Colin J. Yates

Seed supply for broadscale restoration: maximizing evolutionary potential.

Restoring degraded land to combat environmental degradation requires the collection of vast quantities of germplasm (seed). Sourcing this material raises questions related to provenance selection, seed quality and harvest sustainability. Restoration guidelines strongly recommend using local sources to maximize local adaptation and prevent outbreeding depression, but in highly modified landscapes this restricts collection to small remnants where limited, poor quality seed is available, and where harvesting impacts may be high. We review three principles guiding the sourcing of restoration germplasm: (i) the appropriateness of using ‘local’ seed, (ii) sample sizes and population characteristics required to capture sufficient genetic diversity to establish self‐sustaining populations and (iii) the impact of over‐harvesting source populations. We review these topics by examining current collection guidelines and the evidence supporting these, then we consider if the guidelines can be improved and the consequences of not doing so. We find that the emphasis on local seed sourcing will, in many cases, lead to poor restoration outcomes, particularly at broad geographic scales. We suggest that seed sourcing should concentrate less on local collection and more on capturing high quality and genetically diverse seed to maximize the adaptive potential of restoration efforts to current and future environmental change.

Impacts of ecosystem fragmentation on plant populations: generalising the idiosyncratic

Fragmentation of natural vegetation is one of the most pervasive changes in terrestrial ecosystems across the Earth. Developing a general understanding of how fragmentation affects plant and animal populations is essential to meet the pressing need for guidelines for the management of fragmented systems. Nevertheless, this general understanding has to take account of differences in ecosystem types and different biogeographic, evolutionary and ecological backgrounds against which fragmentation impacts are played out in different parts of the world. Here, we examine fragmentation impacts on plant populations by considering the processes underlying fragmentation. We suggest that it is critical to focus on the key processes that are important in particular situations, rather than assuming that the same factors are likely to be important everywhere. In other words, there are inevitable limits to generalisation because of the idiosyncratic nature of the geography, history and biota of different regions. Studies on the effects of fragmentation on plant populations have focused on a limited subset of plant types and have concentrated heavily on reproductive output rather than actual regeneration success. These studies have indicated a clear impact of fragmentation on fecundity, but there is no clear signal in terms of the actual importance of this in relation to population viability. Other factors including local habitat conditions, disturbance and competition from weeds may be just as important as the classical biogeographical impacts of fragmentation. Generalisations based on a clear assessment of key life-history processes may be valuable tools in developing management responses to ecosystem fragmentation, but this requires considerably more emphasis on factors affecting successful recruitment as well as factors affecting fecundity.

Extensive pollen dispersal in a bird-pollinated shrub, Calothamnus quadrifidus, in a fragmented landscape

Pollen dispersal was investigated in six populations of Calothamnus quadrifidus, a bird‐pollinated shrub in the fragmented agricultural region of southern Western Australia. Paternity analysis using six microsatellite loci identified a pollen source within populations for 67% of seedlings, and the remainder were assumed to have arisen from pollen sources outside the populations. Outcrossing was variable, ranging from 5% to 82%, and long‐distance pollen dispersal was observed in all populations with up to 43% of pollen sourced from outside the populations over distances of up to 5 km. This extensive pollen immigration was positively associated with population size but not isolation. Comparison of two populations of similar size but different density showed greater internal pollination and less selfing in the denser population, suggesting an influence of density on pollinator behaviour. The study revealed extensive long‐distance pollen dispersal for C. quadrifidus within this fragmented agricultural landscape and highlighted the interaction between reserve populations and isolated road verge remnants in maintaining genetic connectivity at the landscape scale.

Plant mating systems and assessing population persistence in fragmented landscapes

