Two-scale modelling of Saponaria bellidifolia Sm. (Caryophyllaceae) abundance on limestone outcrops from its northern range periphery (Southeastern Carpathians)

We modelled the effect of habitat heterogeneity on the abundance of the submediterranean Saponaria bellidifolia, a red list species in Romania. The study was designed at two scales: 100 and 0.5 m². At larger scale, generalized additive models and canonical correspondence analysis were used to model the density of ramets, whereas at microscale, binomial logistic regression was employed to model the species’ occurrence. S. bellidifolia abundance responded sensitively to habitat type (classified as “grassy”, “rocky” and “scree”), rather than to microclimatic variables. At both scales, habitat type was the best predictor of ramet abundance, followed by slope and vegetation cover. At 0.5 m², soil depth was also a good predictor of species occurrence. The data revealed that screes are the most suitable habitats for hosting relatively large populations of this rare species, because of occasional natural disturbances and presumably lower interspecific competition.     

Significant patterns of population genetic structure and limited gene flow in a threatened macropodid marsupial despite continuous habitat in southeast Queensland, Australia

Many endangered species worldwide are found in remnant populations, often within fragmented landscapes. However, when possible, an understanding of the natural extent of population structure and dispersal behaviour of threatened species would assist in their conservation and management. The brush-tailed rock-wallaby (Petrogale penicillata), a once abundant and widespread rock-wallaby species across southeastern Australia, has become nearly extinct across much of the southern part of its range. However, the northern part of the species’ range still sustains many small colonies closely distributed across suitable habitat, providing a rare opportunity to investigate the natural population dynamics of a listed threatened species. We used 12 microsatellite markers to investigate genetic diversity, population structure and gene flow among brush-tailed rock-wallaby colonies within and among two valley regions with continuous habitat in southeast Queensland. We documented high and significant levels of population genetic structure between rock-wallaby colonies embedded in continuous escarpment habitat and forest. We found a strong and significant pattern of isolation-by-distance among colonies indicating restricted gene flow over a small geographic scale ( <10 km) and conclude that gene flow is more likely limited by intrinsic factors rather than environmental factors. In addition, we provide evidence that genetic diversity was significantly lower in colonies located in a more isolated valley region compared to colonies located in a valley region surrounded by continuous habitat. These findings shed light on the processes that have resulted in the endangered status of rock-wallaby species in Australia and they have strong implications for the conservation and management of both the remaining ‘connected’8 brush-tailed rock-wallaby colonies in the northern parts of the species’8 range and the remnant endangered populations in the south.     

Conservation of <Emphasis Type="Italic">Hymenocallis coronaria</Emphasis> genetic diversity in the presence of disturbance and a disjunct distribution

Evidence shows that dam construction has affected faunal species distribution and diversity in the southeastern U.S. However, the effects of this perturbation on aquatic flora of the region are heretofore uninvestigated. The objective for this research is to examine the effects of hydroelectric power generation and habitat characteristics on the size and distribution of populations of Hymenocallis coronaria (J. LeConte) Kunth (Amaryllidaceae), an emergent aquatic plant found in the southeastern United States, and to characterize its genetic variability in an effort to target conservation priorities. Multiple linear and logistic regression analyses show that hydropeaking does not have a significant effect on H. coronaria populations located downstream of dams, while other habitat variables associated with the streams are significantly positively related to population size and distribution. Analysis of 24 populations using seven universal cpDNA markers and six nDNA microsatellite markers indicates that the genetic structure for both genomes correlates with the location of a major gap in the species’ range, showing deep divergences in haplotype and population dendrograms between eastern and western regions. An AMOVA confirms that a substantial amount of genetic variation is found among regions, and very little variation is found within regions. The eastern and western regions have diverged for a significant time period, potentially due to the presence of a gene flow barrier in the middle of the species’ range caused by sub-standard habitat. Individual drainage basins contain unique cpDNA haplotypes and should be managed appropriately.     

