Spatial Replication Research Articles

Reducing Sample Quantity and Maintaining High Prediction Quality of Grassland Biomass Properties with near Infrared Reflectance Spectroscopy

Broad-scale ecological research often suffers from insufficient spatial and temporal replication. Near infrared (NIR) reflectance spectroscopy offers the opportunity for rapid and cheap measurements of many chemical constituents in organic materials. However, standard NIR instrumentation requires a certain amount of sample material which strongly restricts the fields of application for the NIR technique. Therefore, we tested if reliable predictions from NIR spectra can be obtained utilising a device that reduces the amount of required sample material by more than 95% compared to standard equipment. For large and small sample quantities, we present two sets of calibration models for C, N, P, K, Ca and Mg concentrations as well as fibre components such as neutral detergent fibre (NDF), acid detergent fibre (ADF) and acid detergent lignin (ADL) in above-ground grassland community biomass. Coefficients of multiple determination ( R2) of calibration models based on spectral data derived from standard equipment for C, N, P, K, Ca, Mg, NDF, ADF and ADL were 0.78, 0.98, 0.78, 0.92, 0.87, 0.89, 0.95, 0.94 and 0.87, respectively. Except for C and P, the ratio of standard deviation of the reference values to the standard error of cross validation and ratio of performance deviation indicated acceptable to high model precision. The application of NIR spectroscopy for C and P measurements was limited due to low variation in concentrations and/or low concentrations in the analysed above-ground grassland biomass. As compared to the deviation of duplicate reference measurements, the standard error of prediction was less than two times higher for C, N, NDF, ADF, ADL and K and up to three times higher for P, Ca and Mg. Prediction models based on the spectral data recorded with a small sample cell (volume of sample material less than 0.25 cm3) were of similar precision. The significant reduction of sample material required for NIR analysis and, at the same time, maintaining (high) precision of calibration models is an important advance towards the wider adoption of the NIR technique in ecological research.

Accounting for the effects of biological variability and temporal autocorrelation in assessing the preservation of species abundance

Quantifying the effects of taphonomic processes on species abundances in time-averaged death assemblages (DAs) is pivotal for paleoecological inference. However, fidelity estimates based on conventional “live-dead” comparisons are fundamentally ambiguous: (1) data on living assemblages (LAs) are based on a very short period of sampling and thus do not account for biological variability in the LA, (2) LAs are sampled at the same time as the DA and thus do not necessarily reflect past LAs that contributed to the DA, (3) compositions of LAs and DAs can be autocorrelated owing to shared cohorts, and (4) fidelity estimates are cross-scale estimates because DAs are time-averaged and LAs are not. Some portion of raw (total) live-dead (LD) variation in species composition thus arises from incomplete sampling of LAs and from biological temporal variation among LAs (together = premortem component of LD variation), as contrasted withnewvariation created by interspecific variation in population turnover and preservation rates and by the time-averaging of skeletal input (together = postmortem component of LD variation). To tackle these problems, we introduce a modified test for homogeneity of multivariate dispersions (HMD) in order to (1) account for temporal autocorrelation in composition between LAs and DAs and (2) decompose total LD compositional variation into premortem and postmortem components, and we use simulations to evaluate the contribution of within-habitat time-averaging on the postmortem component. Applying this approach to 31 marine molluscan data sets, each consisting of spatial replicates of LAs and DAs in a single habitat, we find that total LD variation is driven largely by variation among LAs. However, genuinely postmortem processes have significant effects on composition in 25–65% of data sets (depending on the metric) when the effects of temporal autocorrelation are taken into account using HMD. Had we ignored the effects of autocorrelation, the effects of postmortem processes would have been negligible, inflating the similarity between LAs and DAs. Simulations show that within-habitat time-averaging does not increase total LD variation to a large degree—it increases total LD variation mainly via increasing species richness, and decreases total LD variation by reducing dispersion among DAs. The postmortem component of LD variation thus arises from differential turnover and preservation and multi-habitat time-averaging. Moreover, postmortem processes have less effect on the compositions of DAs in habitats characterized by high variability among LAs than they have on DAs in temporally stable habitats, a previously unrecognized first-order factor in estimating postmortem sources of compositional variation in DAs.

