The predictive accuracy of population viability analysis: a test using data from two small mammal species in a fragmented landscape

This study examines the predictive accuracy of the population viability analysis package, ALEX (Analysis of the Likelihood of EXtinction). ALEX was used to predict the probability of patch occupancy for two species of small native Australian mammals (Antechinus agilis and Rattus fuscipes) among 13 patches of suitable habitat in a matrix of plantation pines (Pinus radiata). The study was retrospective, running each simulation from 1900 until 1997, and the model parameterised without knowledge of the 1997 observed field data of patch occupancy. Predictions were made over eight scenarios for each species, allowing for variation in the amount of dispersal between patches, level of environmental stochasticity, and size of habitat patches. Predicted occupancies were compared to the 1997 field data of patch occupancy using logistic regression, testing H _random, that there was no relationship between observed and predicted occupancy, and H _perfect, that there was a perfect, 1:1 relationship between observed and predicted occupancies. Rejection of H _random and failure to reject H _perfect was taken as a good match between observed and predicted occupancies. Such a match was found for one scenario with R. fuscipes, and no scenarios with A. agilis. In general, patch occupancy was underestimated, with field surveys finding that 9 of the 13 patches were occupied by R. fuscipes and 10 by A. agilis. Nonetheless, PVA predictions were in the right direction, whereby patches predicted to have a high probability of occupancy were generally occupied, and vice versa. A post hoc search over additional scenarios found few scenarios with a better match than the original eight. The results of this study support the notion that PVA is best thought of as a relative, rather than absolute predictor of the consequences of management actions in threatened populations.     

Current state and the future of the crested ibis (Nipponia nippon): A case study by population viability analysis

There is only one population of the crested ibis (Nipponia nippon) surviving in the wild, which has not exceeded 40 individuals since it was rediscovered in 1981. We used detailed data of the annual population size and natality collected by the Crested Ibis Conservation and Observation Station (CICOS) over the past 17years. We were able to obtain a more accurate census of age structure and mortality as all of the fledglings have been identified by leg bands in 1987. Consequently, we analyzed the current state of the crested ibis and predicted its future using Population Viability Analysis (PVA) with the help of the computer simulation model VORTEX (Version 7.2). The results indicated that the extinction probability of the crested ibis was 19.7% in 100years. Sensitivity analysis showed that the extinction probabilities were sensitive to catastrophes and environmental variation, and the long-term existence of the population was dependent on the carrying capacity of the habitat and inbreeding depression. Conservation efforts are critical for the survival of the crested ibis. The prior conservation project involves controlling hunting and predation, as well as improving the life level of these few remnant creatures.     

Using ecological theory to plan restorations of the endangered Beach jacquemontia (Convolvulaceae) in fragmented habitats

Population viability and metapopulation theory and models are heuristic tools that can be used to plan restorations and assess their success. Using examples from South Florida, USA, we review background information and ongoing reintroduction experiments with the federally endangered coastal perennial vine, beach jacquemontia, Jacquemontia reclinata (Convolvulaceae). All known wild populations are declining in isolated habitat fragments varying in size, occupied area, and degree of isolation. Eleven reintroduction sites ranging in size from 422 to 4800 m² within the extant species’ range have been identified that have characteristics suitable for J. reclinata introductions and have land managers amenable to restoration efforts. Previous RAPD analysis indicated that genetic diversity of natural populations was positively correlated with population size; the two largest populations had the highest genetic diversity and the smallest populations had relatively low genetic diversity. Despite habitat fragmentation and large distances between some populations, migration rates were very high among populations (m=4.05). Experimental crosses indicated the species has a mixed mating system. From 2001 to 2005, we have introduced 935 J. reclinata in seven experiments in five locations using plants propagated ex situ at Fairchild Tropical Botanic Garden. Reintroductions have dramatically increased the number of plants in the wild by 72%. Survival from the time of transplant to 2005 ranged from 2% to 97%, was not significantly correlated with metapopulation parameters, such as, founding population size, patch size, or connectivity to extant populations. Reintroduced plants are contributing seed and pollen to the wild populations, but no recruited seedlings have yet been observed. Although it may take decades before we can consider the reintroduced populations to be self-sustainable, we argue that planning restorations for rare species based on predictions from ecological theory is advisable to allow a higher probability of success.     

