Multi-gene phylogenetic analysis of south-east Asian pest members of the Bactrocera dorsalis species complex (Diptera: Tephritidae) does not support current taxonomy

Bactrocera dorsalis sensu stricto, B. papayae, B. philippinensis and B. carambolae are serious pest fruit fly species of the B. dorsalis complex that predominantly occur in south-east Asia and the Pacific. Identifying molecular diagnostics has proven problematic for these four taxa, a situation that cofounds biosecurity and quarantine efforts and which may be the result of at least some of these taxa representing the same biological species. We therefore conducted a phylogenetic study of these four species (and closely related outgroup taxa) based on the individuals collected from a wide geographic range; sequencing six loci (cox1, nad4-3′, CAD, period, ITS1, ITS2) for approximately 20 individuals from each of 16 sample sites. Data were analysed within maximum likelihood and Bayesian phylogenetic frameworks for individual loci and concatenated data sets for which we applied multiple monophyly and species delimitation tests. Species monophyly was measured by clade support, posterior probability or bootstrap resampling for Bayesian and likelihood analyses respectively, Rosenberg's reciprocal monophyly measure, P(AB), Rodrigo's (P(RD)) and the genealogical sorting index, gsi. We specifically tested whether there was phylogenetic support for the four 'ingroup' pest species using a data set of multiple individuals sampled from a number of populations. Based on our combined data set, Bactrocera carambolae emerges as a distinct monophyletic clade, whereas B. dorsalis s.s., B. papayae and B. philippinensis are unresolved. These data add to the growing body of evidence that B. dorsalis s.s., B. papayae and B. philippinensis are the same biological species, which poses consequences for quarantine, trade and pest management.

A systematic review of community-based interventions for emerging zoonotic diseases in Southeast Asia

Background Southeast Asia has been at the epicentre of recent epidemics of emerging and re-emerging zoonotic diseases. Community-based surveillance and control interventions have been heavily promoted but the most effective interventions have not been identified. Objectives This review evaluated evidence for the effectiveness of community-based surveillance interventions at monitoring and identifying emerging infectious disease; the effectiveness of community-based control interventions at reducing rates of emerging infectious disease; and contextual factors that influence intervention effectiveness. Inclusion criteria Participants Communities in Brunei, Cambodia, Indonesia, Laos, Malaysia, Myanmar, the Philippines, Singapore, Thailand and Viet Nam. Types of intervention(s) Non-pharmaceutical, non-vaccine, and community-based surveillance or prevention and control interventions targeting rabies, Nipah virus , dengue, SARS or avian influenza. Types of outcomes Primary outcomes: measures: of infection or disease; secondary outcomes: measures of intervention function. Types of studies Original quantitative studies published in English. Search strategy Databases searched (1980 to 2011): PubMed, CINAHL, ProQuest, EBSCOhost, Web of Science, Science Direct, Cochrane database of systematic reviews, WHOLIS, British Development Library, LILACS, World Bank (East Asia), Asian Development Bank. Methodological quality Two independent reviewers critically appraised studies using standard Joanna Briggs Institute instruments. Disagreements were resolved through discussion. Data extraction A customised tool was used to extract quantitative data on intervention(s), populations, study methods, and primary and secondary outcomes; and qualitative contextual information or narrative evidence about interventions. Data synthesis Data was synthesised in a narrative summary with the aid of tables. Meta-analysis was used to statistically pool quantitative results. Results Fifty-seven studies were included. Vector control interventions using copepods, environmental cleanup and education are effective and sustainable at reducing dengue in rural and urban communities, whilst insecticide spraying is effective in urban outbreak situations. Community-based surveillance interventions can effectively identify avian influenza in backyard flocks, but have not been broadly applied. Outbreak control interventions for Nipah virus and SARS are effective but may not be suitable for ongoing control. Canine vaccination and education is more acceptable than culling, but still fails to reach coverage levels required to effectively control rabies. Contextual factors were identified that influence community engagement with, and ultimately effectiveness of, interventions. Conclusion Despite investment in community-based disease control and surveillance in Southeast Asia, published evidence evaluating interventions is limited in quantity and quality. Nonetheless this review identified a number of effective interventions, and several contextual factors influencing effectiveness. Identification of the best programs will require comparative evidence of effectiveness acceptability, cost-effectiveness and sustainability.