Relative importance of biotic and abiotic soil components to plant growth and insect herbivore population dynamics.

BackgroundPlants are affected by several aspects of the soil, which have the potential to exert cascading effects on the performance of herbivorous insects. The effects of biotic and abiotic soil characteristics have however mostly been investigated in isolation, leaving their relative importance largely unexplored. Such is the case for the dune grass Ammophila, whose decline under decreasing sand accretion is argued to be caused by either biotic or abiotic soil properties.Methodology/Principal FindingsBy manipulating dune soils from three different regions, we decoupled the contributions of region, the abiotic and biotic soil component to the variation in characteristics of Ammophila arenaria seedlings and Schizaphis rufula aphid populations. Root mass fraction and total dry biomass of plants were affected by soil biota, although the latter effect was not consistent across regions. None of the measured plant properties were significantly affected by the abiotic soil component. Aphid population characteristics all differed between regions, irrespective of whether soil biota were present or absent. Hence these effects were due to differences in abiotic soil properties between regions. Although several chemical properties of the soil mixtures were measured, none of these were consistent with results for plant or aphid traits.Conclusions/SignificancePlants were affected more strongly by soil biota than by abiotic soil properties, whereas the opposite was true for aphids. Our results thus demonstrate that the relative importance of the abiotic and biotic component of soils can differ for plants and their herbivores. The fact that not all effects of soil properties could be detected across regions moreover emphasizes the need for spatial replication in order to make sound conclusions about the generality of aboveground-belowground interactions.

Spider Assemblages across Elevational and Latitudinal Gradients in the Yukon Territory, Canada

Arthropod assemblages in the Arctic are set for substantial changes in response to climate change, yet we know little about the ecological structure of many groups in the North. We tested the effects of elevation and latitude on northern spider assemblages by sampling along nine mountains across three latitudes in the Yukon Territory, Canada. Spiders were collected in 216 pitfall traps placed at four elevations along each of the nine mountains, representing 36 sites sampled across three latitudes (i.e., distinct mountain ranges). We collected 1954 individuals representing 89 species, 57 genera, and 12 families of spiders. Using nested ANOVAs, we found significant main effects of latitude, elevation, and an interaction of the two factors on species richness and abundance. Using MRPP and NMS ordination, we also found significant effects of latitude and mountain on species composition, but within each of the three latitudes, only elevation produced significant effects. Our study suggests that changes along spatial gradients associated with changes in habitat can have significant effects on the structure of spider assemblages, but responses vary among mountain ranges. We show that within a given mountain range, individual mountains may be used as spatial replicates for studies about northern arthropod assemblages.

Tigers on trails: occupancy modeling for cluster sampling

Occupancy modeling focuses on inference about the distribution of organisms over space, using temporal or spatial replication to allow inference about the detection process. Inference based on spatial replication strictly requires that replicates be selected randomly and with replacement, but the importance of these design requirements is not well understood. This paper focuses on an increasingly popular sampling design based on spatial replicates that are not selected randomly and that are expected to exhibit Markovian dependence. We develop two new occupancy models for data collected under this sort of design, one based on an underlying Markov model for spatial dependence and the other based on a trap response model with Markovian detections. We then simulated data under the model for Markovian spatial dependence and fit the data to standard occupancy models and to the two new models. Bias of occupancy estimates was substantial for the standard models, smaller for the new trap response model, and negligible for the new spatial process model. We also fit these models to data from a large-scale tiger occupancy survey recently conducted in Karnataka State, southwestern India. In addition to providing evidence of a positive relationship between tiger occupancy and habitat, model selection statistics and estimates strongly supported the use of the model with Markovian spatial dependence. This new model provides another tool for the decomposition of the detection process, which is sometimes needed for proper estimation and which may also permit interesting biological inferences. In addition to designs employing spatial replication, we note the likely existence of temporal Markovian dependence in many designs using temporal replication. The models developed here will be useful either directly, or with minor extensions, for these designs as well. We believe that these new models represent important additions to the suite of modeling tools now available for occupancy estimation in conservation monitoring. More generally, this work represents a contribution to the topic of cluster sampling for situations in which there is a need for specific modeling (e.g., reflecting dependence) for the distribution of the variable(s) of interest among subunits.