Unexpectedly low genetic divergences among populations of the threatened bog turtle (<Emphasis Type="Italic">Glyptemys muhlenbergii</Emphasis>)

We used mitochondrial DNA sequence comparisons to assess range-wide population structure and historical patterns of differentiation among populations of the bog turtle (Glyptemys muhlenbergii). This species is one of North America’s smallest and most endangered pond turtles, and is currently found in three largely disjunct groups of populations: in the southern U.S., in the northeast, and in the Finger Lakes and Lake Ontario Plains region of western and central New York State. All the New York sites and most of the northeastern sites were glaciated during the Pleistocene. We surveyed 2793 bases pairs of mitochondrial DNA spanning three genes (cytb, nd4, and d-loop) in 41 individuals from 21 populations throughout most of the bog turtle’s distribution. We found surprisingly low levels of divergence among populations, even in southern populations that have been hypothesized as refugia during times of climate change. Our data suggest populations of bog turtle’s suffered a bottleneck, followed by a rapid post-Pleistocene expansion into northern segments of the species’ range. We discuss historical changes in habitat availability and climate that may have influenced the historical deployment of lineages in this species, and possible life history traits and habitat dynamics that might also contribute to the overall low genetic diversity across its range.     

Assessing the climatic sensitivity of Douglas-fir at its northern range margins in British Columbia, Canada

Northern hemisphere tree species growing at their northern range margins may be particularly responsive to climate change and could provide important information regarding future broad-scale responses. We analyzed and compared tree-ring data between five Douglas-fir [Pseudotsuga menziesii var. glauca (Mirb.) Franco] populations growing at the species’ northernmost distribution in British Columbia, Canada, and five populations located 150 km to the southeast. We quantified climate–growth relationships using uni- and multivariate techniques at different temporal scales. Our data suggest that (1) even at its northernmost distributions, precipitation limits long-term mature Douglas-fir radial growth more than temperatures, (2) northernmost Douglas-fir populations are distinct from populations located further within the species’ range in terms of certain key short-term growth responses, and (3) northernmost Douglas-fir growth sensitivities to climate may be increasing over time. In the future, mature Douglas-fir productivity in the northern portion of its range may be primarily limited by precipitation, and responses may be strongest at the species’ range margins.     

Genetic structure of peripheral, island-like populations: a case study of Saponaria bellidifolia Sm. (Caryophyllaceae) from the Southeastern Carpathians

Geographically peripheral populations often experience a reduction of genetic diversity and divergence from the core populations. Habitat geometry and quality can induce a local genetic diversity pattern, which overlies the regional variability issued from the range-wide phylogeography. We evaluated the genetic variation and genetic divergence of Saponaria bellidifolia Sm. on limestone outcrops within peripheral island-like populations from the Southeastern Carpathians, using RAPD markers. We also determined the degree of isolation related to other European populations, using AFLP. The Romanian populations had a decreased overall genetic diversity shared among populations, with lower level in small populations. Potential habitat size had a positive effect on genetic diversity estimates. Fisher’s exact tests of genetic differentiation revealed significant divergences only between the geographically most distant populations. Romanian populations were genetically pauperised as compared to Bulgarian and Italian populations and our results suggest that they might have originated from a recent range expansion from southern glacial refugia.     

Prevalence of multiple mating by female common dormice, <Emphasis Type="Italic">Muscardinus avellanarius</Emphasis>

Mating behaviour is an important component of species’ life histories. Knowledge of natural patterns of mating can lead also to more effective management strategies for populations of conservation concern. Despite a high conservation profile many aspects of the biology of the common dormouse (Muscardinus avellanarius) remain unknown, potentially limiting present conservation efforts. We determine the mating behaviour of M. avellanarius at two woodland sites in the UK: (1) Bontuchel (a natural population in Wales) and (2) Wych (a population in England that was established by reintroducing captive-bred animals) by genotyping mothers and litters at a panel of 10 microsatellite loci. Adult female body weight positively correlates with litter size and no apparent reproductive skew was evident. We found that multiple mating by female dormice is prevalent at both sites, with litters containing three or more offspring sired by multiple fathers; moreover, multiple mating is adopted by released animals even after a period of captive breeding where females are mated singly or as a breeding pair. We also present evidence for low proportion of fathers identified in our samples that probably related to unsampled individuals and/or larger than anticipated population sizes. This first report of mating behaviour in M. avellanarius highlights the role of genetic studies to uncover species’ reproductive behaviours and include these data for conservation management.     

