Improving species status assessments under the U.S. Endangered Species Act and implications for multispecies conservation challenges worldwide.

Despite its successes, the U.S. Endangered Species Act (ESA) has proven challenging to implement due to funding limitations, workload backlog, and other problems. As threats to species survival intensify and as more species come under threat, the need for the ESA and similar conservation laws and policies in other countries to function efficiently has grown. Attempts by the U.S. Fish and Wildlife Service (USFWS) to streamline ESA decisions include multispecies recovery plans and habitat conservation plans. We address species status assessment (SSA), a USFWS process to inform ESA decisions from listing to recovery, within the context of multispecies and ecosystem planning. Although existing SSAs have a single-species focus, ecosystem-based research can efficiently inform multiple SSAs within a region and provide a foundation for transition to multispecies SSAs in the future. We considered at-risk grassland species and ecosystems within the southeastern United States, where a disproportionate number of rare and endemic species are associated with grasslands. To initiate our ecosystem-based approach, we used a combined literature-based and structured World Café workshop format to identify science needs for SSAs. Discussions concentrated on 5 categories of threats to grassland species and ecosystems, consistent with recommendations to make shared threats a focus of planning under the ESA: (1) habitat loss, fragmentation, and disruption of functional connectivity; (2) climate change; (3) altered disturbance regimes; (4) invasive species; and (5) localized impacts. For each threat, workshop participants identified science and information needs, including database availability, research priorities, and modeling and mapping needs. Grouping species by habitat and shared threats can make the SSA process and other planning processes for conservation of at-risk species worldwide more efficient and useful. We found a combination of literature review and structured discussion effective for identifying the scientific information and analysis needed to support the development of multiple SSAs. Article impact statement: Species status assessments can be improved by an ecosystem-based approach that groups imperiled species by shared habitats and threats.

Mejoramiento de la Evaluación del Estado de una Especie bajo el Acta de Especies en Peligro de los Estados Unidos y Sus Consecuencias para los Retos de la Conservación Multiespecie a Nivel Mundial Resumen A pesar de su éxito, el Acta de Especies en Peligro de los E.U.A. (AEP) ha sido un reto de implementación por las limitaciones en su financiamiento, el retraso en la carga de trabajo y otros problemas. Conforme se intensifican las amenazas a la supervivencia de las especies y más especies resultan amenazadas, aumenta la necesidad de que la AEP y las políticas similares de otros países funcionen efectivamente. Los intentos por parte del Servicio Estadounidense de Pesca y Fauna (SEPF) para optimizar las decisiones de la AEP incluyen planes multiespecie de recuperación y planes de conservación de hábitat (PRH). Abordamos la evaluación del estado de las especies (EEE), un proceso del SEPF para orientar las decisiones del AEP desde el listado hasta la recuperación, dentro del contexto de la planeación multiespecie y de ecosistemas. Aunque las EEE existentes tienen un enfoque sobre una única especie, la investigación basada en el ecosistema puede orientar eficientemente a múltiples EEE dentro de una región y proporcionar una base para la transición a las EEE multiespecie en el futuro. Consideramos a las especies y los ecosistemas en riesgo de los pastizales del sureste de los Estados Unidos, en donde un número desproporcionado de especies raras y endémicas está asociado con los pastizales. Para iniciar nuestra estrategia basada en el ecosistema, usamos un formato de taller de World Café estructurado y basado en la literatura para identificar la necesidad de tener EEE. Las discusiones se centraron en cinco categorías de amenazas para las especies y ecosistemas de los pastizales, consistentes con las recomendaciones para volver a las amenazas compartidas un foco de la planeación bajo la AEP: (1) pérdida del hábitat, fragmentación y disrupción de la conectividad funcional; (2) cambio climático; (3) regímenes alterados de perturbación; (4) especies invasoras; y (5) impactos localizados. Para cada amenaza, los participantes del taller identificaron las necesidades científicas y de información, incluyendo la disponibilidad de bases de datos, prioridades de la investigación y necesidades de modelado y mapeado. La agrupación de las especies por hábitat y amenaza compartida puede hacer más eficientes y útiles el proceso de EEE y otros procesos de planeación de la conservación de especies en riesgo a nivel mundial. Encontramos una combinación de revisiones bibliográficas y discusiones estructuradas para identificar la información y el análisis necesarios para respaldar el desarrollo de múltiples EEE.