The Effects of Winter Burning and Grazing on Resources and Survival of Texas Horned Lizards in a Thornscrub Ecosystem

ABSTRACT The ecological effects of land‐use practices on reptiles, especially endangered or threatened species, are of conservation and scientific interest. We describe the effects of rotational livestock grazing and prescribed winter burning on resources and survival of the Texas horned lizard (Phrynosoma cornutum) during the summers of 1998 to 2001 in southern Texas, USA. We evaluated survival rates of Texas horned lizards (n = 111) on 6 study sites encompassing 5 different burning and grazing treatments. We also measured indices of cover (i.e., vegetation) and food abundance (i.e., harvester ants [Pogonomyrmex rugosus]). We telemetered and relocated adult lizards daily. We divided the study into 2 seasons, spring (15 Apr–30 Jun) and summer (1 Jul–15 Aug), corresponding to the relative activity of horned lizards. Winter burning provided an increase in food resources and led to increased survival of Texas horned lizards in the second growing season after fire, but grazing‐induced changes in vegetation cover reduced survival, likely by increasing lizard vulnerability. Fire and grazing reduced litter and increased bare ground and forb cover but did not affect woody vegetation. Ant activity was greater in burned sites and varied with grazing level, season, and year. Summer survival functions of horned lizards varied by burning treatment, with higher survival observed on burned sites in the second year after burning. Survival rates were ordered from highest in ungrazed sites to lowest in heavily grazed sites. We recognize the limitations of our work resulting from a lack of spatial replication of treatments. However, our mensurative study provides fertile ground for future hypothesis testing regarding the effects of land management on shrubland and grassland reptiles. We propose that future studies focus on the population consequences of variation in burn frequency, burn timing, and grazing intensity.

Dynamic macroecology on ecological time‐scales

ABSTRACTAim The discipline of macroecology is increasingly being regarded as an effective vehicle for the evaluation of recent population‐ to ecosystem‐level responses to widespread human and environmental influences. However, due to the prevalent use of time‐averaged and cumulative data in macroecological analyses, the majority of the patterns that emerge from research in this field can be regarded as static. Here we review the application of dynamic macroecological analyses to changes in relationships between macroecological variables on seasonal to decadal scales. We illustrate the strength of this perspective for documenting changing patterns and testing hypotheses related to these dynamics on ecological time‐scales.Location Studies were compiled and reviewed from terrestrial and aquatic ecosystems.Methods We review examples of temporal changes in macroecological patterns driven by recent anthropogenic influences and environmental change.Results The dynamic nature of macroecological patterns on ecological time‐scales has been revealed in recent years across a wide range of ecosystems, largely through the development, maintenance and analysis of biotic and environmental monitoring time series. The resultant analyses complement examinations of dynamics over evolutionary time and have similarly revealed that static portrayals can conceal important temporal dynamics that underlie the patterns of interest. As a consequence, static depictions, resting as they do on comparative analyses in which the validity of space‐for‐time substitutions is assumed, may be of limited use for testing hypotheses related to the mechanisms underlying the patterns revealed and, by extension, the development of reliable predictions of future states.Main conclusions Recent dynamic macroecological analyses have demonstrated the utility of combined spatial and temporal replication, and have contributed to hypothesis testing related to the mechanistic processes underlying changes in macroecological patterns on ecological time‐scales. We suggest four specific avenues of future research to further the development and application of temporal approaches on similar time‐scales within the field of macroecology.