Drainage-independent genetic structure and high genetic diversity of endangered freshwater pearl mussels (<Emphasis Type="Italic">Margaritifera margaritifera</Emphasis>) in northern Europe

Freshwater pearl mussels (Margaritifera margaritifera) are among the most critically threatened bivalve molluscs worldwide. An understanding of spatial patterns of genetic diversity is crucial for the development of integrative conservation strategies. We used microsatellites to study the genetic diversity and differentiation of 14 populations of M. margaritifera in central Sweden, an area which was described as a major secondary contact zone in postglacial colonisation for other species. Genetic diversity of Swedish pearl mussel populations was much greater than in central and southern Europe but similar to the genetic diversity observed in the northeastern portion of their European range. Genetic differentiation among populations was pronounced but to a large extent independent from present-day drainage systems. The complex patterns of genetic diversity and differentiation in pearl mussel seem to be strongly influenced by the species’ high degree of specialisation and extraordinary life history strategy which involves facultative hermaphrodism and an obligatory encystment stage on a host fish. Genetic drift effects and anthropogenic disturbances resulting in reduction of population size and loss of connectivity are less pronounced in northern pearl mussel populations compared to those in central and southern Europe.     

High Genetic Differentiation in Endangered <Emphasis Type="BoldItalic">Sedum ussuriense</Emphasis> and Implications for Its Conservation in Korea

Sedum ussuriense Kom. (Crassulaceae) is a succulent perennial herb localized to rocky valleys in southeastern Korea. Although it is an important natural resource with high economic value as an ornamental plant, it is currently endangered because of land-use changes and illegal exploitation. To initiate a proper conservation plan, we selected four populations (Juwang, Okgye, Jeolgol, and Haok) around Mt. Juwang, characterized their phenotypic traits, and evaluated patterns of random amplified polymorphic DNA variation. Despite its small population size, Okgye had the greatest proportion of flowering plants and higher seed production than from the other populations. This population also harbored the greatest genetic diversity. However, recent fragmentation between Okgye and Haok appeared to cause genetic divergence, leading to close genetic relationships of Okgye to Juwang vs. Haok to Jeolgol. In the long term, this raises concerns about the loss of genetic variation and the possibility of a demographic crash in those fragmented populations. Because our results indicated a high degree of divergence among populations, we suggest that conservation activities should focus on maintaining and propagating all populations throughout this species’ range.     

Effects of range position,inter-annual variation and density on demographic transition rates of <Emphasis Type="Italic">Hornungia petraea</Emphasis> populations

The central-marginal model assumes unfavourable and more variable environmental conditions at the periphery of a species’ distribution range to negatively affect demographic transition rates, finally resulting in reduced population sizes and densities. Previous studies on density-dependence as a crucial factor regulating plant population growth have mainly focussed on fecundity and survival. Our objective is to analyse density-dependence in combination with the effect of inter-annual variation and range position on all life stages of an annual plant species, Hornungia petraea, including germination and seed incorporation into the seed bank. As previous studies on H. petraea had revealed a pattern opposite to existing theory with lower population densities at the distribution centre in Italy than at the periphery in Germany, we hypothesised that (1) demographic transition rates are lower, (2) the inter-annual variation in demographic transition rates is higher and (3) the intensity of density-dependence is weaker in Italy than in Germany. To analyse demographic transition rates, we used an autoregressive covariance strategy for repeated measures including density and inter-annual variation. All the three hypotheses were confirmed, but the impact of range position, density-dependence and inter-annual variation differed among the transition steps. All transition rates except fecundity were higher in the German populations than in the Italian populations. Germination rate and incorporation rate into the seed bank were strongly density-dependent. Central populations showed a larger inter-annual variation in fecundity and winter survival rate. Winter survival rate was the only transition step with a stronger density-dependence in peripheral populations. In most cases, these differences between distribution centre and periphery would not have emerged without taking density-dependence and inter-annual variation into account. We conclude that including range position, inter-annual variation and density-dependence in one single statistical model is an important tool for the interpretation of demographic patterns regarding the central-marginal model. Electronic Supplementary Material Supplementary material is available for this article at     

Dynamics of a small re-introduced population of wild dogs over 25 years: Allee effects and the implications of sociality for endangered species’ recovery