Convergence in the ownership and governance of East Asian firms

In the past decade, policymakers in over 70 markets have introduced corporate governance codes or best practice guidelines. In East Asia, they have been introduced in Hong Kong in 1999 and 2006, Indonesia in 2000 and 2007, Malaysia in 2000 and 2007, the Philippines in 2002, Singapore iu 2001 and 2005, South Korea in 2003, Taiwan iu 2002 and Thailand iu 2006. The common focus of these codes is to encourage but not force companies to improve their corporate governance practices to a specified target level, e.g., board independence of 30%. Another commonality is that the guidelines apply to all listed companies regardless of their ownership structure or other characteristics.

The plain truth: Australia, tobacco control, and Southeast Asia

As of December 2012, Australia will be the first country in the world where tobacco products will be sold in olive plain packaging without branding. There has been much discussion as to whether this pioneering initiative will spark an ‘Olive Revolution’ around the world — with other countries adopting the plain packaging of tobacco products. Australia’s neighbours in Southeast Asia would particularly benefit from tobacco control measures such as the plain packaging.

Introduction: Mapping the Contours of East Asian Commercial Law for the Asian Century

The centre of economic gravity in the new century is shifting to the East. Since 200 1, according to the International Monetary Fund (IMF), Asia's contribution to world economic growth has matched that of the United States and Europe combined, and, since 2006, has even exceeded it (IMF, 20 I I; Neumann and Arora, 20 II ). This surge is easy to explain: China has emerged as a global super-power; Japan remains the third-largest world economy, despite only recently emerging from over twenty years of economic stagnation (The Age, 2013); South Korea and the ' tiger ' economies of Taiwan, Hong Kong and Singapore have achieved high-level economic development through capital investment and technological innovation; and Indonesia, Thailand, the Philippines and Malaysia have supplied riches in labour and resources to the regional economy (Macintyre and Naughton, 2005, p. 78). A growing middle class is lifting consumption. ‘Billions of Asians,' writes Mahbubani (2008, p. 3), 'are marching to modernity.’ This book examines scholarly interpretations for the role commercial law has played in East Asia's economic rise. At first blush, this might seem a daunting task. After all, as some theorists have argued, the East Asian experience is largely neglected in writings on Jaw generally and commercial law more broadly (Wolff, 20 12). This is because law, as a discipline, was largely forged in the prior European and American centuries; these 'Anglo-American moorings' ill-serve legal analysis in the new Asian Century (Cossman, 1997, p. 539).

Genetic and Malt Quality Diversity of East Asian Two-Rowed Barley Accessions in Relationship to North American Malting Barley.

Because of epidemics of Fusarium head blight (FHB; caused by Fusarium graminearum Schwabe [teleomorph Gibberella zeae (Schwein.) Petch]) in the northern Great Plains of the United States and Canada in the past two decades, malting barley breeders have been forced to use nonadapted barley (Hordeum vulgare L.) accessions as sources of FHB resistance. Many of the resistant accessions are from East Asia, and limited information is available on their genetic diversity and malt quality. The objectives of this study were to determine the genetic diversity among 30 East Asian accessions and two North American cultivars. Genetic diversity was based on 49 simple-sequence repeat markers. All accessions were tested for barley grain brightness; protein content; 1,000-kernel weight; malting loss; fine-grind malt extract; content of plump kernels, free amino nitrogen, soluble protein, and wort beta-glucan; the Kolbach index (i.e., the ratio of malt soluble protein to malt total protein); a-amylase activity; diastatic power; won color; and wort viscosity. A few accessions had equal quality compared with Harrington and Conlon barley for individual traits but not for all. Qing 2, Mokkei 93-78, and Nitakia 48 could be excellent sources for increased malt extract; Nitakia 48 is a possible source for low wort viscosity; and Mokkei 93-78 and Nitakia 48 are putative sources of low beta-glucan content. The cluster analyses also implied that the malt quality of an accession cannot be predicted based on the country where it was developed.