Mitochondrial genes reveal cryptic diversity in plant-breeding frogs from Madagascar (Anura, Mantellidae, Guibemantis)

One group of mantellid frogs from Madagascar (subgenus Pandanusicola of Guibemantis) includes species that complete larval development in the water-filled leaf axils of rainforest plants. This group consists of six described species: G. albolineatus, G. bicalcaratus, G. flavobrunneus, G. liber, G. pulcher, and G. punctatus. We sequenced the 12S and 16S mitochondrial rRNA genes ( approximately 1.8 kb) from multiple specimens (35 total) of all six species to assess phylogenetic relationships within this group. All reconstructions strongly supported G. liber as part of the Pandanusicola clade, even though this species does not breed in plant leaf axils. This result confirms a striking reversal of reproductive specialization. However, all analyses also indicated that specimens assigned to G. liber include genetically distinct allopatric forms that do not form a monophyletic group. Most other taxa that were adequately sampled (G. bicalcaratus, G. flavobrunneus, and G. pulcher) likewise consist of several genetically distinct lineages that do not form monophyletic groups. These results suggest that many of the recognized species in this group are complexes of cryptic species.     

The influences of habitat, landscape structure and climate on local distribution patterns of the nuthatch (Sitta europaea L.)

The nuthatch, Sitta europaea L., is a small (23 g), cavity-nesting woodland bird which, since the 1970s, has been expanding its range in Britain. However, within this range, the species is notably scarce in an area of eastern England. This gap in the species distribution could arise for several reasons including habitat quality, local landscape structure, regional landscape structure and climate. Field surveys and logistic models of breeding nuthatch presence/absence were used to investigate the relative influences of habitat quality, landscape structure and climate on the prevalence of nuthatches in eastern England. Field surveys of woods in the study area indicated that habitat quality was sufficient to support a nuthatch population. A model of habitat occupancy in relation to local landscape structure, developed in the Netherlands, was applied to the study area. The number of breeding pairs predicted for the study area by the model was lower than expected from habitat area alone, suggesting an additional effect of isolation. However, observed numbers were even lower than those predicted by the model. To evaluate the possible roles of climate and large-scale landscape structure on distribution, presence/absence data of breeding nuthatches at the 10-km grid square scale were related to variables describing climate and the amount and dispersion of broadleaved woodland. While climate in the study area appeared suitable, models including landscape variables suggested that the study area as a whole was unlikely to support nuthatches. Although suitable habitat was available, woodland in the study area appeared to be too isolated from surrounding nuthatch populations for colonisation to be successful. This situation may change if current increases in both national and regional populations continue, thus increasing the number of potential colonists reaching the study area. Received: 3 November 1997 / 22 January 1998     

Demographic analysis of a dominant shrub (Cytisus scoparius): Prospects for encroachment control

Shrub encroachment in pastoral areas subsequent to land-use changes has become a major problem for biodiversity and pastoral resource conservation. Controlling encroachment by grazing is the most promising prospect at this time, but the impact of small domestic ruminants on the population dynamics of dominant shrubby species is still poorly understood. To control the rapid densification of dominant shrub populations means that we have to focus on the early stages of shrub colonisation. In this paper, we present a modelling approach that includes deterministic matrix modelling to understand demography strategy at the very beginning of shrub population growth and to identify potential targeted life stages for control. We collected demographic data over a period of 5 years on the European dominant shrub, Cytisus scoparius (Scotch broom; Fabaceae), which is often part of the heathlands that establish themselves on former permanent grasslands in Europe. Our results reveal a very high variation in flowering frequency between years and a large contribution of the seed bank to the total number of seedlings that emerge each year. Modelling shows that population growth is very vigorous and is not stopped by repeated “bad years” for seed production. Sensitivity analyses indicate that early life stage survival (seedling, juvenile) has the greatest impact on population growth. These results make it possible to identify juveniles as one of the best targets for control since they combine a high degree of sensitivity in terms of population demography, good palatability for domestic herbivores and easy accessibility within a management schedule.     

