. hongdoensis M. Chung & S. Kang), three peninsular KoreanHemerocallis species can be recognized mainly in South Korea:H. hakuunensis Nakai (=H. micrantha Nakai, growing on southern, central, and northwestern Korea);H. thunbergii Baker (=H. coreana Nakai, found on southeastern and central Korea); andH. middendorffii Tr. et Mey. (central and northeastern Korea). Morphological and phenological features contributing to recognition of the three groups were; color of perianth, shape of roots, shape of inflorescence, flowering time, odor, length of inflorescence, width of the lowest bracts, length of perianth tube enclosing a ovary, width of the inner perianth lobes. Natural hybridization seems to be rare in KoreanHemerocallis. It appears that the KoreanHemerocallis species are relatively well characterized by their distribution patterns, phenology, and habitats compared with the JapaneseHemerocallis species.     

Molecular evidence for the taxonomic identity of Korean Adonis (Ranunculaceae)

Although Korean Adonis has been traditionally recognized as Adonis amurensis Regel and Radde with various infraspecific taxa described, its taxonomic identity is still in dispute. We investigated the genetic variation in 60 individuals from 12 populations in Korea to elucidate the taxonomic identity of the Korean Adonis complex. Random amplified polymorphic DNA (RAPD) analysis revealed that Korean Adonis comprises three species: A. amurensis, A. pseudoamurensis, and A. multiflora. Adonis amurensis is distributed in northern central inland regions of Korea and A. pseudoamurensis is found in southern parts of the Korean peninsula. Adonis multiflora grows only on Cheju Island, which is the southernmost part of Korea. Phylogenetic analysis of nuclear ribosomal internal transcribed spacer (ITS) sequences partially supported the presence of three Adonis taxa in Korea as detected by RAPD analysis. The Adonis population on Jangbong Island located in the West Sea, which was referred to as A. pseudoamurensison the basis of morphological examination, was separated from the other populations of A. pseudoamurensis. Otherwise, the molecular evidence is well congruent with the recent morphological study that proposes that Korean Adonis consists of these three species.     

Relationships between floral morphology and pollinator fauna in insular and main island populations of Hosta longipes (Liliaceae)

We compared the floral morphologies and pollinator fauna and morphologies of Hosta longipes var. longipes on the main Japanese island of Honshu and var. latifolia on the Izu Islands to examine the differentiation processes of H. longipes in the island system. The corolla length was shorter on the southern Izu Islands than on the main island and northern Izu Islands. However, the size of other flower parts of H. longipes did not simply decrease across main island Honshu and the Izu Islands, unlike other Izu endemic plant species studied previously. Instead, the floral morphology showed a complicated variation pattern. Pollinator fauna of H. longipes on the Izu Islands varied more widely than those on the main island. The diverse pollinator fauna may have influenced the morphological differentiation of H. longipes on these islands.     

The distribution of diploids and polyploids of the Scilla scilloides complex in Korea

A total of 3 384 individuals of Scilla scilloides Druce were sampled from 40 natural populations at 35 different localities over the Korean Peninsula and analysed cytogenetically. The following cytogenetic types were found: AA (19.6%), AABB (72.4%), ABBB (4.3%), AABBB (3.0%) and aneuploids (0.8%). Of these, AA was found in 9 populations along the west side of the Korean Peninsula facing the Yellow Sea and on Yeongdo and Jeju Islands. Type AABB, the most common type in the complex, was found in 33 natural populations on the Korean Peninsula and Jeju Island. The distributions of ABBB and AABBB are restricted to the southern part of the Korean Peninsula and Jeju Island; they were found in 8 natural populations. In southern Korea the above four types showed overlapping distribution and two or three were often found growing together.Genome A, occurring in the three allopolyploids in the Korean Peninsula, was the same as that in allopolyploids in Japan. However, genome A occurring as AA diploids was karyotypically different from that in the allopolyploids. On the basis of distribution patterns and cytological features, the origin of the allopolyploids is discussed.     