Community-based Natural Resource Management: Issues and Cases from South Asia

Community-based natural resource management (CBNRM) is the joint management of natural resources by a community based on a community strategy, through a participatory mechanism involving all legitimate stakeholders. The approach is community-based in that the communities managing the resources have the legal rights, the local institutions and the economic incentives to take substantial responsibility for sustained use of these resources. This implies that the community plays an active role in the management of natural resources, not because it asserts sole ownership over them, but because it can claim participation in their management and benefits for practical and technical reasons1–4. This approach emerged as the dominant conservation concept in the late 1970s and early 1980s, of the disillusionment with the developmental state. Governments across South and South East Asia, Africa and Latin America have adopted and implemented CBNRM in various ways, viz. through sectoral programmes such as forestry, irrigation or wildlife management, multisectoral programmes such as watershed development and efforts towards political devolution. In India, the principle of decentralization through ‘gram swaraj’ was introduced by Mahatma Gandhi. The 73rd and 74th constitution amendments in 1992 gave impetus to the decentralized planning at panchayat levels through the creation of a statutory three-level local self-government structure5,6. The strength of this book is that it includes chapters by CBNRM advocates based on six seemingly innovative initiatives being implemented by nongovernmental organizations (NGOs) in ecologically vulnerable regions of South Asia: two in the Himalayas (watershed development programme in Lingmutechhu, Bhuthan and Thalisain tehsil, Paudi Grahwal District, Uttarakhand), three in semi-arid parts of western India (watershed development in Hivre Bazar, Maharashtra and Nathugadh village, Gujarat and water-harvesting structures in Gopalapura, Rajasthan) and one in the flood-plains of the Brahmaputra–Jamuna (Char land, Galibanda and Jamalpur districts, Bangladesh). Watersheds in semi-arid regions fall in the low-rainfall region (500–700 mm) and suffer the vagaries of drought 2–3 years in every five-year cycle. In all these locations, the major occupation is agriculture, most of which is rainfed or dry. The other two cases (in Uttarakhand) fall in the Himalayan region (temperate/sub-temperate climate), which has witnessed extensive deforestation in the last century and is now considered as one of the most vulnerable locations in South Asia. Terraced agriculture is being practised in these locations for a long time. The last case (Gono Chetona) falls in the Brahmaputra–Jamuna charlands which are the most ecologically vulnerable regions in the sub-continent with constantly changing landscape. Agriculture and livestock rearing are the main occupations, and there is substantial seasonal emigration for wage labour by the adult males. River erosion and floods force the people to adopt a semi-migratory lifestyle. The book attempts to analyse the potential as well as limitations of NGOdriven CBNRM endeavours across agroclimatic regions of South Asia with emphasis on four intrinsically linked normative concerns, namely sustainability, livelihood enhancement, equity and demographic decentralization in chapters 2–7. Comparative analysis of these case studies done in chapter 8, highlights the issues that require further research while portraying the strengths and limits of NGO-driven CBNRM. In Hivre Bazar, the post-watershed intervention scenario is such that farmers often grow three crops in a year – kharif bajra, rabi jowar and summer vegetable crops. Productivity has increased in the dry lands due to improvement in soil moisture levels. The revival of johads in Gopalpura has led to the proliferation of wheat and increased productivity. In Lingmuteychhu, productivity gains have also arisen, but more due to the introduction of both local and high-yielding, new varieties as opposed to increased water availability. In the case of Gono Chetona, improvements have come due to diversification of agriculture; for example, the promotion of vegetable gardens. CBNRM interventions in most cases have also led to new avenues of employment and income generation. The synthesis shows that CBNRM efforts have made significant contributions to livelihood enhancement and only limited gains in terms of collective action for sustainable and equitable access to benefits and continuing resource use, and in terms of democratic decentralization, contrary to the objectives of the programme. Livelihood benefits include improvements in availability of livelihood support resources (fuelwood, fodder, drinking water), increased productivity (including diversification of cropping pattern) in agriculture and allied activities, and new sources of livelihood. However, NGO-driven CBNRM has not met its goal of providing ‘alternative’ forms of ‘development’ due to impediments of state policy, short-sighted vision of implementers and confrontation with the socio-ecological reality of the region, which almost always are that of fragmented communities (or communities in flux) with unequal dependence and access to land and other natural resources along with great gender imbalances. Appalling, however, is the general absence of recognition of the importance of and the will to explore practical ways to bring about equitable resource transfer or benefit-sharing and the consequent innovations in this respect that are evident in the pioneering community initiatives such as pani panchayat, etc. Pertaining to the gains on the ecological sustainability front, Hivre Bazar and Thalisain initiatives through active participation of villagers have made significant regeneration of the water table within the village, and mechanisms such as ban on number of bore wells, the regulation of cropping pattern, restrictions on felling of trees and free grazing to ensure that in the future, the groundwater is neither over-exploited nor its recharge capability impaired. Nevertheless, the longterm sustainability of the interventions in the case of Ghoga and Gopalpura initiatives as the focus has been mostly on regeneration of resources, and less on regulating the use of regenerated resources. Further, in Lingmuteychhu and Gono Chetona, the interventions are mainly household-based and the focus has been less explicit on ecological components. The studies demonstrate the livelihood benefits to all of the interventions and significant variation in achievements with reference to sustainability, equity and democratic decentralization depending on the level and extent of community participation apart from the vision of implementers, strategy (or nature of intervention shaped by the question of community formation), the centrality of community formation and also the State policy. Case studies show that the influence of State policy is multi-faceted and often contradictory in nature. This necessitates NGOs to engage with the State in a much more purposeful way than in an ‘autonomous space’. Thus the role of NGOs in CBNRM is complementary, wherein they provide innovative experiments that the State can learn. This helps in achieving the goals of CBNRM through democratic decentralization. The book addresses the vital issues related to natural resource management and interests of the community. Key topics discussed throughout the book are still at the centre of the current debate. This compilation consists of well-written chapters based on rigorous synthesis of CBNRM case studies, which will serve as good references for students, researchers and practitioners in the years to come.