Population size and habitat disturbance are key factors likely to shape the mating system of populations in disturbed and fragmented landscapes. They would be expected to influence the availability and behaviour of the pollinator, the ability to find mates in self-incompatible species, inbreeding in self-compatible species and the size of the pollen pool. These in turn might be expected to influence key variables critical for population persistence such as seed production, seed germination and seedling fitness. Here we investigate mating-system variation in six rare species, i.e. Banksia cuneata, B. oligantha, Lambertia orbifolia (Proteaceae), Verticordia fimbrilepis subsp. fimbrilepis, Eucalyptus rameliana (Myrtaceae), Acacia sciophanes (Mimosaceae), and two common species, i.e. Calothamnus quadrifidus (Myrtaceae) and Acacia anfractuosa. All seven species are animal-pollinated relatively long-lived woody shrubs with mixed-mating systems. Population variation in mating-system parameters was investigated in relation to population size and habitat disturbance. We show that although the mating system will vary depending on pollination biology and life-history, as populations get smaller and habitat disturbance increases there is a trend towards increased inbreeding, smaller effective sizes of paternal pollen pools and greater variation in outcrossing among plants. From the species investigated in this study we have found that changes in the mating system can be useful indicators of population processes and can give valuable insight into the development of conservation strategies for the persistence of plant species following anthropogenic disturbance and landscape fragmentation.

Evaluating the influence of different aspects of habitat fragmentation on mating patterns and pollen dispersal in the bird-pollinated Banksia sphaerocarpa var. caesia

Habitat fragmentation can significantly affect mating and pollen dispersal patterns in plant populations, although the differential effects of the various aspects of fragmentation are poorly understood. In this study, we used eight microsatellite loci to investigate the effect of fragmentation on the mating system and pollen dispersal within one large and eight small population remnants of Banksia sphaerocarpa var. caesia, a bird‐pollinated shrub in the southern agricultural region of Western Australia. The large population had a much larger neighbourhood size and lower selfing rate, maternal pollen pool differentiation and within‐plot mean pollen dispersal distance than the small populations. Outcrossing was consistently high and ranged from 85.7% ± 2.6 to 98.5% ± 0.9, and mating patterns suggested nearest‐neighbour pollination. Pollen immigration into small populations ranged from 2.8% ± 1.8 to 16.5% ± 3.2. Using the small populations, we tested for correlations between various fragmentation variables and mating system and pollen dispersal parameters. We found significant negative linear relationships between population isolation and outcrossing rate; population shape and neighbourhood size; and conspecific density and mean pollen dispersal distance. There were significant positive linear relationships between population shape and pollen pool differentiation and between population size and number of different fathers per seed crop. Our results suggest that birds may use a series of fragmented populations as a vegetation corridor while foraging across the landscape and that population connectivity is a critical determinant of pollinator visitation. Our results also suggest that the effect of a linear population shape on the mating system and pollen dispersal is routinely underestimated.

Making decisions to conserve species under climate change

Severe impacts on biodiversity are predicted to arise from climate change. These impacts may not be adequately addressed by conventional approaches to conservation. As a result, additional management actions are now being considered. However, there is currently limited guidance to help decision makers choose which set of actions (and in what order) is most appropriate for species that are considered to be vulnerable. Here, we provide a decision framework for the full complement of actions aimed at conserving species under climate change from ongoing conservation in existing refugia through various forms of mobility enhancement to ex situ conservation outside the natural environment. We explicitly recognize that allocation of conservation resources toward particular actions may be governed by factors such as the likelihood of success, cost and likely co-benefits to non-target species in addition to perceived vulnerability of individual species. As such, we use expert judgment of probable tradeoffs in resource allocation to inform the sequential evaluation of proposed management interventions.

Assessing limitations on population growth in two critically endangered Acacia taxa

Abstract Flowering phenology, soil seedbank and the impact of fire, weeds and grazing were investigated in two rare Acacia taxa restricted to the agricultural district north of Perth Western Australia. Acacia aprica is known from six extant populations and one extinct population. Five of the extant populations are restricted to linear road reserves with the sixth population located in a small native vegetation remnant. Acacia cochlocarpa ssp . cochlocarpa is known from a single population also situated on a road verge. Size class structure, levels of canopy death and an absence of juveniles indicated that all populations are in decline. Flowering intensity and success varied between populations and years in A. aprica and between years in A. cochlocarpa ssp. cochlocarpa. Seed bank analysis indicated that seeds were patchily distributed while experimental fires demonstrated that such events could break seed dormancy and promote germination. Both taxa have similar germination physiologies and showed increased germination after seeds were exposed to heat. Following emergence, however, competition with annual weeds had a negative impact on seedling growth and survival in both taxa. While vertebrate grazing had some influence, weeds were the major inhibitory influence on recruitment. Reduced fire frequencies since fragmentation may be responsible for population decline in both taxa but other site specific factors such as weeds and grazing may affect the establishment of seedlings following fire.

