Conservation genetics of the endangered endemic Sambucus palmensis Link (Sambucaceae) from the Canary Islands

Five polymorphic microsatellites (simple sequence repeat; SSR) markers were used to estimate the levels of genetic variation within and among natural populations from different islands of the endangered endemic from the Canary Islands Sambucus palmensis Link (Sambucaceae). Genetic data were used to infer potential evolutionary processes that could have led to present genetic differentiation among islands. The levels of genetic variability of S. palmensis were considerably high; proportion of polymorphic loci (P = 100%), mean number of alleles per locus (A = 6.8), average expected heterozygosity (He = 0.499). In spite of its small population size and endemic character, 58 different multilocus genotypes were detected within the 165 individuals analyzed. All samples located in different islands always presented different multilocus genotypes. Principal Coordinates Analysis, genetic differentiation analysis (F _ST and G _ST ′) and Bayesian Cluster Analysis revealed significant genetic differences among populations located in different islands. However, this genetic differentiation was not recorded among Tenerife and La Gomera populations, possibly revealing the uncontrolled transfer of material between both islands. AMOVA analysis attributed 77% of the variance to differences within populations, whereas 8% was distributed between islands. The levels of genetic differentiation observed among populations, and the genetic diversity distribution within populations in S. palmensis, indicate that management should aim to conserve as many of the small populations as possible. Concentrating conservation efforts only on the few large populations would result in the likelihood of loss of genetic variability for the species.     

Local endemism and within‐island diversification of shrews illustrate the importance of speciation in building Sundaland mammal diversity

Island systems are important models for evolutionary biology because they provide convenient, discrete biogeographic units of study. Continental islands with a history of intermittent dry land connections confound the discrete definitions of islands and have led zoologists to predict (i) little differentiation of terrestrial organisms among continental shelf islands and (ii) extinction, rather than speciation, to be the main cause of differences in community composition among islands. However, few continental island systems have been subjected to well‐sampled phylogeographic studies, leaving these biogeographic assumptions of connectivity largely untested. We analysed nine unlinked loci from shrews of the genus Crocidura from seven mountains and two lowland localities on the Sundaic continental shelf islands of Sumatra and Java. Coalescent species delimitation strongly supported all currently recognized Crocidura species from Sumatra (six species) and Java (five species), as well as one undescribed species endemic to each island. We find that nearly all species of Crocidura in the region are endemic to a single island and several of these have their closest relative(s) on the same island. Intra‐island genetic divergence among allopatric, conspecific populations is often substantial, perhaps indicating species‐level diversity remains underestimated. One recent (Pleistocene) speciation event generated two morphologically distinct, syntopic species on Java, further highlighting the prevalence of within‐island diversification. Our results suggest that both between‐ and within‐island speciation processes generated local endemism in Sundaland, supplementing the traditional view that the region's fauna is relictual and primarily governed by extinction.     

Phylogeographical structure in the coastal species Senecio rodriguezii (Asteraceae), a narrowly distributed endemic Mediterranean plant

Aim Our goals were (1) to assess the levels of chloroplast DNA variation in a narrowly distributed plant restricted to continental islands, (2) to ascertain whether a phylogeographical structure is present in plants restricted to coastal linear systems, and (3) to interpret the results in the light of the known palaeogeography of these islands. Location The Eastern Balearic Islands (Majorca and Minorca) in the Western Mediterranean Basin. Methods Sampling included 134 individuals from 28 populations of Senecio rodriguezii covering the entire range of the species. Sequences of the chloroplast genome (trnT–trnL spacer) were obtained and parameters of population genetic diversity and substructure were determined (h_sh_t, G_st). The geographical structure of genetic variation was assessed by an analysis of molecular variance (AMOVA). Additionally, a spatial AMOVA (SAMOVA) was used to identify groups of populations that were geographically homogeneous and maximally differentiated from each other. Finally, a pattern of isolation by distance was assessed by testing the correlation between the matrix of pairwise Φ_ST values and the matrix of geographical distances between pairs of populations using a Mantel test. Results Seven haplotypes were detected in S. rodriguezii. Only two of them were shared between islands; all of the others were restricted to Majorca (two) or Minorca (three). Overall, we found high levels of genetic diversity and significant geographical structuring of cpDNA markers. Most of the variation detected can be attributed to differences among populations (84.6%), but there was also a significant differentiation between the islands. Main conclusions Our results support the view that the Balearic Islands constitute a reservoir of genetic diversity, not only for widespread Mediterranean taxa, but also for endemic ones. The intraspecific genetic structure found in S. rodriguezii suggests that its population history was dominated by both expansion and contraction events. This has resulted in a species that is highly structured genetically, showing very few shared haplotypes between islands, and a high number of haplotypes restricted to small geographical areas within the islands. Changes in habitat availability and dynamic processes of population fragmentation and connectivity due to repeated cycles of sea‐level changes during the Quaternary are the possible underlying factors that have shaped the cpDNA pool of this endemic species on a regional scale.     