Lizard Wears Shades. A Spectacled Sphenomorphus (Squamata: Scincidae), from the Sacred Forests of Mawphlang, Meghalaya, North-east India

A new species of lygosomatine scincid lizard is described from the sacred forests of Mawphlang, in Meghalaya, northeastern India. Sphenomorphus apalpebratus sp. nov. possesses a spectacle or brille, an unusual feature within the Scincidae, and a first for the paraphyletic genus Sphenomorphus. The new species is compared with other members of the genus to which it is here assigned, as well as to members of the lygosomatine genera Lipinia and Scincella from mainland India, the Andaman and Nicobar Islands, and south-east Asia, to which it also bears resemblance. The new taxon is diagnosable in exhibiting the following combination of characters: small body size (SVL to 42.0 mm); moveable eyelids absent; auricular opening scaleless, situated in a shallow depression; dorsal scales show a line of demarcation along posterior edge of ventral pes; midbody scale rows 27-28; longitudinal scale rows between parietals and base of tail 62-64; lamellae under toe IV 8-9; supraoculars five; supralabials 5-6; infralabials 4-5; subcaudals 92; and dorsum golden brown, except at dorsal margin of lateral line, which is lighter, with four faintly spotted lines, two along each side of vertebral row of scales, that extend to tail base. The new species differs from its congeners in the lack of moveable eyelids, a character shared with several distantly related scincid genera.

Systematic status of Fejervarya ((Amphibia, Anura, Dicroglossidae) from South and SE Asia with the description of a new species from the Western Ghats of Peninsular India