Seed Biology of Lepidium apetalum (Brassicaceae), with Particular Reference to Dormancy and Mucilage Development.

Lepidium apetalum (Brassicaceae) is an annual or biennial weed widely distributed in Asia and Europe. The outer surface of L. apetalum seeds produces a large amount of mucilage. The primary aim of this study was to explore the dormancy characteristics and to determine how mucilage develops. The role of mucilage in water absorption/dehydration, the effects of after-ripening, gibberellin acid (GA3), cold stratification and seed coat scarification on germination, the role of mucilage in germination and seedling growth during drought, and the progress of mucilage production during seed development were investigated. The results indicate that the best temperature regime for germination was 10/20 °C. After-ripening, GA3 and seed coat scarification helped to break dormancy. Light promoted germination. Seedling growth of mucilaged seeds were significantly higher than those of demucilaged seeds at -0.606 and -1.027 MPa. Anatomical changes during seed development showed that mucilage was derived from the outer layer of the outer integument cells. Our findings suggest that seeds of L. apetalum exhibited non-deep physiological dormancy. The dormancy characteristics along with mucilage production give seeds of L. apetalum a competitive advantage over other species, and thus contribute to its potential as a weed. Effective control of this weed can be achieved by deep tillage.

A research agenda for seed-trait functional ecology.

Trait-based approaches have improved our understanding of plant evolution, community assembly and ecosystem functioning. A major challenge for the upcoming decades is to understand the functions and evolution of early life-history traits, across levels of organization and ecological strategies. Although a variety of seed traits are critical for dispersal, persistence, germination timing and seedling establishment, only seed mass has been considered systematically. Here we suggest broadening the range of morphological, physiological and biochemical seed traits to add new understanding on plant niches, population dynamics and community assembly. The diversity of seed traits and functions provides an important challenge that will require international collaboration in three areas of research. First, we present a conceptual framework for a seed ecological spectrum that builds upon current understanding of plant niches. We then lay the foundation for a seed-trait functional network, the establishment of which will underpin and facilitate trait-based inferences. Finally, we anticipate novel insights and challenges associated with incorporating diverse seed traits into predictive evolutionary ecology, community ecology and applied ecology. If the community invests in standardized seed-trait collection and the implementation of rigorous databases, major strides can be made at this exciting frontier of functional ecology.

Grazing disturbance increases transient but decreases persistent soil seed bank.

Very few studies have examined whether the impacts of grazing disturbance on soil seed banks occur directly or indirectly through aboveground vegetation and soil properties. The potential role of the seed bank in alpine wetland restoration is also unknown. We used SEM (structural equation modeling) to explore the direct effect of grazing disturbance on the seed bank and the indirect effect through aboveground vegetation and soil properties. We also studied the role of the seed bank on the restoration potential in wetlands with various grazing intensities: low (fenced, winter grazed only), medium (seasonally grazed), and high (whole-year grazed). For the seed bank, species richness and density per plot showed no difference among grazing intensities for each depth (0-5, 5-10, 10-15 cm) and for the whole depth (0-15 cm) in spring and summer. There was no direct effect of grazing disturbance on seed bank richness and density both in spring and summer, and also no indirect effect on the seed bank through its direct effect on vegetation richness and abundance. Grazing disturbance indirectly increased spring seed bank density but decreased summer seed bank density through its direct effect (negative correlation) on soil moisture and total nitrogen and its indirect effect on vegetation abundance. Species composition of the vegetation changed with grazing regime, but that of the seed bank did not. An increased trend of similarity between the seed bank and aboveground vegetation with increased grazing disturbance was found in the shallow depth and in the whole depth only in spring. Although there was almost no change in seed bank size with grazing intensities, grazing disturbance increased the quantity of transient seeds but decreased persistent seeds. Persistent seeds stored in the soil could play a crucial role in vegetation regeneration and in restoration of degraded wetland ecosystems. The seed bank should be an integral part of alpine wetland restoration programs.