Bird population density estimated from acoustic signals

Summary1.Many animal species are detected primarily by sound. Although songs, calls and other sounds are often used for population assessment, as in bird point counts and hydrophone surveys of cetaceans, there are few rigorous methods for estimating population density from acoustic data.2.The problem has several parts – distinguishing individuals, adjusting for individuals that are missed, and adjusting for the area sampled. Spatially explicit capture–recapture (SECR) is a statistical methodology that addresses jointly the second and third parts of the problem. We have extended SECR to use uncalibrated information from acoustic signals on the distance to each source.3.We applied this extension of SECR to data from an acoustic survey of ovenbirdSeiurus aurocapilladensity in an eastern US deciduous forest with multiple four‐microphone arrays. We modelled average power from spectrograms of ovenbird songs measured within a window of 0·7 s duration and frequencies between 4200 and 5200 Hz.4.The resulting estimates of the density of singing males (0·19 ha−1SE 0·03 ha−1) were consistent with estimates of the adult male population density from mist‐netting (0·36 ha−1SE 0·12 ha−1). The fitted model predicts sound attenuation of 0·11 dB m−1(SE 0·01 dB m−1) in excess of losses from spherical spreading.5.Synthesis and applications.Our method for estimating animal population density from acoustic signals fills a gap in the census methods available for visually cryptic but vocal taxa, including many species of bird and cetacean. The necessary equipment is simple and readily available; as few as two microphones may provide adequate estimates, given spatial replication. The method requires that individuals detected at the same place are acoustically distinguishable and all individuals vocalize during the recording interval, or that theper capitarate of vocalization is known. We believe these requirements can be met, with suitable field methods, for a significant number of songbird species.

Radial growth response of white oak to climate in eastern North America

Predicting forest responses to climate change requires an understanding of the cause–effect relationships linking climate to tree growth. Dendroecological analyses across sites that span climate gradients provide one means of characterizing such relationships. Dendroecological analyses for white oak ( Quercus alba L.) at 149 sites spanning the species range in eastern North America identified spatially replicated growth–climate associations. Early growing season site water balance variables for the year of annual ring formation had the strongest, most spatially replicated associations with growth. There was little evidence of phenological variation of these associations related to the latitudinal temperature gradient. Most spatial variation in growth–climate associations was along an east-to-west precipitation gradient. Radial growth was most strongly correlated with site water balance at sites in the northwest quadrant of the range, characterized by continental climate and high interannual variability in precipitation. There was little evidence that dormant season temperature affects white oak growth, even at the northern range limit. Correlations with dormant season precipitation were common in the northwest part of the range, where winter soil water recharge may be more variable. Spatial replication was a useful criterion for distinguishing growth–climate correlations that reflect cause–effect relationships.

A Reliable Architecture for Parallel Implementations of the Advanced Encryption Standard

This paper presents an on-line self-test architecture for hardware implementation of the Advanced Encryption Standard (AES). The solution exploits the inherent spatial replications of a parallel architecture for implementing functional redundancy at low cost. We show that the solution is very effective for on-line fault detection while keeping the area overhead very low. Moreover, the architectural modification for on-line test does not weaken the device with respect to side-channel attacks based on power analysis.

Teste do efeito de borda na predação de ninhos naturais e artificiais no Cerrado

The Cerrado is still one of the most important ecosystems in Brazil, even though more than 50% of its area has been altered or converted to pastureland and plantations. Despite its intense degradation, few ecological processes that might affect its biodiversity have been evaluated. The goal of this study was to test the edge effect on the predation rates at natural and artificial nests, at the Ecological Station of Aguas Emendadas, Federal District, Brazil. Natural nests were found and monitored every three to four days from September to December of 2004 in the interior and at the edge of the reserve. Artificial nests were placed at four distances from the edge (0, 500, 1000 and 2000 m) in three spatial replicates in September and again in December of 2004. Each nest received one Japanese Quail and one plasticine egg and was monitored every five days, for 15 days. There was no difference between the rates of predation either in the natural nests or in the artificial nests between treatments. For one bird species, Elaenia chiriquensis (Lawrence, 1865), Tyrannidae, daily survival rates in the incubation and in the hatchling period had opposite values between the edge and the interior. Marks on plasticine eggs suggest that birds are the main predators. Estimates of the abundance of two potential nest predators, Cyanocorax cristatellus (Temminck, 1823), Corvidae and Canis familiaris (Linnaeus, 1758), Canidae, revealed no relationship with distance to the edge, nor with predation rates. Brood parasitism of natural nests was similar between the interior (0%) and the edge (3.8% of the nests). The results described here do not support the edge effect hypothesis for nest predation rates on either natural or artificial nests, nor for brood parasitism rates.