We analysed 25 years (1980–2004) of demographic data on a small re-introduced population of endangered African wild dogs (Lycaon pictus) in Hluhluwe-iMfolozi Park (HiP), South Africa, to describe population and pack dynamics. As small populations of cooperative breeders may be particularly prone to Allee effects, this extensive data set was used to test the prediction that, if Allee effects occur, aspects of reproductive success, individual survival and population growth should increase with pack and population size. The results suggest that behavioural aspects of wild dogs rather than ecological factors (i.e. competitors, prey and rainfall) primarily have been limiting the HiP wild dog population, particularly a low probability of finding suitable mates upon dispersal at low pack number (i.e. a mate-finding Allee effect). Wild dogs in HiP were not subject to component Allee effects at the pack level, most likely due to low interspecific competition and high prey availability. This suggests that aspects of the environment can mediate the strength of Allee effects. There was also no demographic Allee effect in the HiP wild dog population, as the population growth rate was significantly negatively related to population size, despite no apparent ecological resource limitation. Such negative density dependence at low numbers indicates that behavioural studies of the causal mechanisms potentially generating Allee effects in small populations can provide a key to understanding their dynamics. This study demonstrates how aspects of a species’ social behaviour can influence the vulnerability of small populations to extinction and illustrates the profound implications of sociality for endangered species’ recovery. Electronic supplementary material  The online version of this article (doi:) contains supplementary material, which is available to authorized users.     

Population genetic structure and history of fragmented remnant populations of the New England cottontail (<Emphasis Type="Italic">Sylvilagus transitionalis</Emphasis>)

The New England cottontail (Sylvilagus transitionalis) has suffered from extensive loss and fragmentation of its habitat and is now a species of conservation priority in the northeastern United States. Remnant New England cottontail populations currently occur in five geographically disjunct locations: southern Maine and southeastern New Hampshire (MENH); the Merrimack Valley in central New Hampshire (NH-MV); Cape Cod, Massachusetts (CC); parts of eastern Connecticut and Rhode Island (CTRI); and western Connecticut, southeastern New York and southwestern Massachusetts (CTNY). We used microsatellite genotyping to discern patterns of population structure, genetic variability, and demographic history across the species’ range and to assess whether the observed patterns are a consequence of recent habitat loss and fragmentation. Our findings show that the geographic populations are highly differentiated (overall F _ST = 0.145; P < 0.001). Using Bayesian clustering analyses, we identified five genetic clusters, which corresponded closely to the geographic populations, but grouped MENH & NH-MV together (ME/NH) and identified an isolated population in eastern Connecticut (Bluff Point). The genetic clusters showed little evidence of recent gene flow and are highly influenced by genetic drift. The CC and Bluff Point populations show signs they experienced a genetic bottleneck, whereas the ME/NH population shows evidence of ongoing decline. Populations in Bluff Point, CC, and ME/NH also show significantly reduced genetic variation relative to the other clusters (CTNY and CTRI without Bluff Point). Without immediate human intervention, the short-term persistence of New England cottontail populations in Maine, New Hampshire and Cape Cod is at great risk. Conservation efforts at this time should focus on within-population sustainability and eventually restoring connectivity among these isolated populations.     

Patterns of spatial genetic structure and diversity at the onset of a rapid range expansion: colonisation of the UK by the small red-eyed damselfly <Emphasis Type="Italic">Erythromma viridulum</Emphasis>

Species’ geographic ranges may vary in size in response to a change in environmental conditions. The specific genetic consequences of range expansions are context dependent, largely depending upon the rate of colonisation as well as the origins and numbers of founders, and the time since colonisation. Like other “charismatic” taxa, such as birds and lepidopterans, the distributions of odonates (dragonflies and damselflies) are well-known through substantial monitoring programmes co-ordinated by various societies. The small red-eyed damselfly Erythromma viridulum (Odonata: Zygoptera) has undergone a substantial, northward range expansion in Europe in the last 30 years and has recently-colonised two distinct areas in the UK. We quantify the immediate genetic consequences of this rapid colonisation by genotyping more than 1,400 E. viridulum from 39 sites across the northwest margin of this species’ geographic range. Levels of genetic diversity and spatial structure are impacted by this species recent range expansion and non-equilibrium conditions that drive weak genetic divergence, even at regional spatial scales. Populations of E. viridulum become less diverse towards the edge of this species’ distribution, presumably as a consequence of colonisation through a series of founder events. Specifically, there is a significant reduction in genetic diversity in the smallest, most recent focus of colonisation in the UK; however, there are generally low levels of genetic diversity across this E. viridulum’s northern range margin. While most populations are generally poorly differentiated, E. viridulum nonetheless consists of two distinct lineages that broadly differentiate between eastern and western Europe. Genetic divergence between the two UK colonisation foci are indicative of distinct immigration events from separate sources; however a general lack of spatial structure prevents us from pinpointing the specific origins of these migrant damselflies.     