Aerial and soil seed banks enable populations of an annual species to cope with an unpredictable dune ecosystem.

BACKGROUND AND AIMS: Simultaneous formation of aerial and soil seed banks by a species provides a mechanism for population maintenance in unpredictable environments. Eolian activity greatly affects growth and regeneration of plants in a sand dune system, but we know little about the difference in the contributions of these two seed banks to population dynamics in sand dunes. METHODS: Seed release, germination, seedling emergence and survival of a desert annual, Agriophyllum squarrosum (Chenopodiaceae), inhabiting the Ordos Sandland in China, were determined in order to explore the different functions of the aerial and soil seed banks. KEY RESULTS: The size of the aerial seed bank was higher than that of the soil seed bank throughout the growing season. Seed release was positively related to wind velocity. Compared with the soil seed bank, seed germination from the aerial seed bank was lower at low temperature (5/15 °C night/day) but higher in the light. Seedling emergence from the soil seed bank was earlier than that from the aerial seed bank. Early-emerged (15 April-15 May) seedlings died due to frost, but seedlings that emerged during the following months survived to reproduce successfully. CONCLUSIONS: The timing of seed release and different germination behaviour resulted in a temporal heterogeneity of seedling emergence and establishment between the two seed banks. The study suggests that a bet-hedging strategy for the two seed banks enables A. squarrosum populations to cope successfully with the unpredictable desert environment.

Seed germination and seedling development ecology in world-wide populations of a circumboreal Tertiary relict.

BACKGROUND AND AIMS: Temperate forests are disjunct in the Northern Hemisphere, having become fragmented from the earlier widespread (Tertiary) boreotropical forest. We asked 'What are the contemporary patterns of population variation in ecological traits of a Tertiary relict in a macroecological context?'. This issue underpins our understanding of variation in populations occurring in the same biome but on different continents. METHODOLOGY: We examined characters associated with root and shoot emergences among populations of Viburnum opulus in temperate forests of Asia, North America and Europe. This species has complex seedling emergence extending over several years and requiring various temperature cues. PRINCIPAL RESULTS: Populations varied in germination responses and clustered into groups that were only partly related to varietal status. Whereas roots (at warm temperatures) and shoots (following a cold period) simultaneously emerged from seeds of all populations when simulated dispersal occurred in winter, they were delayed in some populations when dispersal occurred in summer. CONCLUSIONS: Viburnum opulus populations, some separated by 10 300 km, showed high similarity in seedling development and in germination phenology, and we suggest that stabilizing selection has played a key role in maintaining similar dormancy mechanisms. Nevertheless, there was some degree of variation in other germination characters, suggesting local adaptation.

Sympatric species of Hibbertia (Dilleniaceae) vary in dormancy break and germination requirements: implications for classifying morphophysiological dormancy in Mediterranean biomes.

BACKGROUND AND AIMS: Several ecologically important plant families in Mediterranean biomes have seeds with morphophysiological dormancy (MPD) but have been poorly studied. The aim of this study was to understand the seed ecology of these species by focusing on the prominent, yet intractably dormant Australian genus Hibbertia. It was hypothesized that the slow germination in species of this genus is caused by a requirement for embryo growth inside the seed before germination, and that initiation of embryo growth is reliant upon a complex sequence of environmental cues including seasonal fluctuations in temperature and moisture, and an interplay with light and smoke. Using the results, the classification of the MPD level in species of Hibbertia is considered. METHODS: Four species of Hibbertia in winter rainfall south-western Australia were selected. These species, whilst differing in geographic distributions, are variously sympatric, and all are important understorey components of plant communities. The following aspects related to dormancy break, embryo growth and germination were investigated: temperature and moisture requirements; effects of karrikinolide, gibberellic acid and aerosol smoke; and phenology. KEY RESULTS: Following exposure to wet/dry cycles at low or high temperatures, embryo growth and germination occurred, albeit slowly in all species at low temperatures when moisture was unlimited, corresponding to winter in south-west Australia. Photo regime influenced germination only in H. racemosa. Aerosol smoke triggered substantial germination during the 1st germination season in H. huegelii and H. hypericoides. CONCLUSIONS: Although the study species are con-generic, sympatric and produce seeds of identical morphology, they possessed different dormancy-break and germination requirements. The physiological component of MPD was non-deep in H. racemosa but varied in the other three species where more deeply dormant seeds required >1 summer to overcome dormancy and, thus, germination was spread over time. Embryos grew during winter, but future studies need to resolve the role of cold versus warm stratification by using constant temperature regimes. To include Mediterranean species with MPD, some modifications to the current seed-dormancy classification system may need consideration: (a) wet/dry conditions for warm stratification and (b) a relatively long period for warm stratification. These outcomes have important implications for improving experimental approaches to resolve the effective use of broadcast seed for ecological restoration.