Phylogeography of Eastern Polynesian sandalwood (Santalum insulare), an endangered tree species from the Pacific: a study based on chloroplast microsatellites

Aim Patterns of genetic variation within forest species are poorly documented in island ecosystems. The distribution of molecular variation for Santalum insulare, an endangered tree species endemic to the islands of eastern Polynesia, was analysed using chloroplast microsatellite markers. The aims were to quantify the genetic diversity; to assess the genetic structure; and to analyse the geographical distribution of the diversity within and between archipelagoes. The ultimate goal was to pre‐define evolutionary significant units (ESUs) for conservation and restoration programmes of this species, which constitutes a natural resource on small, isolated islands. Location Eleven populations, each representative of one island, covering most of the natural occurrence of S. insulare were sampled: five populations from the Marquesas Archipelago; three from the Society Archipelago; and three from the Cook–Austral Archipelago. These South Pacific islands are known for their high degree of plant endemism, and for their human occupation by Polynesian migrations. The extensive exploitation of sandalwood by Europeans nearly 200 years ago for its fragrant heartwood, used overseas in incense, carving and essential oil production for perfume, has dramatically reduced the population size of this species. Methods We used chloroplast microsatellites, which provide useful information in phylogeographical forest tree analyses. They are maternally inherited in most angiosperms and present high polymorphism. Among the 499 individuals sampled, 345 were genotyped successfully. Classical models of population genetics were used to assess diversity parameters and phylogenetic relationships between populations. Results Four microsatellite primers showed 16 alleles and their combinations provided 17 chlorotypes, of which four exhibited a frequency > 10% in the total population. The gene diversity index was high for the total population (H_e = 0.82) and varied among archipelagoes from H_e = 0.40 to 0.67. Genetic structure is characterized by high levels of differentiation between archipelagoes (36% of total variation) and between islands, but differentiation between islands varied according to archipelago. The relationship between genetic and geographical distance confirms the low gene flow between archipelagoes. The minimum spanning tree of chlorotypes exhibits three clusters corresponding to the geographical distribution in the three main archipelagoes. Main conclusions The high level of diversity within the species was explained by an ancient presence on and around the hotspot traces currently occupied by young islands. Diversity in the species has enabled survival in a range of habitats. Relationships between islands show that the Cook–Austral chlorotype cluster constitutes a link between the Marquesas and the Society Islands. This can be explained by the evolution of the island systems over millions of years, and extinction of intermediary populations on the Tuamotu Islands following subsidence there. Based on the unrooted neighbour‐joining tree and on the genetic structure, we propose four ESUs to guide the conservation and population restoration of Polynesian Sandalwood: the Society Archipelago; the Marquesas Archipelago; Raivavae Island; and Rapa Island.     