Predictive models of weed population dynamics

SummaryMany of the challenges faced by weed ecologists can be met only by the capability to predict the responses of weed populations to changes in their environment or management. In spite of this, a review of papers published in Weed Research suggests that weed ecologists are remarkably reluctant to produce detailed, quantitative predictions. This may result from uncertainty in the accuracy of predictions and indeed, a variety of reasons have been put forward to suggest that the potential utility of weed models may be limited in this regard. In this study, we review the applications to which weed models have been put. Focusing on predictive population modelling, we highlight several limitations that can lead to failures of this approach and we discuss the likely prospects for weed population modelling. We make three points regarding the future of weed modelling. First, owing to prohibitive data requirements, the development of highly mechanistic models that attempt to make detailed predictions of weed population numbers is unlikely to be very successful. Second, data collection for developing weed models needs to be rethought. Weed models are most commonly compromised by a lack of spatial and temporal replication, preventing modellers from measuring parameter variability and error effectively and limiting assessments of model uncertainty. Finally, the utility of models needs to be better appreciated; models are key tools in making long range predictions of how management will affect weed populations, but, we estimate, they are used in only a small fraction of studies. Without the further development of models for weed population dynamics, our ability to predict long‐term dynamics will be restricted.

Optimising sample volume and replicates using the Bou-Rouch method for the rapid assessment of hyporheic fauna

Despite the widespread use of the Bou-Rouch method in obtaining hyporheic samples in ecological studies, problems persist in comparing data because of differences in volume and number of samples taken in various studies. Towards standardising this method, we conducted a study in the Delour River, Ireland, between March and September 2006. The hyporheic habitat was divided into three lateral zones: the flowing stream (SS); the stream–terrestrial ecotone (EC); and the terrestrial margin (TM). Between 3 and 6 random insertions of the stand pipe were made into the hyporheic habitat at 0.2 m and 0.5 m in each zone. Ten consecutive 1-L samples were pumped at each point to determine an optimum sample volume and replicates needed to estimate taxon richness and abundance. The optimum sample combinations range between 6 and 15 3- to 10-L samples depending on depth and habitat zone. Our results show that both spatial and temporal scales are important factors in considering the optimum combinations of sample volume and number of independent spatial replicates needed to sample stream hyporheos, with the zone closer to the flowing stream requiring many small-volume samples whereas areas further away towards the alluvial groundwater need larger volumes with small numbers of replicates.

Experimental evidence that competition and habitat use shape the individual fitness surface

A key prediction made by theories of density-dependent competition is that resource overlap should increase the intensity of competition. By extension, we can predict that competition should lead to density-dependent natural selection. I studied natural selection on limb length and body size in a total of seven populations of Anolis sagrei over 3 years in the Bahamas. Experimental manipulations of population density on small off-shore cays revealed that the strength of natural selection on body size increased with density, suggesting that density-dependent intraspecific competition drives natural selection. At low density, reduced competition revealed significant selection on limb length driven by changes in perch diameter, indicating that selection favoured a match between morphology and habitat. The role habitat played in shaping selection was further illuminated by inter-annual changes in vegetation structure stemming from variation in precipitation among years. Thus, changes in both the intensity of competition across spatial replicates, and in resource availability through time, revealed changes in the targets of natural selection. Results provide empirical support for the long-standing hypothesis that density-dependent natural selection shapes the fitness surface of Greater Antilles anoles.

SPATIAL DENSITY DEPENDENCE SCALES UP BUT DOES NOT PRODUCE TEMPORAL DENSITY DEPENDENCE IN A REEF FISH.

