Allozyme diversity and the evolution ofSymplocos (Symplocaceae) on the Bonin (Ogasawara) Islands

To study the origin and speciation of plants in oceanic islands, electrophoretic analyses have been done on three endemic species ofSymplocos in the Bonin Islands as well as on three other species;S. kuroki, S. nakaharae andS. tanakae which are considered to be closely related to the Bonin endemics. There occur three species:S. kawakamii, S. pergracilis andS. boninensis in Bonin. The genusSymplocos is one which is considered to be diversified in the Bonin Islands. Seven enzyme systems presumed to be encoded by 18 loci were examined. The genetic diversity was low in the island species, as reported in some oceanic island plants of Hawaii and the Bonin Islands. The three endemics share high genetic identities and they clustered together in the tree drawn by the UPGMA method, suggesting that they are a monophyletic group, that is, they result from a single introduction.     

Allozyme diversity ofPittosporum (Pittosporaceae) on the Bonin (Ogasawara) Islands

The genusPittosporum includes about 160 species. Four species ofPittosporum occur in the Bonin Islands, and all of these are endemic to the islands. Electrophoretic studies of the four endemic species,P. tobira, from the Japanese mainland, andP. lutchuense var.denudatum from the Ryukyu Islands, were used to determine the origin and speciation pattern of the endemic species. 259 individuals were sampled from ten populations. Twenty loci in nine enzyme systems were resolved and used to calculate the gene frequencies for each population. A low genetic diversity was observed in three of the Bonin Island species, as is reported for other oceanic island plants. The exception,P. boninense, has the largest population size and widest distribution. A dendrogram generated by the UPGMA method shows two clusters. One consists of only the Bonin endemics, suggesting a monophyletic origin for these species.     

Phylogenetic relationships ofCrepidiastrum (Asteraceae) of the Bonin (Ogasawara) Islands based on chloroplast DNA restriction site variation

RFLPs of cpDNA were investigated for seven species ofCrepidiastrum, of which three are endemic to the Bonin Islands. As an outgroup for rooting the tree,Paraixeris denticulata was also examined. Approximately 350 restriction sites were surveyed using 16 restriction enzymes. A total of 26 restriction site mutations were detected, and seven of them were phylogenetically informative. Wagner parsimony analysis resulted in four most parsimonious trees. In the tree obtained, the Bonin endemics are monophyletic with four mutations and a bootstrap value of 0.98 for the branch. This result agrees with that obtained from a previous electrophoretic analysis (Ito and Ono 1990), and supports the hypothesis that the three Bonin endemics have been evolved from a common ancestor. The Bonin endemics cluster together withC. lanceolatum, suggesting that the endemics andC. lanceolatum share a common ancestor, although the bootstrap probability is not very high (0.78) and thus other possibilities cannot be rejected.     

Origin and evolution of endemic plants of the Bonin (Ogasawara) Islands

The Bonin Islands are typical oceanic islands, located at the western part of the North Pacific Ocean and approximately 1,000 km south of mainland Japan. This archipelago consists of about 20 small islands. Although floristic diversity is low due to the small area and limited environmental diversity, the Bonin Islands harbor unique endemic flora as in other well-known oceanic islands. This paper presents a brief summary of the results obtained from recent studies on the endemic flora of the Bonin Islands. The results are reviewed in relation to the four stages of the evolution of endemic flora in the oceanic islands; migration, establishment, enlargement and diversification. The ancestors of the flora originated mostly from tropical and subtropical Southeast Asia or mainland Japan by rare events of long distance dispersal. The proportion of bird-dispersed species is relatively high as for other oceanic islands. Genetic data sets obtained from allozyme variation in some endemic species suggest that migration occurred several million years ago and genetic diversity is correlated with current population size. At the time of establishment, self-compatible plants are expected to have an advantage. However, the percentage of dioecious plants is relatively high. This is partly due to evolutionary changes from hermaphroditic ancestors to dioecy which occurred in two genera in the Bonin Islands. In addition, there are some examples of evolutionary changes from herbaceous ancestors to woody endemics. Adaptive radiation is found in some genera, although the number of congeneric endemic species is less than five. Studies of allozyme variation inPittosporum, Symplocos andCrepidiastrum showed that genetic identity is generally very high between congeneric species in spite of their distinct morphologies. This result suggests that divergence of these species occurred rather recently and distinct morphological differences are based on a limited number of genetic changes.     