Genetic diversity in micronesian island populations of the tropical tree campnosperma brevipetiolata (anacardiaceae)

Tropical plant species have been the focus of considerable attention in regard to their potential economic and social importance in the face of rapidly diminishing biodiversity in the tropics. Pacific Island species represent an even more fragile resource because different island populations are widely scattered and overall population sizes are small. We examined the distribution of genetic variation in Campnosperma brevipetiolata (Anacardiaceae), an upland rainforest tree species that is of potential use for both lumber and reforestation efforts in the Federated States of Micronesia (FSM). Seeds were collected from multiple populations on four island groups in the Caroline Islands (Kosrae, Pohnpei, and Yap in the FSM; and the Republic of Palau) and subjected to an electrophoretic analysis involving four polymorphic genetic loci. We hypothesized that variation on these islands would decrease with increasing distance from the presumed Indo-Malayan source of these island floras. Indeed, we found a trend of decreasing variation from west to east indicated by the mean number of alleles per locus (A = 1.50-1.33), effective number of alleles per locus (A_e = 1.14-1.12) and mean genetic diversity (H_e = 0.123-0.107). We also found little genetic differentiation among the islands (F_pt = 0.174) and among subpopulations within islands (F_sp = 0.047), indicating that either there are high levels of gene flow among the islands by seed dispersal or that these populations have not been established long enough for divergence to have occurred. The lack of divergence among islands observed for Campnosperma brevipetiolata suggests that germplasm sampled from any one island population would be a suitable starting point for plant breeding or reforestation efforts.     

Genetic and morphometric diversity in Wallacea: geographical patterning in the horse shoe bat, Rhinolophus affinis

Genetic and morphometric variation was examined in eleven island populations of the horse‐shoe bat, Rhinolophus affinis, at the easterly end of this widespread species’ range and encompassing the Australian–Oriental biogeographic interface. Allozyme variation revealed mean heterozygosity levels within islands of 0.047, which is near the mammalian average. However, heterozygosity tended to decline from west to east as populations approached the periphery of the species’ distribution, and was lowest in those islands that were separated by the greatest sea‐crossing from source populations. There is extensive between‐island genetic differentiation (mean F_ST = 0.40) and relationships between islands are associated with their arrangement in geographical space; genetic distance is correlated with geographical distance and the genetic arrangement of islands is associated with longitude. The arrangement of islands as indicated by variation in body and skull metrics is also associated with their geographical positions, and the metric and genetic measures are themselves associated. While other taxa in the region have shown genetic‐geographical concordances, R. affinis is the only one that displays concordant patterns in metrical features. These patterns in biological diversity are interpreted as arising from the sequential island population structure and clines in key biogeographic gradients.     

Persistent inter‐ and intraspecific gene exchange within a parallel radiation of caterpillar hunter beetles (Calosoma sp.) from the Galápagos

When environmental gradients are repeated on different islands within an archipelago, similar selection pressures may act within each island, resulting in the repeated occurrence of ecologically similar species on each island. The evolution of ecotypes within such radiations may either result from dispersal, that is each ecotype evolved once and dispersed to different islands where it colonized its habitat, or through repeated and parallel speciation within each island. However, it remains poorly understood how gene flow during the divergence process may shape such patterns. In the Galápagos islands, three phenotypically similar species of the beetle genus Calosoma occur at higher elevations of different islands, while lowlands are occupied by a fourth species. By genotyping all major populations within this radiation for two nuclear and three mitochondrial gene fragments and seven microsatellite markers, we found strong support that the oldest divergence separates the highland species of the oldest island from the remaining species. Despite their morphological distinctness, highland species of the remaining islands were genetically closely related to the lowland population on each island and within the same magnitude as lowland populations sampled at different islands. Repeated evolution of highland ecotypes out of the lowland species appears the most likely scenario and estimates of geneflow rates revealed extensive admixture among ecotypes within islands, as well as between islands. These findings indicate that gene exchange among the different populations and species may have shaped the phylogenetic relationships and the repeated evolution of these ecotypes.     