Host-parasitoid extinction and colonization in a fragmented prairie landscape

Few field studies of natural populations have examined the factors influencing local extinctions and colonization of empty habitat patches for a prey species and its predator. In this study, I carried out a census of planthopper (Prokelisia crocea; Hemiptera: Delphacidae) and egg parasitoid (Anagrus columbi; Hymenoptera: Mymaridae) incidence and densities in 147 host-plant patches (Spartina pectinata; Poaceae) over seven planthopper generations in a tall-grass prairie landscape. For both species, the likelihood of going extinct in a patch was related to a number of patch-specific variables: density, temporal variability in density, proportion of hosts parasitized (planthopper only), host-plant density, patch size, patch isolation, and composition of the surrounding matrix. Colonization likelihood was related only to the physical attributes of the patch. There was high patch turnover in this prairie landscape. On average, planthoppers went extinct in 23% of the patches and A. columbi went extinct in 51% of the patches in each generation. For the planthopper, extinction likelihood increased with a decrease in patch size and the proportion of the matrix composed of mudflat. Parasitism of eggs had no effect on the extinction likelihood of local P. crocea populations, suggesting that A. columbi may not play a major role in the patch dynamics of its host. The likelihood of extinction for A. columbi was dependent on factors that spanned three trophic levels. An increase in plant density, decrease in host density and decrease in parasitoid density all increased the likelihood of A. columbi extinction within a patch. The dependency on multiple trophic levels may explain the higher extinction risk for the parasitoid than its host. A. columbi extinction was also affected by the matrix habitat surrounding the patch—the effect was the opposite of that for P. crocea. Finally, vacant patches were colonized at rates of 53% and 34% per generation for the planthopper and parasitoid, respectively. For both species, colonization probabilities decreased with an increase in patch isolation. High host densities in a patch also favored high rates of colonization by A. columbi. I discuss how anthropogenic changes to the prairie landscape can affect the metapopulation dynamics and persistence time of this host-parasitoid interaction.     

Viability and Management of an Endangered Capercaillie (Tetrao urogallus) Metapopulation in the Jura Mountains, Western Switzerland

The populations of Capercaillie (Tetrao urogallus), the largest European grouse, have seriously declined during the last century over most of their distribution in western and central Europe. In the Jura mountains, the relict population is now isolated and critically endangered (about 500 breeding adults). We developed a simulation software (TetrasPool) that accounts for age and spatial structure as well as stochastic processes, to perform a viability analysis and explore management scenarios for this population, capitalizing on a 24 years-long series of field data. Simulations predict a marked decline and a significant extinction risk over the next century, largely due to environmental and demographic stochasticity (average values of life-history parameters would otherwise allow stability). Variances among scenarios mainly stem from uncertainties about the shape and intensity of density dependence. Uncertainty analyses suggest to focus conservation efforts on enhancing, not only adult survival (as often advocated for long-lived species), but also recruitment. The juvenile stage matters when local populations undergo extinctions, because it ensures connectivity and recolonization. Besides limiting human perturbations, a silvicultural strategy aimed at opening forest structure should improve the quality and surface of available patches, independent of their size and localization. Such measures are to be taken urgently, if the population is to be saved.     

Utilisation of a tropical bay as a nursery area by sharks of the families Carcharhinidae and Sphyrnidae

Synopsis At least eight species of sharks of the families Carcharhinidae and Sphyrnidae use Cleveland Bay in northern Australia as a communal nursery area.Carcharhinus dussumieri, C. fitzroyensis, C. limbatus andC. tilstoni use the bay as a seasonal primary nursery, with juveniles occurring in it for only a few months each year immediately after birth. Alternatively,Carcharhinus sorrah, Rhizoprionodon acutus andR. taylori use the bay as a year-round primary and secondary nursery, with juveniles remaining in it up to the size at maturity. AdultR. taylori also persist in the bay, a behavioural pattern possibly explained by their small maximum size. While present immediately after birth the type of utilisation pattern displayed bySphyrna lewini could not be clarified in this study. Although diets of these species in the bay are similar, there is probably little direct competition for food due to the highly productive habitats in the bay supporting an abundance of food resources. The highest numbers of juveniles occur when prey species are the most abundant, and when temporal separation of some seasonally-occurring species of sharks in effect.     