Seed dormancy in Trillium camschatcense (Melanthiaceae) and the possible roles of light and temperature requirements for seed germination in forests.

PREMISE OF THE STUDY: Seeds of some temperate plants require multiple treatments to break complex forms of dormancy, such as deep simple double morphophysiological dormancy (MPD), but embryo growth and radicle and cotyledon emergence have not been studied in depth for this level of MPD. Here we studied Trillium camschatcense, a species that is purported to have this MPD and that is sensitive to habitat fragmentation with decreased recruitment at forest edges. METHODS: Trillium camschatcense seeds were collected and experiments conducted in Hokkaido, Japan over 5 years. Growth of embryos and emergence of radicles and cotyledons were tracked in relation to field temperature to determine specific temperature and light requirements for these stages under laboratory conditions. KEY RESULTS: Low (winter) temperatures overcame dormancy in the underdeveloped embryos, and embryo growth and radicle emergence occurred between July and September, ca. 1 year after seed dispersal. Radicles emerged optimally over a narrow temperature range (20-25°C), in darkness, and at constant temperatures. Roots developed during the second autumn. Cotyledons needed a second low temperature (second winter) to emerge from seeds with roots, doing so in April, slightly over 1.5 years after dispersal. CONCLUSIONS: Seeds of T. camschatcense have deep simple double MPD and requirements for radicle emergence: darkness and constant temperatures. Ecologically, edges of forests may be deleterious for germination of the species due to increased light and to higher temperature fluctuations as compared to the interiors of forests. Thus, these specific requirements may play an important role in reducing seed germination of this plant at forest edges.

Understanding the germination of bulbils from an ecological perspective: a case study on Chinese yam (Dioscorea polystachya).

BACKGROUND AND AIMS: Bulbils serve as a means of vegetative reproduction and of dispersal for many plants; this latter aspect making them analogous to seeds. However, germination of bulbils may differ considerably from seeds due to dissimilar anatomical structures and perhaps environmental cue perception. The few laboratory studies done on bulbils suggest that their germination is similar to that of seeds in the same habitats and to vegetative buds of winter-dormant plants. The present study is the first to examine how bulbil germination is controlled in nature in relation to dispersal (before vs. after winter of the same cohort) and to ambient temperatures. METHODS: Under laboratory conditions, temperature and light requirements for root and shoot emergences from bulbils of Dioscorea polystachya collected in September, 2005, February, 2006 (produced in 2005) and July, 2006 were determined. Effects of cold stratification and dry storage for releasing dormancy were tested on September and July bulbils. The phenology of dormancy release and of root and shoot emergences and the persistence of bulbils in soil were followed over time under field conditions. KEY RESULTS: Although a low percentage of bulbils collected in July or in September produced roots, but no shoots, in the laboratory and field, these roots died within approx. 1 month. Regardless of collection date, cold stratification markedly increased root and shoot emergences. Bulbils sown outdoors in October produced roots and shoots the following March and April, respectively. The soil bulbil bank is short lived. CONCLUSIONS: Bulbils of D. polystachya are similar to seeds of many temperate plants being mostly dormant when dispersed in summer or autumn and overcoming dormancy with cold stratification during winter. Adaptively, bulbil germination primarily occurs in spring at the beginning of a favourable period for survivorship and growth.

Agroecosystem management for rare species of Paysonia (Brassicaceae): integrating their seed ecology and life cycle with cropping regimens in a changing climate.