Rapid morpho‐functional changes among insular populations of the greater white‐toothed shrew

Islands are often considered to be natural laboratories where repeated ‘evolutionary experiments’ have taken place. Consequently, islands have been key model systems in our understanding of evolutionary theory. The greater white‐toothed shrew (Crocidura russula) is of interest as it has invaded French Atlantic islands within the last few thousand years and is considered to be morphologically and genetically stable in this area. In this article, we study the shape of the mandible of the greater white‐toothed shrew on four islands and compare it with that of individuals from populations on the mainland to quantify the effects of insularity. The degree of insularity (i.e. island size and distance to the continent) is thought to be linked to differences in ecological characteristics of islands compared with the mainland. We used geometric morphometric analyses to quantify differences in size and shape between populations and employed a simple biomechanical model to evaluate the potential effects of shape differences on bite force. Specimens from island populations are different from continental populations in shape and mechanical potential of the mandible. Among islands, the mandible shows various shapes that are correlated with both the distance from the coast and island area. The shape differences are located on different parts of the mandible, suggesting different ecological constraints on each island. Moreover, these shapes are linked to the ‘mechanical potential’, which is markedly different between islands. Mechanical potential has been suggested to evolve in response to prey size and or mechanical properties. In conclusion, our results show that, in spite of the relatively recent colonization of the Atlantic Islands, the mandible of C. russula possesses a distinct shape. Moreover, the shape differs among islands and is probably linked to the consumption of different prey. © 2012 The Linnean Society of London, Biological Journal of the Linnean Society, 2012, ?? , ??–??.     

Recent origin and adaptive diversification of <Emphasis Type="Italic">Ainsliaea</Emphasis> (Asteraceae) in the Ryukyu Islands: molecular phylogenetic inference using nuclear microsatellite markers

We investigated the phylogeny and phylogeography of four closely related Ainsliaea species (Asteraceae) on the continental Ryukyu Islands of eastern Asia, which consist of two flood-adapted “rheophyte” and two non-rheophyte (inland) species, based on 12 nuclear microsatellite loci. Phylogenetic analyses using 420 individuals from 26 populations showed that rheophytic A. linearis and A. oblonga are genetically distinct. Each species was clustered with the inland species that occur on the same islands, suggesting a different ancestry for the two rheophytes that evolved independently by local adaptation to flooded habitats. The results from the neighbor-joining clustering and principal coordinate analysis (PCoA) indicate that the southern populations of A. macroclinidioides are distinct lineages and ancestral to the northern populations as well as the other diverse species complex in the Ryukyus. These results suggest a pattern of colonization initially from the Asian mainland to the southern islands, followed by the northern islands via land bridges generated during the Quaternary glaciations. After isolation from southern populations, species radiation and regional differentiation within the northern clade occurred possibly via local adaptation and/or geographic isolation of the subdivided island bridge.     

Adaptive radiation versus intraspecific differentiation: morphological variation in Caribbean Anolis lizards

Lizards in the genus Anolis have experienced adaptive radiation in the Greater Antilles, producing a suite of species morphologically adapted to use different parts of the environment. In the Lesser Antilles, adaptive radiation has not occurred, but on some islands, interpopulational variation is high and represents adaptation to different habitats. We compared the extent of morphological differentiation among Greater Antillean habitat specialists with that exhibited among populations of two species, Anolis marmoratus and A. oculatus, from the Lesser Antillean islands of Guadeloupe and Dominica. Although extensive, intraspecific divergence in the Lesser Antilles is substantially less in magnitude than the differences among habitat specialists in the Greater Antilles. All populations of A. marmoratus are most similar to Greater Antillean trunk‐crown habitat specialists, but populations of A. oculatus differ in their affinities: some are similar to trunk‐crown anoles, but others are more similar to trunk‐ground habitat specialists.     