We carried out a large-scale phylogenetic analysis of fejervaryan (dicroglossid frogs with `Fejervaryan lines' on the ventral side of the body) frogs, distributed in South and SE Asia, using published and newly generated sequences of unidentified individuals from the northern Western Ghats. The results corroborate the presence of a larger fejervaryan clade with a sister relationship to a clade composed of Sphaerotheca. Two sister clades could be discerned within the lager fejervaryan clade. The unidentified individuals formed a monophyletic group and showed a strong support for a sister relationship with Minervarya sahyadris. The species was found to be highly divergent (16S rRNA-4% and tyr-1%) from its sister lineage Minervarya sahyadris, and the clade composed of these two lineages were found to be deeply nested within the larger clade of Fejervarya. Based on this, the genus Minervarya Dubois, Ohler and Biju, 2001 is synonymized under the genus Fejervarya Bolkay, 1915. The unidentified lineage is recognized, based on phylogenetic position, genetic divergence and morphological divergence, as a distinct species and named here as Fejervarya gomantaki sp. nov. The presence of rictal glands was observed to be a synapomorphic character shared by the nested clade members, Fejervarya sahyadris and Fejervarya gomantaki sp. nov. Based on the presence of rictal gland and small size, Minervarya chilapata, a species from a lowland region in the Eastern Himalayas, is synonymized under Fejervarya and evidence for morphological separation from the new species, Fejervarya gomantaki sp. nov. is provided. For the fejervaryan frogs, currently three generic names (Frost, 2015) are available for the two phylogenetic subclades; the genus Fejervarya Bolkay, 1915 for the species of fejervaryan frogs having distribution in the South East Asia; the genus Zakerana Howlader, 2011 for the species of fejervaryan frogs having distribution in the South Asia and the genus Minervarya Dubois, Ohler and Biju, 2001 nested within the `Zakerana clade'. In the phylogenetic analysis Minervarya sahyadris, the new species described herein as Fejervarya gomantaki sp. nov. are nested within the `Zakerana clade', if the `Zakerana clade' for the fejervaryan frogs having distribution in the South Asia is provided a generic status the nomen `Minervarya' should be considered as per the principle of priority of the ICZN Code. Taking into consideration the overlapping distribution ranges of members of the sister clades within the larger fejervaryan clade and the absence of distinct morphological characteristics, we also synonymize the genus Zakerana Howlader, 2011, a name assigned to one of the sister clades with members predominantly distributed in South Asia, under the genus Fejervarya Bolkay, 1915. We discuss the need for additional sampling to identify additional taxa and determine the geographical ranges of the members of the sister clades within Fejervarya to resolve taxonomy within this group.

Y-chromosome evidence of southern origin of the East Asian - Specific haplogroup O3-M122

The prehistoric peopling of East Asia by modern humans remains controversial with respect to early population migrations. Here, we present a systematic sampling and genetic screening of an East Asian-specific Y-chromosome haplogroup (O3-M122) in 2,332 individuals from diverse East Asian populations. Our results indicate that the O3-M122 lineage is dominant in East Asian populations, with an average frequency of 44.3%. The microsatellite data show that the O3-M122 haplotypes in southern East Asia are more diverse than those in northern East Asia, suggesting a southern origin of the O3-M122 mutation. It was estimated that the early northward migration of the O3-M122 lineages in East Asia occurred similar to 25,000-30,000 years ago, consistent with the fossil records of modern humans in East Asia.

The ADH1B Arg47His polymorphism in East Asian populations and expansion of rice domestication in history

Background: The emergence of agriculture about 10,000 years ago marks a dramatic change in human evolutionary history. The diet shift in agriculture societies might have a great impact on the genetic makeup of Neolithic human populations. The regionally restricted enrichment of the class I alcohol dehydrogenase sequence polymorphism (ADH1BArg47His) in southern China and the adjacent areas suggests Darwinian positive selection on this genetic locus during Neolithic time though the driving force is yet to be disclosed. Results: We studied a total of 38 populations (2,275 individuals) including Han Chinese, Tibetan and other ethnic populations across China. The geographic distribution of the ADH1B*47His allele in these populations indicates a clear east-to-west cline, and it is dominant in south-eastern populations but rare in Tibetan populations. The molecular dating suggests that the emergence of the ADH1B*47His allele occurred about 10,000 similar to 7,000 years ago. Conclusion: We present genetic evidence of selection on the ADH1BArg47His polymorphism caused by the emergence and expansion of rice domestication in East Asia. The geographic distribution of the ADH1B*47His allele in East Asia is consistent with the unearthed culture relic sites of rice domestication in China. The estimated origin time of ADH1B*47His allele in those populations coincides with the time of origin and expansion of Neolithic agriculture in southern China.

Phylogeny of East Asian mitochondrial DNA lineages inferred from complete sequences

The now-emerging mitochondrial DNA ( mtDNA) population genomics provides information for reconstructing a well-resolved mtDNA phylogeny and for discerning the phylogenetic status of the subcontinentally specific haplogroups. Although several major East Asian mtDNA haplogroups have been identified in studies elsewhere, some of the most basal haplogroups, as well as numerous minor subhaplogroups, were not yet determined or fully characterized. To fill the lacunae, we selected 48 mtDNAs from >2,000 samples across China for complete sequencing that cover virtually all ( sub) haplogroups discernible to date in East Asia. This East Asian mtDNA phylogeny can henceforth serve as a solid basis for phylogeographic analyses of mtDNAs, as well as for studies of mitochondrial diseases in East and Southeast Asia.