Over-winter survival in subadult European rabbits: weather effects,density dependence,and the impact of individual characteristics

The survival probability of an individual may be limited by density-dependent mechanisms and by environmental stochasticity, but can also be modified by individual characteristics. In our study, we investigated over-winter survival of subadults of an enclosed European rabbit Oryctolagus cuniculus population in a temperate zone habitat over the period 1992–2002. We: (1) selected for appropriate models to explain individual variation in over-winter survival and the animals autumn body mass, the latter was used as a measure of the individual pre-winter body condition; and (2) aimed to compare the sensitivity of the target variables on the realised variation of the factors considered. Model selection based on information theory revealed that individual over-winter survival was best explained by the combination of autumn body mass, winter temperature, population density and sex, where the probability of survival was higher in females than in males. According to this model, the probability of survival reacted most sensitively to variation in the autumn body mass and in winter temperature. Individual autumn body mass was best explained by the combination of the date of birth, population density, and weather conditions by means of the percentage of rainy days during the first 2 months after the animals had emerged above ground, where the autumn body mass was negatively related to the percentage of rainy days. The chosen model suggested that the autumn body mass reacted most sensitively to variation in the date of birth. Combining these models, we found that weather conditions during two different periods of time as well as population density, sex and the date of birth operated together to determine the probability of over-winter survival. In particular, the study points out the high impact of environmental stochasticity on over-winter survival: (1) by direct effects of winter temperature conditions, and (2) by the indirect action of weather conditions to which the animals were exposed during the early period of juvenile development.     

Is one parasitoid enough? A test comparing one with a pair of parasitoid species in the biological control of arrowhead scales

In classic biological control using natural enemies, the question of whether a single species or multiple species should be introduced has been a matter of debate. The introduction of two parasitoids, Aphytis yanonensis and Coccobius fulvus (Hymenoptera: Aphelinidae), to control the arrowhead scale, Unaspis yanonensis (Hemoptera: Diaspididae), which is a serious pest in Japanese citrus orchards, has been one of the most successful biological control projects in Japan. The success of this program may be explained by two alternative hypotheses: (1) the parasitoid species work complementarily, or (2) only one of them plays a major role. To test which hypothesis is applicable to this host-parasitoid system, we conducted caging experiments and observed temporal changes in the proportion of the parasitisms and the densities of arrowhead scales enclosed with one of the following combinations of parasitoids: (1) A. yanonensis and C. fulvus together, (2) A. yanonensis alone, (3) C. fulvus alone, or (4) neither parasitoid. Parasitisms in the cohorts with A. yanonensis and C. fulvus together and C. fulvus alone rapidly increased to approximately 70%; parasitism with A. yanonensis alone also increased slightly, although it remained consistently lower that those with A. yanonensis and C. fulvus together and C. fulvus. At the end of the experiment, parasitisms with A. yanonensis and C. fulvus together and C. fulvus alone were significantly higher than that with A. yanonensis alone. Parasitism by C. fulvus constituted most of (74%) the parasitism in the cohort with A. yanonensis and C. fulvus together. Further, only C. fulvus suppressed the population growth rates of scales significantly. These results suggest that C. fulvus alone successfully suppresses scale populations as efficiently as both species together do.     

Defining core habitat of local populations of the gray treefrog (Hyla versicolor) based on choice of oviposition site

Concern over amphibian population declines and loss of terrestrial and aquatic habitat have emphasized the need to define habitat requirements for each stage in a species' life history. The realization that pond-breeding amphibians spend most of their lives in the terrestrial environment suggests the need to protect terrestrial as well as aquatic habitat. Many studies on amphibian populations have focused on emigration from breeding sites to define habitat use; however these studies do not typically elucidate terrestrial activities of adults within the breeding season. We measured colonization rates of artificial pools by gray treefrogs (Hyla versicolor) at multiple distances from natural breeding ponds. We found a non-random distribution of egg deposition among distances, with 95% of eggs deposited within 15 m of the breeding pond. Additionally, we found that the time to first colonization of artificial pools increased with respect to distance. Our results indicate that adult gray treefrogs may travel up to 200 m within a breeding season, and that multiple breeding ponds may be considered part of a single population. We suggest that a minimum core terrestrial habitat of 60 m surrounding breeding sites is appropriate for protection of local populations of gray treefrogs.     