Dioecism of the genusCallicarpa (Verbenaceae) in the Bonin (Ogasawara) Islands

The morphology and function of the flowers of three species of the genusCallicarpa, C. glabra, C. nisnimurae andC. subpubescens, endemic to the Bonin Islands, revealed that all three species were dioecious. Male plants had short-styled flowers with sterile ovaries, while female plants had long-styled flowers with non-germinating inaperturate pollen grains. Such dioecism has never been reported from the family Verbenaceae. This curious sexual system with non-germinating pollen grains may have been maintained as a reward to insect pollinators.     

Evolution ofPeperomia (Piperaceae) in the Juan Fernandez Islands,Chile

Four species ofPeperomia (Piperaceae) occur in the Juan Fernandez Islands, Chile:P. berteroana, P. margaritifera, P. skottsbergii, andP. fernandeziana. The last species is found also in continental Chile, whereas the other three are endemic to the archipelago.Peperomia margaritifera is found only on the older island of Masatierra, whereasP. skottsbergii is confined to the younger island of Masafuera, andP. berteroana occurs on both islands. Phenetic analyses of mainland taxa suggest thatP. fernandeziana belongs to subg.Sphaerocarpidium whereas the endemic taxa form their own subg.Tildenidium connecting to subg.Tildenia. Cladistic analyses indicate thatP. margaritifera is the most primitive species in the archipelago and thatP. berteroana is the most derived, especially patristically. Chromosomally, the four species are all n = 22, which may be tetraploid on a base of x = 11. Sulfated flavones occur only inP. berteroana andP. skottsbergii, which are otherwise unknown for the family. Dispersal of propagules to the islands from the continent and between islands is believed to have been accomplished by birds.     

Phylogenetic relationships inCrossostylis (Rhizophoraceae) inferred from restriction site variation of chloroplast DNA

Phylogenetic relationships among the nine species ofCrossostylis (Rhizophoraceae) were elucidated using cladistic analysis of restriction site variations of chloroplast DNA. As a result, this genus was found to comprise two pronounced monophyletic groups as follows:C. biflora, C. grandiflora, C. multiflora andC. sebertii; andC. cominsii, C. pachyantha, C. parksii, C. richii andC. seemannii. Moreover, the monophyly ofC. biflora, C. grandiflora andC. sebertii in the former group and the monophyly ofC. pachyantha, C. parksii, C. richii andC. seemannii in the latter group were also suggested. The molecular tree corresponded well with that inferred from morphological data and no discrepancy was recognized. Many of the floral morphological characters reflected lineage, but all seed coat characters were homoplasious. Evolutionary trends in some morphological characters were optimized on the cpDNA tree obtained. Species from New Caledonia and Polynesia were monophyletic, as were those from the Solomon Islands, Vanuatu and the Fiji Islands. All species endemic to the Fiji Islands made a cluster, and this suggests that speciation occurred from a single ancestral species on the Islands.     

Dioecy in the endemic genusDendrocacalia (Compositae) on the Bonin (Ogasawara) Islands

Dendrocacalia crepidifolia Nakai (Compositae, Senecioneae), the only species of this arboreal genus endemic to Haha Island in the Bonin Islands, was found to be dioecious. Male flowers differ from female ones in having a stunted style (style in female exserted from corolla and deeply bifurcating) and anthers filled with fertile pollen (anthers in the female lacking pollen). The size of the corolla and number of florets per head were similar between male and female flowers. The crown area of this arboreal species was also similar in male and female plants. The sex ratio was 0.55 male, not significantly different from 0.5. Both sexes produced nectar of similar sugar concentration (ca. 50%). The flowers are pollinated by feral honeybees (Apis mellifera), but they are thought to have been pollinated by small, lesshairy, endemic solitary bees before honeybees were introduced and subsequently became the dominant bee species on the island. The evolution of dioecy ofDendrocacalia on the island is thought to stem from the deleterious effects of inbreeding that are inherent in plants with geitonogamy. The increased geitonogamy on the island has resulted from increased woodiness (i.e., increased number of flowers per plant) and the original dependence on endemic bee pollinators, which are now endangered.     