Anagenetic evolution in island plants

Aim  Plants in islands have often evolved through adaptive radiation, providing the classical model of evolution of closely related species each with strikingly different morphological and ecological features and with low levels of genetic divergence. We emphasize the importance of an alternative (anagenetic) model of evolution, whereby a single island endemic evolves from a progenitor and slowly builds up genetic variation through time.
Location  Continental and oceanic islands.
Methods  We surveyed 2640 endemic angiosperm species in 13 island systems of the world, both oceanic and continental, for anagenetic and cladogenetic patterns of speciation. Genetic data were evaluated from a progenitor and derivative species pair in Ullung Island, Korea, and Japan.
Results  We show that the anagenetic model of evolution is much more important in oceanic islands than previously believed, accounting for levels of endemic specific diversity from 7% in the Hawaiian Islands to 88% in Ullung Island, Korea, with a mean for all islands of 25%. Examination of an anagenetically derived endemic species in Ullung Island reveals genetic (amplified fragment length polymorphism) variation equal or nearly equal to that of its continental progenitor.
Main conclusions  We hypothesize that, during anagenetic speciation, initial founder populations proliferate, and then accumulate genetic variation slowly through time by mutation and recombination in a relatively uniform environment, with drift and/or selection yielding genetic and morphological divergence sufficient for the recognition of new species. Low-elevation islands with low habitat heterogeneity are highly correlated with high levels of anagenetic evolution, allowing prediction of levels of the two models of evolution from these data alone. Both anagenetic and adaptive radiation models of speciation are needed to explain the observed levels of specific and genetic diversity in oceanic islands.     

Molecular evidence for the taxonomic identity of Korean Adonis (Ranunculaceae)

 Although Korean Adonis has been traditionally recognized as Adonis amurensis Regel and Radde with various infraspecific taxa described, its taxonomic identity is still in dispute. We investigated the genetic variation in 60 individuals from 12 populations in Korea to elucidate the taxonomic identity of the Korean Adonis complex. Random amplified polymorphic DNA (RAPD) analysis revealed that Korean Adonis comprises three species: A. amurensis, A. pseudoamurensis, and A. multiflora. Adonis amurensis is distributed in northern central inland regions of Korea and A. pseudoamurensis is found in southern parts of the Korean peninsula. Adonis multiflora grows only on Cheju Island, which is the southernmost part of Korea. Phylogenetic analysis of nuclear ribosomal internal transcribed spacer (ITS) sequences partially supported the presence of three Adonis taxa in Korea as detected by RAPD analysis. The Adonis population on Jangbong Island located in the West Sea, which was referred to as A. pseudoamurensis on the basis of morphological examination, was separated from the other populations of A. pseudoamurensis. Otherwise, the molecular evidence is well congruent with the recent morphological study that proposes that Korean Adonis consists of these three species. Received: November 6, 2001 / Accepted: February 26, 2002     

Morphometric analysis and taxonomic revision of the Vriesea paraibica complex (Bromeliaceae)

The species related to Vriesea paraibica (Bromeliaceae, Tillandsioideae) have controversial taxonomic limits. For several decades, this group has been identified in herbarium collections as V. × morreniana, an artificial hybrid that does not grow in natural habitats. The aim of this study was to assess the morphological variation in the V. paraibica complex through morphometric analyses of natural populations. Two sets of analyses were performed: the first involved six natural populations (G1) and the second was carried out on taxa that emerged from the first analysis, but using material from herbarium collections (G2). Univariate ANOVA was used, as well as discriminant analysis of 16 morphometric variables in G1 and 18 in G2. The results of the analyses of the two groups were similar and led to the selection of diagnostic traits of four species. Lengths of the lower and median floral bracts were significant for the separation of red and yellow floral bracts. Vriesea paraibica and V. interrogatoria have red bracts; these two species are differentiated by the widths of the lower and median portions of the inflorescence and by scape length. These structures are larger in the former and smaller in the latter. Of the species with yellow floral bracts, V. eltoniana is distinguished by longer leaf blades and scapes and V. flava is characterized by its shorter sepal lengths. © 2009 The Linnean Society of London, Botanical Journal of the Linnean Society, 2009, 159 , 163–181.     

Genetic homogeneity between Korean and Japanese populations of the broad-leaved evergreen tree Machilus thunbergii (Lauraceae): A massive post-glacial immigration through the Korea Strait or something else?

The warm-temperate vegetation of Korea, currently limited to southern coastal areas, shifted during the Last Glacial Maximum (LGM) towards glacial refugia, putatively located in southern Japan. We hypothesized two scenarios of post-glacial re-colonization of warm-temperate species in relation to current levels of genetic diversity within their populations. If extant Korean populations originated from a single source (a single glacial refugium), we expect significantly lower levels of genetic diversity relative to those from Japan due to founder effects. Alternatively, if they were derived from multiple source populations, levels of genetic diversity within Korean populations will not be significantly reduced compared to those of Japanese ones. To test which of these scenarios is more likely, we investigated the patterns of genetic diversity in 14 populations (seven from southern Korea and seven from southern Japan) of the broad-leaved evergreen tree Machilus thunbergii, employing 11 allozyme loci. High levels of genetic variation in M. thunbergii were found both in southern Korea and southern Japan, with a considerable genetic homogeneity not only between the two regions but also between populations within the two regions. These results suggest a pattern of re-colonization after the LGM fitting the second scenario (immigration from multiple refugia), probably through multiple waves and/or with large founder populations.     