Field experiments provide rigorous tests of ecological hypotheses but are typically of short duration and use small spatial replicates. We assessed empirically whether the results of experiments testing for density dependence applied at larger spatial domains and explained temporal population dynamics. We studied a small coral reef fish, the goldspot goby (Gnatholepis thompsoni), in the Bahamas. We assessed the effects of interactions with conspecifics and with an ecologically similar species, the bridled goby (Coryphopterus glaucofraenum). Two density manipulations on small reef patches revealed that goldspot goby mortality over one month increased as conspecifics became crowded. On five large natural reefs, we correlated the initial year-class density of both species (annual larval settlement) with the subsequent decline of goldspot goby year-classes for five years. Mortality was correlated with conspecific density among reefs for all years, but not among years for all reefs. Thus, spatial density dependence in mortality scaled up qualitatively from small patches to entire reefs but was not associated with temporal density dependence. Our results support the conclusion that field experiments may be extrapolated to larger spatial domains with care, but that using small spatial comparisons to predict temporal responses is difficult without knowing the underlying biological mechanisms.

Optimal denoising in redundant representations.

Image denoising methods are often designed to minimize mean-squared error (MSE) within the subbands of a multiscale decomposition. However, most high-quality denoising results have been obtained with overcomplete representations, for which minimization of MSE in the subband domain does not guarantee optimal MSE performance in the image domain. We prove that, despite this suboptimality, the expected image-domain MSE resulting from applying estimators to subbands that are made redundant through spatial replication of basis functions (e.g., cycle spinning) is always less than or equal to that resulting from applying the same estimators to the original nonredundant representation. In addition, we show that it is possible to further exploit overcompleteness by jointly optimizing the subband estimators for image-domain MSE. We develop an extended version of Stein's unbiased risk estimate (SURE) that allows us to perform this optimization adaptively, for each observed noisy image. We demonstrate this methodology using a new class of estimator formed from linear combinations of localized "bump" functions that are applied either pointwise or on local neighborhoods of subband coefficients. We show through simulations that the performance of these estimators applied to overcomplete subbands and optimized for image-domain MSE is substantially better than that obtained when they are optimized within each subband. This performance is, in turn, substantially better than that obtained when they are optimized for use on a nonredundant representation.

High speed fabrication of aluminum nanostructures with 10 nm spatial resolution byelectrochemical replication

A high fidelity electrochemical replication technique for the rapid fabrication of Alnanostructures with 10 nm lateral resolution has been successfully demonstrated. Aluminumis electrodeposited onto a lithographically patterned Si master using a non-aqueous organichydride bath of aluminum chloride and lithium aluminum hydride at room temperature.Chemical pretreatment of the Si surface allows a clean detachment of the replicated Al foilfrom the master, permitting its repetitive use for mass replication. This high throughputtechnique opens up new possibilities in the fabrication of Al-related nanostructures,including the growth of long range ordered anodic alumina nanochannel arrays.

A comparison of the relative contributions of temporal and spatial variation in the density of drifting invertebrates in a Dorset (U.K.) chalk stream

Summary1. Invertebrate drift is commonly investigated in streams, with the majority of studies focussed on temporal (typically diel) variation. In comparison, few studies have investigated spatial variation in drift and there is little consensus among them. We tested the hypothesis that spatial variation in invertebrate drift is as important as temporal variation.2. The density of drifting invertebrates in a chalk stream was sampled using an array of nets arranged to determine vertical, lateral and longitudinal variation. Samples were collected at dawn, during the day, at dusk and by night, on four separate monthly occasions. Insecta and Crustacea were analysed separately to identify the effect of differing life history strategies. The density of drifting debris was also recorded, to act as a null model.3. Time of day and vertical position together explained the majority of the variance in invertebrate drift (79% for Insecta and 97% for Crustacea), with drift densities higher at dusk and night, and nearer the stream bed. Independently, time of day (38%, Insecta; 52%, Crustacea) and vertical position (41%, Insecta; 45%, Crustacea) explained a similar amount of the observed variance. Month explained some of the variance in insect drift (9%) but none for Crustacea.4. Variation in the density of drifting debris showed little in common with invertebrate drift. There was little variation associated with time of day and only 27% of the observed variation in debris could be explained by the factors investigated here, with month explaining the largest proportion (20%). We suggest the difference in drifting debris and invertebrates provides further evidence for a strong behavioural component in invertebrate drift.5. Spatial variation in invertebrate drift can be of the same order of magnitude as the much‐described diel temporal variation. The extent of this spatial variation poses problems when attempting to quantify invertebrate drift and we recommend that spatial replication should be incorporated into drift studies.