Growth and Survival of Early Juveniles of the Marine Sponge Hymeniacidon perlevis (Demospongiae) Under Controlled Conditions

To resolve “the supply problem” in sponge-derived drug development and other biotechnological applications, current research is exploring the possibility of obtaining an alternative sustainable supply of sponge biomass through intensive aquaculture of sponges utilizing artificial seed rearing. This study aimed to investigate the technology of early juvenile sponge cultivation under controlled conditions. The effects of food, temperature, water flow, and light on the growth and survival of early juveniles of the marine sponge Hymeniacidon perlevis were examined. The concentrations of four types of food elements [microalgae (Isochrysis galbana), photosynthetic bacteria (Rhodopseudomonas), Fe³⁺ (FeCl₃), and Si (Na₂SiO₃)] were investigated for early H. perlevis juvenile growth. Interestingly, temperature changes have striking effects on juvenile growth. Juvenile sponges grow faster when they are shifted to higher temperatures (18°C to 23°C) than when they are shifted to lower temperatures (18°C to 4°C to 23°C) or kept at a constant temperature (18°C). Periodic water flow and light cycles favor early juvenile sponge growth. Light was found to be a key factor in the color loss of early H. perlevis juveniles. Overall, size (area) increased as much as 29 times for H. perlevis juveniles under the tested controlled conditions.     

Genetic variation for foot-rot and Fusarium head-blight resistances among full-sib families of a self-incompatible winter rye (Secale cereale L.) population

The amount of genetic variation for resistance to foot rot caused by Pseudocercosporella herpotrichoides, Fusarium spp., and Microdochium nivale and for resistance to head blight caused by Fusarium culmorum are important parameters when estimating selection gain from recurrent selection in winter rye. One-hundred and eighty-six full-sib families of the selfincompatible population variety Halo, representing the Petkus gene pool, were tested for foot-rot resistance at five German location-year combinations (environments) and for head-blight resistance in three environments with artificial inoculation in all but one environment. Foot-rot rating was based on 25 stems per plot scored individually on a 1–9 scale. Head-blight resistance was plotwise scored on a 1–9 scale and, additionally, grain-weight per spike was measured relative to the non-inoculated control plots. Significant estimates of genotypic variance and medium-sized heritabilities (h²=0.51–0.69) were observed in the combined analyses for all resistance traits. In four out of five environments, the amount of genetic variance was substantially smaller for foot-rot than for head-blight rating. Considerable environmental effects and significant genotype-environment interactions were found for both foot-rot and head-blight resistance. Coefficients of error-corrected correlation among environments were considerably closer than phenotypic correlations. No significant association was found between the resistances to both diseases (r=-0.20 to 0.17). In conclusion, intra-population improvement by recurrent selection should lead to substantial higher foot-rot and head-blight resistances due to significant quantitative genetic variation within Halo. Selection should be carried out in several environments. Lack of correlation between foot-rot and head-blight resistance requires separate infection tests for improving both resistances.     

Spatial population structure in the banner-tailed kangaroo rat,Dipodomys spectabilis

I attempted to characterize spatial units of local dynamics and dispersal in banner-tailed kangaroo rats (Dipodomys spectabilis), to determine if spatial structure influenced population dynamics in the way predicted by current metapopulation models. D. spectabilis exhibited a hierarchical spatial structure. Local populations that appeared as discrete entities on a scale of kilometers were subdivided into clusters of mounds on a scale of meters. This structure, however, cannot be characyerized in terms of the discrete habitat patches envisioned by the metapopulation models. Occupied areas were statistically distinguishable from the surrounding matrix, but this difference was only quantitative. There were no discrete boundaries between occupied areas and the matrix. Habitat within occupied areas was heterogeneous, and occupied areas in different locations were statistically distinguishable from each other. High heterogeneity within occupied areas, and high contrast among them, make it difficult to define what is a suitable habitat patch for D. spectabilis. On a smaller spatial scale, there was significant aggregation of resident mounds within occupied areas. These aggregations, however, do not correspond to discrete habitat patches. Rather, they appear to result from an interaction between fine-scale habitat heterogeneity and limited dispersal due to natal philopatry and low adult vagility. These complications make it difficult to identify habitat patches independent of the species' distribution. For species like D. spectabilis that are patchily distributed but do not occupy discrete habitat patches, a patch occupancy approach does not seem appropriate for describing spatial structure. Hierarchical spatial structure underscores the need for a framework that incorporates multiple scales of spatial structure, rather than one that pre-imposes a single spatial scale as being important for population dynamics. A framework that (i) considers patchiness as a combination of both habitat heterogeneity, and life-history and behavioral characteristics, and (ii) incorporates hierarchical spatial structure, appears to be the most suitable for conceptualizing spatial dynamics of behaviorally complex vertebrates such as D. spectabilis.     