Genetic variation and population structure in Korean endemic species: III.Hosta minor (Liliaceae)

Hosta minor, an insect-pollinated and rhizomatous herbaceous perennial, occurs in eastern and southern Korean Peninsula. AlthoughH. minor is a Korean endemic species and most populations of the species are scattered, the species maintains considerably high levels of genetic variation within the species, with a moderate level of variation (14%) found among populations. Nineteen of the 29 putative loci resolved (66%) were polymorphic within the species, the mean number of alleles per locus was 2.10 across all populations and averaged 1.68 within populations. In addition, genetic diversity was a considerably higher (for species and population level, mean estimates of genetic diversity were 0.275 and 0.230, respectively) than the average for other long-lived herbaceous perennials. Indirect estimate of the number of migrants per generation (Nm=1.03, calculated from mean G_ST) was moderate. Factors contributing to the high levels of genetic diversity found within populations ofH. minor include population maintenance via sexual and asexual modes of reproduction, primarily outcrossing breeding system, long generation time, probable ancient polyploid origin of the species, and moderate levels of seed dispersal by wind. Human disturbance in South Korea such as road and apartment constructions appears to be the major threat to this genetically diverse species.     

Phylogeography of the Asian lesser white-toothed shrew, <Emphasis Type="Italic">Crocidura shantungensis</Emphasis>, in East Asia: role of the Korean Peninsula as refugium for small mammals

Many peninsulas in the temperate zone played an important role as refugia of various flora and fauna, and the southern Korean Peninsula also served as a refugium for many small mammals in East Asia during the Pleistocene. The Asian lesser white-toothed shrew, Crocidura shantungensis, is a widely distributed species in East Asia, and is an appropriate model organism for exploring the role of the Korean Peninsula as a refugium of small mammals. Here, we investigated phylogenetic relationships and genetic diversity based on the entire sequence of the mitochondrial cytochrome b gene (1140 bp). A Bayesian tree for 98 haplotypes detected in 228 C. shantungensis specimens from East Asia revealed the presence of three major groups with at least 5 subgroups. Most haplotypes were distributed according to their geographic proximity. Pairwise F_ST’s and analysis of molecular variance (AMOVA) revealed a high degree of genetic differentiation and variance among regions as well as among populations within region, implying little gene flow among local populations. Genetic evidence from South Korean islands, Jeju-do Island of South Korea, and Taiwan leads us to reject the hypothesis of recent population expansion. We observed unique island-type genetic characteristics consistent with geographic isolation and resultant genetic drift. Phylogeographic inference, together with estimates of genetic differentiation and diversity, suggest that the southern most part the Korean Peninsula, including offshore islands, played an important role as a refugium for C. shantungensis during the Pleistocene. However, the presence of several refugia on the mainland of northeast Asia is also proposed.     

Ecomorphological patterns in the Blackcap Sylvia atricapilla: insular versus mainland populations

Capsule Blackcap Sylvia atricapilla populations from the Azores archipelago show morphological differences to continental birds which are consistent with the ‘Island Rule’.

Aims The morphology of insular vertebrates is usually the result of the evolution in their particular environment and leads to predictable morphological patterns, according to the Island Rule. We test the predictions of the Island Rule, using the Blackcap of the Azores archipelago as our model.

Methods We compared morphological variation (body size and wing shape) of populations from the nine islands of the Azores to continental birds, using multivariate indexes. Also, we looked at the relationship between these patterns and possible insular ecological drivers of morphological divergence.

Results Our findings are concordant with Island Rule predictions, as in general birds from the Azores are larger than continental populations, especially birds from the most distant islands. Wing shape also differs significantly, as Azorean Blackcaps tend to have rounder wings than continental birds with a migratory-like phenotype.

Conclusion Overall, we conclude that the observed morphological patterns in Blackcap in the Azores conform in general to the Island Rule predictions.     