Song dialects on the Atlantic islands: goldcrests of the Azores ( Regulus regulus azoricus, R. r. sanctae-mariae, R. r. inermis)

Territorial songs in island populations of songbirds are often highly divergent from those of neighbouring continental relatives. This is shown for the three goldcrest subspecies (Regulus regulus azoricus, R. r. sanctae-mariae and R. r. inermis) endemic on six islands of the Azorean archipelago. All investigated populations display a high intra- and inter-individual acoustic variation. On each island, up to six different song types have been found; and a single male sings up to three types. In contrast, all northwestern European populations of R. r. regulus and R. r. anglorum share only a single song type. In playback experiments, none of 18 tested dialect songs of Azorean goldcrests evoked notable territorial reaction in German and Czech goldcrest males (ssp. regulus). Two differing dialect groups of the goldcrest can be distinguished on the Azores. Populations of the eastern islands, São Miguel and Santa Maria, share common song types which are not found on the islands of the central and western groups. Dialect repertoires on the westernmost islands, Flores, Faial and Pico, are dominated by a different song type. In the geologically younger western crater of São Miguel, both western and eastern song types coexist. Acoustic similarities to a population from neighbouring Terceira suggest the western part of São Miguel as the origin for the westward expansion of R. regulus on the Azores.     

Phylogenetic analysis of Asian<Emphasis Type="Italic"> Symplocos</Emphasis> (Symplocaceae) based on nuclear and chloroplast DNA sequences

Thirty species and one variety of Symplocos (Symplocaceae), including all taxa distributed in Japan, were phylogenetically analyzed with DNA sequence data. The evolution of morphological characters is discussed on the basis of the phylogenetic relationships obtained. All species were Asian, except one, S. austromexicana, from Mexico. The nuclear ribosomal internal transcribed spacer (ITS) region and two intergenic spacer regions, between trnL and trnF, and between trnH and psbA of chloroplast DNA were used. The topologies of trees obtained from ITS and the chloroplast intergenic spacers are largely congruent. S. sonoharae, the representative of the subgenus Symplocos in Asia, was sister to all other species. The position of the American species, S. austromexicana, which also belongs to the subgenus Symplocos, was not well resolved. The phylogenetic tree based on combined sequence data largely supports the monophyletic origin of the infrageneric sections proposed earlier. However, the phylogenetic relationship between them is not well resolved, probably due to rapid diversification. The section Palura, a deciduous group, is well defined in the DNA analysis, suggesting its independent status in the genus Symplocos. In spite of their morphological divergence, the three endemic species of the Bonin Islands are monophyletic. The occurrence of curved seeds seems to be homoplastic, scattered over the phylogenetic tree without showing a particular infrageneric relationship.     

Morphological Variation of Three Endemic Species of Callicarpa (Verbenaceae) in the Bonin (Ogasawara) Islands

Abstract A total of 22 population samples of three endemic species of Callicarpa in the Bonin Islands (the Chichijima Islands and the Hahajima Islands) were examined for nine leaf characters to clarify the present status of morphological variations. Results showed that three species are distinetly recognizable and have their own patterns of variation. Callicarpa glabra and C. nishimurae were relatively less variable and adapted to limited habitats only in the Chichijima Islands. Thus these species were regarded as specialized species. C. subpubescens was more variable than the above two species and distributed widely in both of the two island groups. This species was morphologically differentiated between the two island groups. In the Hahajima Islands where two other species were not distributed, several plants of C. subpubsecens were similar to C. glabra. The populations of C. subpubescens in the Hahajima Islands were inter-populationally more variable than those in the Chichijima Islands. C. subpubescens occupied various habitats within the Bonin Islands and was regarded as a diversified species.     

Low genetic differentiation betweenAdiantum capillusveneris L. andA. ogasawarense Tagawa (Adiantaceae), an endemic pteriodophyte of the bonin (ogasawara) islands

Genetic differentiation was examined betweenAdiantum capillus-veneris L. andA. ogasawarense Tagawa. A total of 136 individuals from two populations ofA. ogasawarense and three populations ofA. capillus-veneris were sampled. Allozymes encoded by 19 putative loci for nine enzyme systems were observed. The populations ofA. ogasawarense are genetically differentiated from those ofA. capillus-veneris; conspecific populations cluster together. However, genetic identity between two species was estimated to be 0.815, and indicates that the degree of allozyme differentiation is lower than those between endemic flowering plants and their progenitors. It is possible that the low genetic differentiation observed betweenA. ogasawarense andA. capillus-veneris results from a recent origin of the endemic species,A. ogasawarense.     