ROLES OF SURVIVAL AND DISPERSAL IN REINTRODUCTION SUCCESS OF GRIFFON VULTURE (GYPS FULVUS)

The success of reintroduction programs greatly depends on the amount of mortality and dispersal of the released individuals. Although local environmental pressures are likely to play an important role in these processes, they have rarely been investigated because of the lack of spatial replicates of reintroduction. In the present study, we analyzed a 25-year data set encompassing 272 individuals released in five reintroduction programs of Griffon Vultures (Gyps fulvus) in France to examine the respective roles of survival and dispersal in program successes and failures. We use recent developments in multi-strata capture-recapture models to take into account tag loss in survival estimates and to consider and estimate dispersal among release areas. We also examined the effects of sex, age, time, area, and release status on survival, and we tested whether dispersal patterns among release areas were consistent with habitat selection theories. Results indicated that the survival of released adults was reduced during the first year after release, with no difference between sexes. Taking into account local observations only, we found that early survival rates varied across sites. However when we distinguished dispersal from mortality, early survival rates became equal across release sites. It thus appears that among reintroduction programs difference in failure and success was due to differential dispersal among release sites. We revealed asymmetrical patterns of dispersal due to conspecific attraction: dispersers selected the closest and the largest population. We showed that mortality can be homogeneous from one program to another while, on the contrary, dispersal is highly dependent on the matrix of established populations. Dispersal behavior is thus of major interest for metapopulation restoration and should be taken into account in planning reintroduction designs.

Cellular differentiation in moss protonemata: a morphological and experimental study.

Previous studies of protonemal morphogenesis in mosses have focused on the cytoskeletal basis of tip growth and the production of asexual propagules. This study provides the first comprehensive description of the differentiation of caulonemata and rhizoids, which share the same cytology, and the roles of the cytoskeleton in organelle shaping and spatial arrangement. Light and electron microscope observations were carried out on in vitro cultured and wild protonemata from over 200 moss species. Oryzalin and cytochalasin D were used to investigate the role of the cytoskeleton in the cytological organization of fully differentiated protonemal cells; time-lapse photography was employed to monitor organelle positions. The onset of differentiation in initially highly vacuolate subapical cells is marked by the appearance of tubular endoplasmic reticulum (ER) profiles with crystalline inclusions, closely followed by an increase in rough endoplasmic reticulum (RER). The tonoplast disintegrates and the original vacuole is replaced by a population of vesicles and small vacuoles originating de novo from RER. The cytoplasm then becomes distributed throughout the cell lumen, an event closely followed by the appearance of endoplasmic microtubules (MTs) in association with sheets of ER, stacks of vesicles that subsequently disperse, elongate mitochondria and chloroplasts and long tubular extensions at both poles of the nucleus. The production of large vesicles by previously inactive dictysomes coincides with the deposition of additional cell wall layers. At maturity, the numbers of endoplasmic microtubules decline, dictyosomes become inactive and the ER is predominantly smooth. Fully developed cells remain largely unaffected by cytochalasin; oryzalin elicits profound cytological changes. Both inhibitors elicit the formation of giant plastids. The plastids and other organelles in fully developed cells are largely stationary. Differentiation of caulonemata and rhizoids involves a remarkable series of cytological changes, some of which closely recall major events in sieve element ontogeny in tracheophytes. The cytology of fully differentiated cells is remarkably similar to that of moss food-conducting cells and, in both, is dependent on an intact microtubule cytoskeleton. The disappearance of the major vacuolar apparatus is probably related to the function of caulonema and rhizoids in solute transport. Failure of fully differentiated caulonema and rhizoid cells to regenerate is attributed to a combination of endo-reduplication and irreversible tonoplast fragmentation. The formation of giant plastids, most likely by fusion, following both oryzalin and cytochalasin treatments, suggests key roles for both microtubules and microfilaments in the spatial arrangement and replication of plastids.