Dormancy break and germination of seeds are governed by climatic cues, and predicted changes in climate may impact the ecology and conservation of species. Paysonia perforata and P. stonensis are rare brassicaceous winter annuals occurring primarily in fields on floodplains, where corn or soybeans are recommended for habitat maintenance. We tested the effects of precipitation, based on two predictions of changes in climate, on seed germination in these two species and placed the results into a management framework. Seeds of both species, collected during peak dispersal in late April/early May, were given various periods of light (or darkness) followed by darkness (or light) at summer temperatures before placement in darkness during late summer/early autumn in both laboratory and field. The light requirement was met earliest at 10 wk (mid-July) on alternating wet/dry substrate (simulating current climatic conditions). However, seeds of P. perforata and P. stonensis were photostimulated earliest at 2 wk (mid-May) and 6 wk (mid-June), respectively, on a continuously moist substrate (simulating predicted future conditions). The soil seed bank could be depleted if plowing coincides with photostimulation of seeds. Fields should be prepared after dispersal but before seeds are photostimulated and harvesting completed before seed germination in early September. Because seeds are highly photostimulated in late summer, disturbance from harvesting must be low to prevent burial. Cultivation of soybean, particularly for forage, is better matched to the seed biology and life cycle of Paysonia than that of corn under current and predicted climates.

Complex combination of seed dormancy and seedling development determine emergence of Viburnum tinus (Caprifoliaceae).

BACKGROUND AND AIMS: The shrub Viburnum tinus is widely distributed in mattoral vegetation of the Mediterranean basin. The purpose of the present study was to classify the seed dormancy type and examine the requirements for embryo growth, root protrusion and shoot emergence. METHODS: Overwintered fruits were collected in western Spain in April 2001 and prepared in three ways: entire pericarp was removed, exocarp and mesocarp were removed or fruits were left intact. Fruits treated in these three ways were subjected to artificial annual temperature cycles or to constant temperature regimes for 1.5 years. KEY RESULTS: Removal of exocarp and mesocarp was necessary for embryo growth and germination. High temperature favoured dormancy alleviation and embryo growth, intermediate to low temperatures favoured root protrusion, and intermediate temperature shoot emergence. There was substantial germination at constant temperature regimes, indicating an overlap between temperature intervals suitable for the different stages of embryo and seedling development. Functionally, V. tinus has the same root and shoot emergence pattern that is described for other Viburnum species considered to have epicotyl dormancy. However, the requirement for high and low temperatures for radicle protrusion and epicotyl emergence, respectively, was missing in V. tinus; these characters are the foundation for the epicotyl dormancy classification. CONCLUSIONS: It is concluded that V. tinus does not have epicotyl dormancy. Instead, there is a combination of a weak morphophysiological dormancy and a slow germination process, where different temperatures during an annual cycle favour different development stages. The present study suggests that the first complete seedlings would emerge in the field 1.5 years after fruit maturation in October, i.e. seed dispersal during winter, embryo growth during the first summer, root protrusion and establishment during the second autumn and winter, and cotyledon emergence during the second spring.

Seed germination ecophysiology of the Asian species Osmorhiza aristata (Apiaceae): comparison with its North American congeners and implications for evolution of types of dormancy.

Osmorhiza aristata is an herbaceous perennial that grows primarily in Japan, through southern China, to the Himalayas. It closely resembles the eastern North American species O. claytonii and O. longistylis, and, together, the three species are an example of the well-known North American-Asian pattern of disjunction. Requirements for dormancy break and embryo growth were determined for seeds of O. aristata collected in Japan during the summers of 1998-2000. Embryos in fresh seeds were ca. 0.5 mm long, and they had to grow to 9 mm before the radicle emerged from the mericarp. Embryo growth and germination occurred during cold stratification at 5°C, the optimum temperature for germination. Gibberellic acid did not substitute for cold stratification. Thus, O. aristata seeds have deep complex morphophysiological dormancy (MPD). The type of MPD in O. aristata is similar to that in two western North American congeners but different from that in eastern North American congeners (nondeep complex MPD). Mapping the types of MPD onto a phylogeny of the genus suggests that nondeep complex MPD is derived from deep complex MPD. Although eastern North American-Asian disjuncts often exhibit morphological stasis, the taxa may differ greatly in physiological traits, such as seed dormancy.