Genetic diversity and conservation units in wild and captive populations of endangered freshwater fishes: a case of <Emphasis Type="Italic">Hemigrammocypris rasborella</Emphasis> in Shizuoka,Japan

Population structure and genetic diversity were examined using partial mitochondrial cytochrome b gene sequences of four wild, one reintroduced, and five captive populations of the endangered cyprinid Hemigrammocypris rasborella from three river systems in the easternmost region of the species’ range in Shizuoka Prefecture, central Honshu, Japan. We detected loss of genetic diversity from portions of the wild and captive populations, as well as suspected nonindigenous haplotypes in some captive, reintroduced, and even wild populations. Given the population structure revealed, we suggest that the populations should be managed with consideration for both the endemism and viability (avoidance of inbreeding depression) of the local populations.     

Environmental constraints on the invasion of <Emphasis Type="Italic">Triadica sebifera</Emphasis> in the eastern United States: an experimental field assessment

Identifying the environmental constraints that affect the distribution of an invasive species is fundamental to its effective control. Triadica sebifera (Chinese tallow tree) has invaded the southeastern United States, but its potential for further range and habitat extension has been unresolved. We explored experimentally environmental factors in macro- and microhabitats that affect its persistence at five widely separated sites along the Atlantic seaboard of the United States and at two sites inland; three sites occur well beyond the tree’s current range. At each site, seeds and young vegetative plants (0.5–0.65 m tall) of T. sebifera were placed in four microhabitats (closed-canopy upland, closed-canopy lowland, open-canopy upland, and open-canopy lowland). Plant growth, leaf CO₂ assimilation rates, leaf N concentrations and δ¹³C ratios, and stem water potential were measured for two growing seasons. Percent seed germination was consistently higher in open-canopy microhabitats and lowest at northern and inland sites. T. sebifera grew in all open-canopy microhabitats, even 300–500 km beyond its current distribution. Plant growth in closed-canopy habitats was lower, attributable to lower carbon gain per unit leaf area in shaded compared with open-canopy environments, especially at northern and inland sites. Neither competition, other than canopy shade, nor grazing was a key constraint on distribution at any scale. Our results demonstrate that T. sebifera is dispersal limited at landscape scales but limited locally by dispersal and overstory shade; it has yet to occupy the full extent of its new range in North America. Quantifying environmental factors both within and well beyond a species’ current range can effectively highlight the limits on its distribution.     

The genetic structure of <Emphasis Type="Italic">Quercus crispula</Emphasis> in northeastern Japan as revealed by nuclear simple sequence repeat loci

Previous studies have reached different discussions about the genetic variation and genetic structure of Quercus crispula populations in northeastern Japan. This is a common oak species in Eastern Asia. Some studies have suggested that the populations in northeastern Japan were derived from those remaining in the southwest after the last glacial maximum (LGM), whilst other studies have found evidence that populations persisted in northeastern Japan during the LGM. Using seven highly polymorphic nuclear simple sequence repeat loci, we investigated the genetic structure of 16 Q. crispula populations along a latitudinal gradient in northeastern Japan (northern Honshu and Hokkaido), spanning about half of the species’ biogeographic range in the country. Although the level of population differentiation was low (F _ST = 0.021; G^￠_\textST G^{\prime}_{\text{ST}}  = 0.090), two geographically differentiated clusters were detected by STRUCTURE analysis. The first cluster included most of the populations in Hokkaido, and may indicate continued survival throughout past glacial periods. We found a significant decrease in allelic richness with latitude, so the second cluster may represent an expansion of the lineage from Honshu during the post-glacial period. These results should enhance our understanding of historical north–south migrations of this species in northeastern Japan.     

Chinese mitten crab <Emphasis Type="Italic">Eriocheir sinensis</Emphasis> in the Baltic Sea—a supply-side invader?

Although the Chinese mitten crab Eriocheir sinensis (H. Milne-Edwards, 1853) (Crustacea, Decapoda, Varunidae) invaded the Baltic Sea about 80 years ago, published information on its present distribution and abundance in this region is lacking. We provide here information on its Baltic-wide distribution and long-term population dynamics. The species has been found all over the coastal Baltic Sea and also in some adjacent rivers and lakes. The Chinese mitten crab appears to have increased in abundance in recent years in the northeastern part of the Baltic Sea (Gulf of Finland, Gulf of Riga, northern Baltic Proper). Higher catch rates were observed in spring (April–June) and autumn (September–November). The size variation of crabs in different samples was low (mean carapace width 6.1–6.3 cm). Despite findings of gravid females, the reproduction of the mitten crab in the central, northern and eastern Baltic region is considered unlikely due to low salinity and the individuals caught are assumed to actively migrate into the region from the species’ main European distribution area (southeastern North Sea), certainly over 1500 km migration distance. Thus, the dynamics of the North Sea population is probably regulating, at least in part, the occurrence of the Chinese mitten crab in the Baltic Sea area.     