Adaptive radiation and genetic differentiation in the woodySonchus alliance (Asteraceae:Sonchinae) in the Canary Islands

The woodySonchus alliance consists of 19 species ofSonchus subg.Dendrosonchus, one species ofSonchus subg.Sonchus and species of five genera (i.e.Babcockia, Sventenia, Taeckholmia, Lactucosonchus, Prenanthes), and is restricted primarily to the archipelago of the Canaries in the Macaronesian phytogeographical region. An enzyme electrophoretic study, including 13 loci, was conducted to assess genetic diversity within and divergence among species of the alliance. Nei's genetic identities (distances) between genera and/or subgenera range from 0.490 (0.714) to 0.980 (0.013), and pairwise comparisons of all populations show relatively high genetic identities, with a mean of 0.804. The high identities further support the genetic cohesiveness of the alliance and its single origin on the Macaronesian islands. Species in the alliance also show about 50%; higher total genetic diversity (H_T) than the mean for other oceanic endemics. There is greater divergence between endemics or species on older islands compared to those on younger islands, which suggests that time is a factor for divergence at allozyme loci. Furthermore, populations on older islands have higher total genetic diversities and lower identities than conspecific populations on younger islands. These results imply early colonization, radiation, and divergence of the woodySonchus alliance on older islands followed by subsequent colonization to younger islands. The taxonomic distribution of alleles in the alliance indicates lineage sorting also played a role in divergence among species. Lineage sorting may also produce nonconcordance with either taxonomic designation or the pattern of variation obtained from other molecular markers such as ITS sequences of nrDNA. Timing for the origin and radiation of the alliance agrees with the estimate based on ITS sequences, and suggests that the early divergence and rapid radiation took place during the Late Tertiary on either Gran Canaria or Tenerife.     

Allozyme diversity in the federally threatened golden paintbrush, <Emphasis Type="Italic">Castilleja levisecta</Emphasis> (Scrophulariaceae)

Castilleja levisecta (Scrophulariaceae), the golden paintbrush, is an insect-pollinated herbaceaous perennial found in the Pacific Northwest. Currently restricted to two island populations off British Columbia and nine populations (eight on islands) in Washington, C. levisecta is a rare species threatened with extinction. Allozymes were used to describe genetic diversity and structure in these eleven populations. Despite its threatened status and small geographic range, exceptionally high levels of genetic diversity are maintained within C. levisecta. All sixteen of the loci resolved were polymorphic within the species (P_s=100%), while the mean percentage of loci polymorphic within populations (P_p) was 65.7%. The mean number of alleles per polymorphic locus (AP_s) was 2.94 within the species and averaged 2.38 within populations (AP_p). Genetic diversity (H_es) was 0.285 for the species, whereas mean population genetic diversity (H_ep) was 0.213. Smaller populations had, on average, fewer observed alleles and less genetic diversity. A significant negative correlation (r = –0.72) was found between genetic identity and geographic distance, indicating reduced gene flow between distant populations. The most geographically isolated population was one of the larger populations, one of the most genetically diverse and the most genetically divergent. A wide range of pairwise population genetic identities (I = 0.771 – 0.992) was found, indicating considerable genetic divergence between some populations. Overall, 19% of the total genetic diversity was distributed among populations. Results of this survey indicate that genetic augmentation of existing populations is unnecessary. The high allelic diversity found for the species and within its populations holds promise for conservation and restoration efforts to save this rare and threatened plant species.     

Allozyme Variation and the Genetic Structure of Populations of Trochodendron aralioides, a Monotypic and Narrow Geographic Genus

Trochodendron aralioides Siebold & Zucc., a primitive angiosperm that is insect-pollinated, has wind-dispersed seeds and grows only in eastern Asia. Nineteen populations were analyzed, including 16 from Taiwan, two from the main islands of Japan and one from Iriomote Island in the Ryukyu Islands. Genetic variability (0.133) at the species level was more similar to that of a narrow geographic species than an endemic species. Based on genotype frequencies, mating within populations was non-random (F _{I s}=0.065) and there was significant genetic differentiation (F _ST =0.164) among populations. Genetic diversity was largely within, rather than among, populations (H _s =0.132, 85.14%). Bisexual flowers of T. aralioides exhibit synchronized dimorphism. The protogynous and protandrous morphs are self-incompatible, which probably promotes outcrossing and contributes to the high levels of heterozygosity within populations. Using a maximum likelihood tree, populations of T. aralioides were separated into two distinct clusters: Japan and Taiwan/ Iriomote Island. Populations of T. aralioides on Iriomote Island and in northern Taiwan exhibited a high degree of genetic similarity. The isolation-by-distance model does not fit populations in Taiwan, but suitably describes the relationship between populations of T. aralioides in Japan and Taiwan. Received 28 April 2000/ Accepted in revised form 13 December 2000     