Development of microsatellite markers to genetically differentiate populations of Octopus minor from Korea and China

Of the more than 300 octopus species, Octopus minor is one of the most popular and economically important species in Eastern Asia, including Korea, along with O. vulgaris, O. ocellatus, and O. aegina. We developed 19 microsatellite markers from Octopus minor and eight polymorphic markers were developed to analyze the genetic diversity and relationships among four octopus populations from Korea and three from China. The number of alleles per locus varied from 10 to 49, and allelic richness per locus ranged from 2 to 16.4 across all populations. The average allele number among the populations was 11.1, with a minimum of 8.3 and a maximum of 13.6. The mean allelic richness was 8.7 in all populations. The Hardy-Weinberg equilibrium (HWE) test revealed significant deviation in 19 of the 56 single-locus sites, and null alleles were presumed in five of eight loci. The pairwise F ( ST ) values between populations from Korea and China differed significantly in all pairwise comparisons. The genetic distances between the China and Korea samples ranged from 0.161 to 0.454. The genetic distances among the populations from Korea ranged from 0.033 to 0.090, with an average of 0.062; those among populations from China ranged from 0.191 to 0.316, with an average of 0.254. The populations from Korea and China formed clearly separated into clusters via an unweighted pair group method with arithmetic mean dendrogram. Furthermore, a population from muddy flats on the western coast of the Korean Peninsula and one from a rocky area on Jeju Island formed clearly separated subclusters. An assignment test based on the allele distribution discriminated between the Korean and Chinese origins with 96.9 % accuracy.     

The spatial analysis of biological interactions: morphological variation responding to the co‐occurrence of competitors and resources

By sharing geographic space, species are forced to interact with one another and the contribution of this process to evolutionary and ecological patterns of individual species is not fully understood. At the same time, species turnover makes that species composition varies from one area to another, so the analysis of biological interaction cannot be uncoupled from the spatial context. This is particularly important for clades that show high degree of specialization such as hummingbirds, where any variation in biotic pressures might lead to changes in morphology. Here, we describe the influence of biological interactions on the morphology of Hylocharis leucotis by simultaneously considering potential competition and diet resources. We characterized the extent of local potential competition and local available floral resources by correlating two measurements of hummingbird diversity, floral resources and the size of morphological space of H. leucotis along its geographic distribution. We found that H. leucotis shows an important morphological variability across its range and two groups can be recognized. Surprisingly, morphological variation is not always linked to local hummingbird richness or the phylogenetic similarity of. Only in the southern part of its distribution, H. leucotis is morphologically more variable in those communities where it coexist with closely related hummingbird species. We also found that morphological variation in H. leucotis is independent from the availability of floral resources. Our results suggest that abiotic factors might be responsible for morphological differences across populations in Hylocharis leucotis being biological interactions of minor importance.     

Habitat fragmentation and body‐shape variation of African hind Cephalopholis taeniops (Valenciennes) in an archipelago system (Cape Verde,eastern Atlantic Ocean)

Morphological variations of the serranid fish Cephalopholis taeniops were studied in relation to habitat fragmentation in the Cape Verde Archipelago. While a significant allometric effect existed (11·5% of total body‐shape variation), differences in morphology associated with sex and ontogeny were not significant. MANOVA followed by CVA showed that each island presented a particular allometric pattern. Average body shape for all islands was well discriminated with CVA models. Pair‐wise comparisons of the ontogeny of morphological change between islands revealed that northern islands (Santo Antão, São Vicente and Santa Luzia) along with Boavista Island showed a similar direction in shape ontogeny, while all other paired comparisons indicated different ontogenetic patterns. When comparing directions of inter‐population shape changes, individuals from Fogo Island, the southernmost locality, departed far from the orthogonal relation, suggesting that they were undergoing independent body‐shape trajectories. Physical isolation by geographic distance and depth was positively correlated with morphological divergence among populations from different islands. This finding supports the hypothesis that habitat fragmentation in the Cape Verde Archipelago can be interpreted in terms of marine population structure.     

Phenotypic and genetic divergence among island populations of sika deer (<Emphasis Type="Italic">Cervus nippon</Emphasis>) in southern Japan: a test of the local adaptation hypothesis

Transplant and common garden experiments have been used in studies on local adaptation, but are difficult to be conducted for large animals with long life span. A previous study on the southern Japanese islands demonstrated that relative limb lengths of sika deer (Cervus nippon) were short on islands with steep slopes. We hypothesized that this morphological variation was evidence for local adaptation, and tested this hypothesis by comparing phenotypic divergence with neutral genetic divergence among eight populations of the sika deer in the southern Japanese islands. Divergence patterns differed between the phenotypic and neutral genetic features. Genetic similarity was high among individuals on Kyushu (OI, KGS, and KGK). Individuals on Tanegashima (TN) and Yakushima (YK) also constituted a group, whereas individuals on Tsushima (TS), Wakamatsujima (WM), and Kuchinoerabujima (KE) formed a genetically distinct group. Phenotypic data indicated that individuals from TS, OI, KGS, and KGK exhibited similarity, whereas individuals on YK formed an isolated group that was separated from the other populations. The degree of phenotypic divergence was larger than that of neutral genetic divergence between TN and YK. These results suggest that divergent selection worked between two of the eight island populations (TN and YK). The morphological trait of captive-bred individuals from TN and YK, which had never experienced their original environments, retained their original morphological features. By combining the results of multiple analyses, we found that the difference in relative limb length between the two populations was consistent with local adaptation hypothesis, although conclusive results were not obtained for the other populations.     