Genetic variation of pantropical Terminalia catappa plants with sea‐drifted seeds in the Bonin Islands: suggestions for transplantation guidelines

The Bonin Islands are endowed with endemic species. However, these species are at risk of extinction because of the exuberance of invasive alien plants. Therefore, native plant species should be revegetated after eradicating alien plants. We investigated the genetic variation of Terminalia catappa populations in the Bonin Islands by using nuclear (n) microsatellites (simple sequence repeats [SSRs]) and chloroplast (cp) DNA. No significant differences were observed in the genetic diversity of nSSRs among 22 populations. However, recent bottlenecks were detected in three populations on the Chichijima Island group. nSSR variation and cpDNA haplotypes suggested the presence of two genetically distinct groups in the Mukojima and Chichijima Island groups and the Hahajima Island group. A similar genetic structure was observed in plants and animals in the Bonin Islands. Populations on the three islands, which were separated from other islands in each island group when the water depth was 50‐m lower than the present level, were dominated by unique nSSRs clusters, suggesting that historical changes in island connections during the Pleistocene era affected genetic substructuring. These results suggested that different factors contributed to the genetic structure of T. catappa on different geographic scales. At the whole‐island level, the genetic structure was determined by long‐distance seed dispersal by ocean currents. At the island‐group level, the genetic structure was determined by historical changes in island connections caused by changes in the sea level due to glacial–interglacial transition. These findings would help in establishing transplantation zone borders for revegetating T. catappa on the Bonin Islands.     

Systematic study of the genusDelilia (Asteraceae,Heliantheae)

A taxonomic study of the genusDelilia is presented. Two species are recognized:D. biflora, a widespread weed of tropical and subtropical regions of the New World, extending from Mexico to northern Argentina and recently introduced in the Cape Verde Islands; andD. repens, endemic to the Galapagos islands. A key to the species, illustrations, distribution maps, and complete synonymy are presented.     

Molecular evidence for adaptive radiation of Micromeria Benth. (Lamiaceae) on the Canary Islands as inferred from chloroplast and nuclear DNA sequences and ISSR fingerprint data

The Canary Islands have been a focus for phylogeographic studies on the colonization and diversification of endemic angiosperm taxa. Based on phylogeographic patterns, both inter island colonization and adaptive radiation seem to be the driving forces for speciation in most taxa. Here, we investigated the diversification of Micromeria on the Canary Islands and Madeira at the inter- and infraspecific level using inter simple sequence repeat PCR (ISSR), the trnK-Intron and the trnT-trnL-spacer of the cpDNA and a low copy nuclear gene. The genus Micromeria (Lamiaceae, Mentheae) includes 16 species and 13 subspecies in Macaronesia. Most taxa are restricted endemics, or grow in similar ecological conditions on two islands. An exception is M. varia, a widespread species inhabits the lowland scrub on each island of the archipelago and could represent an ancestral taxon from which radiation started on the different islands. Our analyses support a split between the "eastern" islands Fuerteventura, Lanzarote and Gran Canaria and the "western" islands Tenerife, La Palma and El Hierro. The colonization of Madeira started from the western Islands, probably from Tenerife as indicated by the sequence data. We identified two lineages of Micromeria on Gomera but all other islands appear to be colonized by a single lineage, supporting adaptive radiation as the major evolutionary force for the diversification of Micromeria. We also discuss the possible role of gene flow between lineages of different Micromeria species on one island after multiple colonizations.     

Electrophoretic evidence for the origins of tetraploids and hybrids ofSphenomeris chinensis sensu lato (Lindsaeaceae) in Japan

Hypotheses for the origins of tetraploids and hybrids ofSphenomeris chinensis sensu lato in Japan were tested using electrophoretic analysis of isozymes. Comparison of banding patterns supported the hypothesis that plants having abortive spores had resulted following hybridization between diploidS. chinensis var.chinensis andS. biflora and between diploid var.chinensis andS. gracilis. Results from isozymic analyses also revealed thatS. intermedia probably originated through allotetraploid speciation following hybridization between var.chinensis andS. biflora. There were two electrophoretic variants among tetraploids ofS. chinensis. One variant, restricted to the southern Ryukyu Islands, is probably an allotetraploid of hybrid origin between diploidS. chinensis andS. gracilis. It has been described as a new species,S. yaeyamensis Lin, Kato & K. Iwatsuki. The other variant, morphologically corresponding to var.chinensis, is distributed at least throughout southern Japan and also in northern Thailand. Its origin is uncertain, but one of the parents is probably diploidS. chinensis. Sphenomeris intermedia andS. yaeyamensis occur inland, whereas the parentalS. biflora andS. gracilis are halophilous and rheophllous, respectively.     