Multiple invasions of europe by east asian cobitid loaches (Teleostei : Cobitidae)

While it has been widely suggested that freshwater fishes from East Asia invaded the western Palaearctic, details about this process are largely unknown. Here, using the cytochrome b gene, we evaluated the phylogenetic relationships of a small group of Eurasian primary freshwater fishes (Cobitidae), which are widely distributed and species rich in East Asia and Europe, with the purpose of inferring their invasion process of Europe from East Asia. Though phylogenetic relationships of cobitids were not well resolved, our analysis could identify three sister groups formed by the European and East Asian cobitids, which brought new insights into the biogeography of the genera Cobitis, Misgurnus, and Sabanejewia. The present results support the view that Asian cobitid fishes may have invaded Europe at least five times independently, and once reverse colonization of European cobitids to East Asia could also be found. Ancestral Sabanejewia might have been the first cobitids to cross Siberia and invade the EMZS (Euro-Mediterranean zoogeographic subregion) about 33.54 million years ago (MYA). One lineage of Cobitis and the ancestor of Misgurnus fossilis (Linnaeus) almost in the same time invaded the Europe, responding to 16.71 MYA and 16.59 MYA, respectively. Three different lineages of Cobitis were found to have invaded the EMZS from East Asia, and once reverse invasion to East Asia occurred to one subclade of European Cobitis. And our data also suggest that the diversity of East Asian cobitid fishes, especially of the genus Cobitis, is greatly underestimated.

Molecular evidence for the monophyly of East Asian groups of Cyprinidae (Teleostei : Cypriniformes) derived from the nuclear recombination activating gene 2 sequences

The family Cyprinidae is one of the largest families of fishes in the world and a well-known component of the East Asian freshwater fish fauna. However, the phylogenetic relationships among cyprinids are still poorly understood despite much effort paid on the cyprinid molecular phylogenetics. Original nucleotide sequence data of the nuclear recombination activating gene 2 were collected from 109 cyprinid species and four non-cyprinid cypriniform outgroup taxa and used to infer the cyprinid phylogenetic relationships and to estimate node divergence times. Phylogenetic reconstructions using maximum parsimony, maximum likelihood, and Bayesian analysis retrieved the same clades, only branching order within these clades varied slightly between trees. Although the morphological diversity is remarkable, the endemic cyprinid taxa in East Asia emerged as a monophyletic clade referred to as Xenocypridini. The monophyly for the subfamilies including Cyprininae and Leuciscinae, as well as the tribes including Labeonini, Gobionini, Acheilognathini, and Leuciscini, was also well resolved with high nodal support. Analysis of the RAG2 gene supported the following cyprinid molecular phylogeny: the Danioninae is the most basal subfamily within the family Cyprinidae and the Cyprininae is the sister group of the Leuciscinae. The divergence times were estimated for the nodes corresponding to the principal clades within the Cyprinidae. The family Cyprinidae appears to have originated in the mid-Eocene in Asia, with the cladogenic event of the key basal group Danioninae occurring in the early Oligocene (about 31-30 MYA), and the origins of the two subfamilies, Cyprininae and Leuciscinae, occurring in the mid-Oligocene (around 26 MYA). (c) 2006 Elsevier Inc. All rights reserved.

Records of the East Asian winter monsoon from the mud area on the inner shelf of the East China Sea since the mid-Holocene

AMS(14)C dating and analysis of grain size, major elements and clay minerals were applied to Core MZ01 from the mud area on the inner shelf of the East China Sea. Based on the environmentally sensitive grain size, clay mineral and major element assemblages, the history of the East Asia winter monsoon since the mid-Holocene could be reconstructed. These three proxies, mean grain size (>9.71 mu m), chemical index of alteration (CIA) and ratio of smectite to kaolinite in particular, show similar fluctuation patterns. Furthermore, 10 extreme values corresponding to the contemporary cooling events could be recognized since the mid-Holocene; these extreme values are likely to have been caused by the strengthening of the East Asia winter monsoon. The cooling events correlated well with the results of the delta O-18 curves of the Dunde ice core and GISP2, which therefore revealed a regional response to global climate change. Four stages of the East Asia winter monsoon were identified, i.e. 8300-6300 a BP, strong and unstable; 6300-3800 a BP, strong but stable; 3800-1400 a BP, weak and unstable; after 1400 a BP, weak but stable.