Plant communities of the ironstone ranges of South Western Australia: hotspots for plant diversity and mineral deposits

Plant communities of ancient banded iron formation ranges of South Western Australia occur as islands in a matrix of the largest remaining area of Mediterranean woodlands and shrub lands on the planet. These xeric shrub lands are structurally and compositionally different from the surrounding matrix and exhibit high levels of endemism and species turnover that cannot be ascribed to geology or current climatic gradients. The pattern of the vegetation and flora on these ranges appears to be related to local topographical factors and the long period of time these landscapes have remained unglaciated and above sea level. Similar patterns have recently been described for iron outcrop communities in Brazil. In both regions these communities are being significantly impacted by mining. The high correlation between local endemism and restricted plant communities and high grade mineral deposits presents difficult challenges for achieving a comprehensive, adequate and representative reserve system in South Western Australia.

Comparative population structure and reproductive biology of the critically endangered shrub Grevillea althoferorum and two closely related more common congeners

Grevillea althoferorum is a critically endangered, sprouting shrub known from two disjunct populations within the South-West Botanical Province of Western Australia. This study compares the conservation biology of G. althoferorum and two closely related but more common congeners, the non-sprouter G. rudis and the sprouter G. synapheae subsp. pachyphylla in order to determine whether there are differences in reproductive and ecological attributes that might explain why G. althoferorum is rare. In contrast to the more common species, neither population of G. althoferorum exhibited evidence of seedling recruitment. However, the northern population was confirmed to be clonal and was actively recruiting from root suckers. Both populations of G. althoferorum were found to have reduced amounts of viable pollen on stigmas in comparison with the other species. The fruit set at the southern population of G. althoferorum was considerably lower than that found for the common species, with only 0.15% of flowers setting fruit and no fruit was produced in the northern population. In addition no evidence of a soil seed bank was found for either population of G. althoferorum, but G. rudis and G. synapheae subsp. pachyphylla both had soil stored seed which germinated following treatment with aqueous smoke solution. Sexual recruitment at both populations of G. althoferorum was absent, and reproduction appears to be predominantly clonal in the northern population. Management strategies for G. althoferorum should therefore focus on the protection of adult plants from accidental destruction.

Hierarchies of cause : Understanding rarity in an endemic shrub Verticordia staminosa (Myrtaceae) with a highly restricted distribution

Verticordia staminosa C.Gardner & A.C.George subsp. staminosa is an extremely rare shrub occurring as an isolated population of ∼1200 plants on a granite outcrop in the semi-arid agricultural region of Western Australia, separated from its closest relative V. staminosa subsp. cylindracea by 400 km. We aimed to determine a hierarchy of causes for explaining the extremely restricted distribution of subsp. staminosa, and to determine the genetic relationships among populations within both subspecies. We measured allozyme variation in all known populations of the two subspecies. There were exceptionally high levels of genetic divergence between subsp. staminosa and subsp. cylindracea, including an apparent duplication of the gene encoding phosphoglucomutase, leading to an additional gene in subsp. cylindracea. These findings combined with UPGMA analysis indicate a very long period of historical separation, perhaps originating in the early Pleistocene. Genetic variation was partitioned mostly between rather than within populations, with very low levels of genetic variation within populations of both subspecies. For subsp. staminosa we quantified seed production for three consecutive years and demography for five consecutive years. We used transition matrix models to describe the shrubs population dynamics and stochastic simulations to explicitly compare the effects of low rainfall and disturbance on population viability. Verticordia staminosa subsp. staminosa produces large numbers of seeds each year and has flower to fruit ratios greater than reported for related rare and common congeners. Seedling recruitment occurs in most years, with pulses in the wettest years. The mean finite population growth rate was 1.031. Elasticity analyses showed that population growth rate was more sensitive to stasis of established plants than to seedling recruitment. Population viability declined with lower rainfall and increased fire-related mortality of adult plants. Rarity in subsp. staminosa is best explained by evolutionary history and the interaction of climate change and disturbances such as fire that kill plants. Climatic fluctuations since the late Pliocene might have led to stochastic extinction episodes of populations on other granite outcrops, resulting in the currently restricted distribution. We discuss the implications of our findings for management of the species.