Upwelling and eddies affect connectivity among local populations of the goldeye rockfish,Sebastes thompsoni (Pisces,Scorpaenoidei)

The goldeye rockfish, Sebastes thompsoni, commercial rockfish catch in the Northwest Pacific Ocean, may influence its population structure. To clarify the population genetic structure of Korean S. thompsoni and its degree of hybridization with the most close species, Sebastes joyneri, we analyzed a mitochondrial (mt) DNA control region and eleven polymorphic microsatellite (ms) loci. S. joyneri individuals were clearly distinguished from S. thompsoni by the mtDNA control region and ms loci results, with single interspecific hybridization between two species suggesting no impact on genetic structure of S. thompsoni. Analysis of mtDNA revealed no population structure within S. thompsoni, suggesting the survival of a single population in southern refugia during the glacial period. The ms loci results, in contrast, showed two genetically distinct clusters within S. thompsoni: One was predominant throughout Korean coasts (from the Yellow Sea, via the Korea Strait to the East Sea); the other was predominant at Dokdo Island in the East Sea; and both occurred in similar ratios at Wangdolcho Reef in the East Sea. A possible factor that restricts gene flow between Korean coastal and offshore populations in the East Sea may be related to the complex oceanic current patterns such as eddies and upwelling, which represent impermeable barriers to population connectivity for this species. Our findings highlight that these two populations might be representative of two separate stock within Korean waters and maintain their geographically related genetic structure.     

Genetic diversity and historical migration patterns of an endemic evergreen oak,Quercus acuta,across Korea and Japan,inferred from nuclear microsatellites

Forests dominated by broad-leaved evergreen trees cover a narrow zone of the southern coast of Korea as well as warm-temperate and subtropical regions of Japan. However, little is known about their establishment history, especially in Korea. Endemic Quercus acuta grows in Korea and Japan. We evaluated its genetic diversity, population structure, and degree of lineage admixture in terms of its distribution history. Analyses utilized seven nuclear microsatellites that were genotyped from 330 individuals of 16 populations. Genetic diversity was highest for the Japanese Kyushu populations. The allelic diversity of populations was similar between Jeju Island and Kyushu, implying that they are putative refugia. Although the mean F _ST value was very low, patterns for isolation-by-distance were strong and significant. Results from Bayesian clustering and Monmonier’s algorithm indicated that populations are roughly partitioned from west (Korea and Kyushu) to east (Japanese Honshu). Therefore, the two divisions within this species are demographically independent and may have arisen due to past fragmentation. In considering the mixed genetic structure between Korea and Kyushu populations, and the higher levels of genetic diversity in mainland Korea relative to Honshu, we hypothesized that this species has been able to survive within Jeju Island and Kyushu. Consequently, mainland Korea populations might have been shaped by range expansion that kept them well mixed from a continuous and large genetic resource.     

First record of the termite family Kalotermitidae (Blattodea: Termitoidae) in Korea

The drywood termite Kalotermitidae Froggatt, represented by Glyptotermes nakajimai Morimoto, is identified for the first time in Korea. They were collected in Yeoseo-do Island, the southern part of the Korean peninsula. A key to the termite species in Korea, habitus photographs of all castes, ecological information, and illustrations of diagnostic characters are provided.     

Evolution of Mating Systems in Island Populations of Campanula microdonta: Pollinator Availability Hypothesis

Abstract Island populations of Campanula microdonta and mainland Honshu populations of C. punctata have several different mating systems: self-incompatible (SI) and therefore obligately outcrossing in mainland Honshu and Oshima Island; self-compatible (SC) and largely outcrossing in the northern islands of Toshima and Niijima; and SC and predominantly inbreeding southern ones of Miyake and Hachijo. Several features possibly associated with the mating systems are described. A hypothesis is proposed that the mean and variance of pollinator availability together with inbreeding depression could explain the observed patterns of mating system as the results of evolution.