Isolation and characterization of microsatellite loci in <Emphasis Type="Italic">Melicope quadrilocularis</Emphasis> (Rutaceae), an endemic plant species of the Bonin Islands,Japan, and cross-species amplification in closely related taxa

Twelve microsatellite loci were isolated and characterized for Melicope quadrilocularis, an insular endemic tree species of the Bonin Islands. The observed number of alleles at each locus ranged from 1 to 18. The range of expected heterozygosity was 0.0000–0.9445. The inter-specific applicability of these loci was evaluated by analyzing two other endemic species and one endemic variety of Melicope that are also distributed on the Bonin Islands. All primer pairs for the 12 loci tested successfully amplified the loci in all taxa, except for primers for four loci in M. nishimurae.     

Distyly and incompatibility in Psychotria homalosperma (Rubiaceae), an endemic plant of the oceanic Bonin (Ogasawara) Islands

Distyly rarely occurs on oceanic islands. Indeed, this form of polymorphism is absent on the Hawaiian Islands and in New Zealand. However, Psychotria homalosperma (Rubiaceae), which is endemic to the oceanic Bonin Islands of Japan, was presumed to maintain a distylous breeding system and floral morphology. We investigated the floral and reproductive characteristics of this species and examined incompatibility in self- and intramorph pollination. Psychotria homalosperma is morphologically and functionally distylous, based on findings with the examined populations, but its stigma and anther heights are not exactly reciprocal between the two morphs. This is the second example of distyly among the flora from the Bonin Islands, the other being the endemic P. boninensis. Both of these distylous species contrast with the Hawaiian Psychotria species, which have already become dioecious. The flowers of the long (L)-styled (pin) morph of the Bonin Islands plants are completely self- and intramorph incompatible, while the flowers of the short (S)-styled (thrum) morphs are partially intramorph compatible. The S-styled morph was more abundant in the examined populations from three islands of the archipelago, probably due to the existence of partial S–S intramorph compatibility.     

Phylogenetic analysis of the generaCladonia andCladina (Cladoniaceae, lichenizedAscomycota)

A cladistic analysis of 44 species of the generaCladina andCladonia is presented.Pycnothelia papillaria, Cladia aggregata andC. retipora were used as outgroup taxa. The consensus of all equally parsimonious trees suggests a common ancestral origin for species inCladonia andCladina while the generaPycnothelia andCladia cluster outside this group. The results do not support distinction ofCladina at genus level although it is distinguished as a monophyletic group but within the genusCladonia. The current sectional division ofCladonia is not supported. SectionsCocciferae andHelopodium are best supported but even these groups as currently delimited seem to in include elements that are more closely related to species of the other sections, e.g.C. carneola of sect.Cocciferae andC. pityrophylla of sect.Helopodium. Sections such asUnciales andPerviae seem to be artificial assemblages. In general the evolutionary scheme of the genus seems to be more complicated than revealed by the current sectional division.     

A review on the impacts of feral cats (Felis silvestris catus) in the Canary Islands: implications for the conservation of its endangered fauna

Feral cats have been directly responsible for the extinction of numerous species on islands worldwide, including endemic species of mammals, birds and reptiles. The diet of feral cats in the main habitats of the Canary Islands, as generally occurred on oceanic islands, is mainly composed of introduced mammals, and native species of birds, reptiles and insects. The impact of feral cat upon the endangered species was assessed by evaluating their relative abundance in the cats’ diet and by considering their current conservation status. A total of 68 different preys were identified at species level in all studies carried out in the Canary Islands (5 mammals, 16 birds, 15 reptiles and 32 invertebrates). From all the species preyed by feral cats in the Canary Islands, only four of them are considered threatened by the IUCN Red List of Threatened Species: one endemic bird Saxicola dacotiae and three endemic giant lizards, Gallotia simonyi, Gallotia intermedia, and Gallotia gomerana. Although some efforts on management control have been carried out, it is necessary to enforce these conservation activities on those areas of Tenerife, La Gomera and El Hierro where giant lizards are still present. Furthermore some local areas where endangered bird species are highly predated should be protected. Nevertheless, it is important to take into account the presence of other introduced species such as rats, mice or rabbits in order to avoid problems derived from the hyperpredation process and mesopredator release effect.