Distance-dependent pollen limitation of seed set in some insect-pollinated dioecious plants

By applying hand pollination, we studied pollen limitation of seed set of female plants in four dioecious plant species with insect-pollination. The effect of hand pollination increased with distance to the nearest male plant. Distances at which seed set was 50% of its maximal value (after hand pollination) were: 2.3 m for Valeriana dioica, 5.3 m for Salix repens, 8.5 m for Asparagus officinale and 10.6 m for Bryonia dioica. We discuss to what extent the reduced seed set was caused by either fewer visits or by visits of a lower quality. We argue that quantifying distance-dependent seed set in dioecious plants may be a good way of studying effects of habitat fragmentation on the breakdown of mutualistic pollination systems.     

Herbivory and seedling performance in a fragmented temperate forest of Chile

Forest fragmentation alters plant-animal interactions, including herbivory. Relying manipulative experiments, we test if the reduction in insect herbivory associated with forest fragmentation translates into increased seedling growth and survival of three tree common species (Aristotelia chilensis, Cryptocarya alba and Persea lingue) in forest fragments and continuous forests in coastal Maulino forest, central Chile. Furthermore, we test if after protecting seedlings from herbivorous insects, plant performance is increased regardless of forest fragmentation. Nursery grown seedlings were transplanted into four forest fragments and a continuous forest during 2002. Insects, important herbivores in this forest, were excluded from half the seedlings by repeated applications of insecticides. Compared to continuous forests, in forest fragments, herbivory was reduced in all three species, seedling growth was greater in A. chilensis and C. alba but not in P. lingue, and survivorship was unaffected by herbivory or fragmentation in all three species. Protecting seedlings from insects reduced herbivory in the continuous forest to similar levels attained in the forest fragments. No change in herbivory results from by protecting seedlings in forest fragments. These results confirm that insects are important herbivores in the Maulino forest and also support the hypothesis that fragmentation can have strong indirect effects on plant communities as mediated through trophic interactions.     

The sensitivity of the pseudomurein-containing genus Methanobacterium to inhibitors of murein synthesis

The sensitivity to inhibitors of various steps of murein synthesis was studied with six strains of methanogenic bacteria. Four of the strains belong to the genus Methanobacterium, which contains pseudomurein in its cell walls. This polymer-as well as murein-is not present in the two control organisms, Methanosarcina barkeri and Methanospirillum hungatii, which were found to be resistant to all inhibitors of murein synthesis. The four strains of Methanobacterium were resistant to the antibiotics fosfomycin, D-cycloserine, vancomycin, penicillin G and cephalosporin C, all of which inhibit the synthesis or function of the peptide subunits of murein. On the other hand, the four strains were susceptible to bacitracin, nisin, gardimycin and enduracidin. It is therefore assumed that the biosynthesis of murein and pseudomurein, respectively, may have some reactions of the so-called lipid cycle and the polymerization of the heteroglycan strands in common.     

Survival of the Black bog ant (<Emphasis Type="Italic">Formica transkaucasica</Emphasis> Nasanov) in relation to the fragmentation of its habitat

In the past, extensive areas in Drenthe (The Netherlands) were covered by peat bogs and wet heath lands, but nowadays only relatively small fragments are left. During the second half of the 20th century the quality of these fragments decreased, due to lowering of the water table and the input of nutrients. These factors will have a negative effect on the survival of species which are adapted to these stable type of biotopes, like the Black bog ant. The distribution pattern was analysed within a study area of 750 km², in order to find out if this species will survive in a landscape where its habitat is severely fragmented. Using multiple logistic regression analysis it appears that size and quality of the habitat patches, as well as openness of the environment, contribute significantly to patch-occupancy. No correlation was found between the probability of a patch being occupied and its distance to the nearest occupied patch. It appears that the spatial cohesion of local populations by means of flying queens is weak or absent on the scale of the study area. Only in parts of the area, where the distance between habitat patches is less than 3 km in open field, a habitat network may still exist. However, with ongoing habitat loss a threshold will be passed and the species will ultimately become extinct.