Microsatellites for tree of heaven (Ailanthus altissima)

Nine (CT)_n microsatellites were developed for tree of heaven, Ailanthus altissima, from invasive populations on the Mediterranean islands. These loci had seven to 12 alleles in 96 trees from five islands. Two loci had significant deficits of heterozygotes within islands while the other loci were in Hardy–Weinberg equilibrium, and four pairs of loci had significant linkage disequilibrium within a single island. These loci were also polymorphic in one to three individuals of the tree of heaven varieties, Ailanthus altissima erythrocarpa, Ailanthus altissima sutchuenensis and Ailanthus altissima tanakai, and the related species Ailanthus giraldii and Ailanthus vilmariniana.     

Larger body size on islands affects silvereye Zosterops lateralis song and call frequency

Differences in song repertoires and characteristics of island and mainland populations of the same avian species are usually explained by dispersal, cultural evolution and/or habitat differences. The influence of morphology is often overlooked, even though island populations are frequently morphologically distinct from mainland populations, and morphology could affect vocalizations. I compared morphological features, songs, contact calls and alarm calls of six isolated island populations of silvereye Zosterops lateralis with those of two mainland populations to examine whether differences between mainland and island vocalizations were consistent across vocalization types, and whether these differences could be linked to morphological differences. Vocalizations were lower in frequency on islands. Island individuals were larger (both in mass and body structure), and body mass was an important predictor of frequency in contact and alarm calls. I argue that this strong association results from the island rule (islands promote larger body sizes) and cascading effects of morphology on vocalization frequency in this species.     

Cryptic diversity within the endemic prehensile‐tailed gecko Urocotyledon inexpectata across the Seychelles Islands: patterns of phylogeographical structure and isolation at the multilocus level

The deciphering of the process of genetic differentiation of species with insular distributions is relevant for biogeographical and conservation reasons. Despite their importance as old gondwanic islands and part of the western Indian Ocean biodiversity hotspot, little is known about the genetic structure of taxa from the Seychelles Islands. We have examined the patterns of structure and isolation within Urocotyledon inexpectata (Reptilia: Geckkonidae), an endemic species from this archipelago. Genetic diversity was screened from populations across the archipelago for both mitochondrial and nuclear genes. Gene genealogies and model‐based inference were used to explore patterns and timings of isolation between the main lineages. High levels of genetic diversity were found for the mitochondrial and some of the nuclear markers. This species harbours at least two highly differentiated lineages, exclusively distributed across the northern and southern groups of the islands. The main split between these was dated back to the Miocene–late Pliocene, but isolation events throughout the Pliocene and Pleistocene were also inferred. Migration between groups of islands was apparently nonexistent, except for one case. The low dispersal capabilities of this species, together with the intrinsic fragmented nature of its geographical distribution, seem to have resulted in highly structured populations, despite the cyclic periods of contact between the different island groups. These populations may currently represent more than one species, making U. inexpectata another example of a morphologically cryptic lineage with deep genetic divergence within gekkonids. The observed patterns suggest a hypothetical biogeographic scenario (of a main north–south phylogeographic break) for the Seychelles that can be further tested with the exploration of the phylogeographic structure of other Seychellois taxa. © 2011 The Linnean Society of London, Biological Journal of the Linnean Society, 2011, 104 , 177–191.     