Inter-specific relationships and hierarchical spatial genetic structuring in Nervilia nipponica, an endangered orchid in Japan

An understanding of the extent to which reproductive strategy and seed dispersal lead to the structuring of genetic diversity in space is required when planning measures towards the conservation of endangered plant species. In this study, genetic structure in the endangered terrestrial orchid Nervilia nipponica was investigated using amplified fragment length polymorphisms following extensive sampling throughout the species’ range in Japan and intensive sampling at a single population. Limited diversity was found within the species as a whole, but significant structuring was detected between populations. One genotype was common to two widely separated sites, possibly indicative of long-range dispersal. Significant structure was also detected at the intensively sampled site, as a result of the presence of two distinct putative clones. These findings are consistent with observations of the species’ ability to set seed autogamously and propagate vegetatively. Given the strong colonising capability inferred for the species, attention should focus on identifying and securing habitat conditions conducive to seed germination and seedling establishment in the development of a conservation strategy. As presently circumscribed, N. nipponica is shown to comprise two polyphyletic taxa, both endemic to Japan, and both distinct from N. taiwaniana, a species that some authors have considered conspecific.     

Evidence of autumn reproduction in female European hares (Lepus europaeus) from southern Europe

Information on reproductive biology of the European hare (Lepus europaeus) in different environmental and landscape conditions comprises part of fundamental knowledge regarding species’ adaptive responses as well as many aspects of its biology. Most of the studies conducted on European hare reproduction are confined to midlatitude and northern populations, whereas no data exist on the indigenous southern populations. Here, we present information on reproductive characteristics of European hares inhabiting Mediterranean ecosystems on the island of Crete, Greece for two successive hunting seasons. Although the annual reproductive cycle of the species is well known, with an autumn sexual inactivity, the duration of this period is subjected to fluctuations in different years and for different areas. According to our data, hare populations of Crete present an autumn–early winter reproductive activity with high proportions of pregnant females observed in all the months of the study. Furthermore, the estimated mean litter size (1.54 SE ± 0.07) while signed to the lowest values ever observed for European hares is similar to values obtained in continuous breeding species of the same genus, Lepus granatensis, Lepus corsicanus, Lepus (capensis) mediterraneus, and Lepus capensis also inhabiting warm climates. In conclusion, our results suggest that Cretan European hare populations exhibit a reproductively active period during autumn–early winter where proportions of pregnant females and litter size give a strong indication of a continuous reproduction throughout the year.     

Effect of reproductive modes and environmental heterogeneity in the population dynamics of a geographically widespread clonal desert cactus

The dynamics of plant populations in arid environments are largely affected by the unpredictable environmental conditions and are fine-tuned by biotic factors, such as modes of recruitment. A single species must cope with both spatial and temporal heterogeneity that trigger pulses of sexual and clonal establishment throughout its distributional range. We studied two populations of the clonal, purple prickly pear cactus, Opuntia macrocentra, in order to contrast the factors responsible for the population dynamics of a common, widely distributed species. The study sites were located in protected areas that correspond to extreme latitudinal locations for this species within the Chihuahuan Desert. We studied both populations for four consecutive years and determined the demographic consequences of environmental variability and the mode of reproduction using matrix population models, life table response experiments (LTREs), and loop and perturbation analyses. Although both populations seemed fairly stable (population growth rate, λ∼1), different demographic parameters and different life cycle routes were responsible for this stability in each population. In the southernmost population (MBR) LTRE and loop and elasticity analyses showed that stasis is the demographic process with the highest contributions to λ, followed by sexual reproduction, and clonal propagation contributed the least. The northern population (CR) had both higher elasticities and larger contributions of stasis, followed by clonal propagation and sexual recruitment. Loop analysis also showed that individuals in CR have more paths to complete a life cycle than those in MBR. As a consequence, each population differed in life history traits (e.g., size class structure, size at sexual maturity, and reproductive value). Numerical perturbation analyses showed a small effect of the seed bank on the λ of both populations, while the transition from seeds to seedlings had an important effect mainly in the northern population. Clonal propagation (higher survival and higher contributions to vital rates) seems to be more important for maintaining populations over long time periods than sexual reproduction.