Genetic variations and phylogeography of the swallowtail butterfly Papilio machaon on the Japanese Islands

The swallowtail butterfly Papilio machaon Linnaeus, 1758 is widely distributed in the Holarctic region, including all of the main islands of Japan, as well as Sakhalin, and on other smaller islands south to Yakushima Island. The Japanese population is situated at the margin of the Eurasian distribution range of this species. It is morphologically different from other populations and has been classified as the subspecies hippocrates C. & R. Felder, 1864. The population of the Japanese Islands is considered to be genetically distinct from the continental populations in relation to the geographical history of the Japanese Islands. Therefore, we examined a part of the ND5 gene sequence of the mitochondrial DNA for P. machaon individuals of various localities in Japan and some nearby countries, and found 68 haplotypes in 400 individuals from the Japanese Islands and Sakhalin. A DNA polymorphism analysis revealed that the genetic structure of the Hokkaido population was significantly different from that of the southern populations on the main Japanese islands. These results imply that P. machaon expanded its range from the Amur region of Russia southward through Sakhalin to the Japanese Islands, and that the Tsugaru Strait between Hokkaido and Honshu may have subsequently limited their gene flow as a geographical barrier.     

Genetic diversity within and among populations of Shorea leprosula Miq. and Shorea parvifolia Dyer (Dipterocarpaceae) in Indonesia detected by AFLPs

The genetic diversity within and among populations of Shorea leprosula and Shorea parvifolia from Indonesia was investigated using amplified fragment length polymorphisms (AFLPs). The results indicated that S. leprosula is genetically more variable than S. parvifolia. At the population level, a higher level of genetic diversity was revealed for S. leprosula with a percentage of polymorphic loci (PPL_p) of 53.32% and an expected heterozygosity (H _ep) of 0.16 in comparison with S. parvifolia showing PPL_p of 51.79% and H _ep of 0.14. At the species level, S. leprosula showed PPL_s of 92.86% and H _es of 0.21, while S. parvifolia showed PPL_s of 85.71% and H _es of 0.21. Genetic differentiation (G _st) indicated that 25 and 31% of total genetic diversity in S. leprosula and S. parvifolia, respectively, were attributed to the differences among populations. An analysis of molecular variance (AMOVA) at two hierarchical levels exhibited that most genetic variation resided within populations with proportion of 70.2% for S. leprosula and 66.2% for S. parvifolia. The AMOVA at three hierarchical levels performed for S. leprosula and S. parvifolia together revealed that the genetic difference between the two species was remarkably higher with a proportion of 44.1% than the differences within and among populations (38.1 and 17.8%, respectively). The genetic differentiation between islands was significant for S. leprosula but not for S. parvifolia. The observed genetic diversity agreed with the life history traits of Shorea species. Highly differentiating individual AFLP markers were found for each species, which will serve as diagnostic markers for the identification of wood of different species, from different islands and regions.     

ALLOZYME VARIATION WITHIN AND BETWEEN LASTHENIA MINOR AND ITS DERIVATIVE SPECIES,L. MARITIMA (ASTERACEAE)

Enzyme electrophoresis was employed to examine genetic variation at 20 loci in 16 populations of Lasthenia minor and 18 populations of its presumed derivative species L. maritima. The purposes of the study were to ascertain levels of genetic variation in each species, to assess how the variation at enzyme-coding genes is apportioned within and among populations of each species, and to determine the level of divergence between the two species. The two species are both diploid annuals, similar morphologically, and produce fertile F₁ hybrids when crossed. Lasthenia minor is self-incompatible and restricted to mainland California, whereas L. maritima is self-compatible and probably largely autogamous; it occurs on seabird rocks from central California to British Columbia. Mean genetic identities for pair-wise comparisons of populations of the two species are similar to values for populations of the same species, indicating they have not diverged at the 20 genes coding for soluble enzymes. Despite its more extensive geographical range, L. maritima exhibits only 50% of the genetic diversity of L. minor. The latter species apportions a greater amount of its diversity within populations, whereas the former harbors more diversity among populations than within them. This is probably a reflection of the different breeding systems of the two species. Six unique alleles were detected in L. minor, whereas only one novel allele was found in a single individual of L. maritima. The electrophoretic data are concordant with the suggestion that L. maritima is relatively recently derived from L. minor. The switch from outcrossing to selfing and selection of genotypes adapted to the chemically and physically unusual substrate on the seabird rocks are considered the critical steps in the evolution of L. maritima.