Nonrandom movement behavior at habitat boundaries in two butterfly species: implications for dispersal

We observed meadow brown (Maniola jurtina) and gatekeeper (Pyronia tithonus) butterflies at habitat boundaries and observed spontaneous movements out of suitable habitat in order to investigate such movements in relation to dispersal. We found that butterflies of both species were aware of the position of a highly permeable habitat boundary without needing to cross it. Nevertheless, a considerable proportion of butterflies close to the boundary left their habitat (25-43%). Butterflies that crossed the boundary, and moved substantial distances into unsuitable habitat (up to 350 m in M. jurtina and 70 m in P. tithonus), usually returned to their original habitat patch (98-100%). Movement trajectories, at least in M. jurtina, were significantly different from, and more directed and systematic than, a correlated random walk. Approximately 70-80% of spontaneous movements into unsuitable habitat in both species were "foray" loops comparable to those described in mammals and birds. We conclude that, since migrants seemed to have considerable control over leaving their patch and over their subsequent movement trajectories, chance encounter rates with habitat boundaries, and indeed habitat leaving rates, might be less crucial in determining dispersal rates than is usually assumed. In addition, random dispersal trajectories should not be taken for granted in population or evolution models.     

Statistical selection of perimeter-area models for patch mosaics in multiscale landscape analysis

This paper presents a statistical method for detecting distinct scales of pattern for mosaics of irregular patches, by means of perimeter–area relationships. Krummel et al. (1987) were the first to develop a method for detecting different scaling domains in a landscape of irregular patches, but this method requires investigator judgment and is not completely satisfying. Grossi et al. (2001) suggested a modification of Krummel's method in order to detect objectively the change points between different scaling domains. Their procedure is based on the selection of the best piecewise linear regression model using a set of statistical tests. Even though the change points were estimated, the null distributions used for testing purposes were those appropriate for known change points. The present paper investigates the effect that estimating the change points has on the underlying distribution theory. The procedure we suggest is based on the selection of the best piecewise linear regression model using a likelihood ratio (LR) test. Each segment of the piecewise linear model corresponds to a fractal domain. Breakpoints between different segments are unknown, so the piecewise linear models are non-linear. In this case, the frequency distribution of the LR statistic cannot be approximated by a chi-squared distribution. Instead, Monte Carlo simulation is used to obtain an empirical null distribution of the LR statistic. The suggested method is applied to three patch types (CORINE biotopes) located in the Val Baganza watershed of Italy.     

General methods for sensitivity analysis of equilibrium dynamics in patch occupancy models

Sensitivity analysis is a useful tool for the study of ecological models that has many potential applications for patch occupancy modeling. Drawing from the rich foundation of existing methods for Markov chain models, I demonstrate new methods for sensitivity analysis of the equilibrium state dynamics of occupancy models. Estimates from three previous studies are used to illustrate the utility of the sensitivity calculations: a joint occupancy model for a prey species, its predators, and habitat used by both; occurrence dynamics from a well-known metapopulation study of three butterfly species; and Golden Eagle occupancy and reproductive dynamics. I show how to deal efficiently with multistate models and how to calculate sensitivities involving derived state variables and lower-level parameters. In addition, I extend methods to incorporate environmental variation by allowing for spatial and temporal variability in transition probabilities. The approach used here is concise and general and can fully account for environmental variability in transition parameters. The methods can be used to improve inferences in occupancy studies by quantifying the effects of underlying parameters, aiding prediction of future system states, and identifying priorities for sampling effort.     

Pattern formation and individual-based models: The importance of understanding individual-based movement

This paper demonstrates that while pattern formation can stabilize individual-based models of predator–prey systems, the same individual-based models also allow for stabilization by alternate mechanisms, particularly localized consumption or diffusion limitation. The movement rules of the simulation are the critical feature which determines which of these mechanisms stabilizes any particular predator–prey individual-based model. In particular, systems from well-connected subpopulations, in each of which a predator can attack any prey, generally exhibit stabilization by pattern formation. In contrast, when restricted movement within a (sub-)population limits the ability of predators to consume prey, localized consumption or diffusion limitation can stabilize the system. Thus while the conclusions from differential equations on the role of pattern formation for stability may apply to discrete and noisy systems, it will take a detailed understanding of movement and scales of interaction to examine the role of pattern formation in real systems. Additionally, it will be important to link an understanding of both foraging and inter-patch movement, since by analogy to the models, both would be critical for understanding how real systems are stabilized by being discrete and spatial.     

Group navigation and the "many-wrongs principle" in models of animal movement

Traditional studies of animal navigation over both long and short distances have usually considered the orientation ability of the individual only, without reference to the implications of group membership. However, recent work has suggested that being in a group can significantly improve the ability of an individual to align toward and reach a target direction or point, even when all group members have limited navigational ability and there are no leaders. This effect is known as the "many-wrongs principle" since the large number of individual navigational errors across the group are suppressed by interactions and group cohesion. In this paper, we simulate the many-wrongs principle using a simple individual-based model of movement based on a biased random walk that includes group interactions. We study the ability of the group as a whole to reach a target given different levels of individual navigation error, group size, interaction radius, and environmental turbulence. In scenarios with low levels of environmental turbulence, simulation results demonstrate a navigational benefit from group membership, particularly for small group sizes. In contrast, when movement takes place in a highly turbulent environment, simulation results suggest that the best strategy is to navigate as individuals rather than as a group.     

Elephants as agents of habitat creation for small vertebrates at the patch scale

Ecologists increasingly recognize the ability of certain species to influence ecological processes by engineering the physical environment, but efforts to develop a predictive understanding of this phenomenon are in their early stages. While many believe that the landscape-scale effects of ecosystem engineers will be to increase habitat diversity and therefore the abundance and richness of other species, few generalities exist about the effects of engineering at the scale of the engineered patch. According to one hypothesis, activities that increase structural habitat complexity within engineered patches will have positive effects on the abundance or diversity of other organisms. Here I show that, by damaging trees and increasing their structural complexity, browsing elephants create refuges used by a common arboreal lizard. Observational surveys and a lizard transplant experiment revealed that lizards preferentially occupy trees with real or simulated elephant damage. A second experiment showed that lizards vacate trees when elephant-engineered refuges are removed. Furthermore, local lizard densities increased with (and may be constrained by) local densities of elephant-damaged trees. This facilitative effect of elephants upon lizards via patch-scale habitat modification runs contrary to previously documented negative effects of the entire ungulate guild on lizards at the landscape scale, suggesting that net indirect effects of large herbivores comprise opposing trophic and engineering interactions operating at different spatial scales. Such powerful megaherbivore-initiated interactions suggest that anthropogenic changes in large-mammal densities will have important cascading consequences for ecological communities.     

A new index for measuring perpendicularity of animal movements in relation to patch boundaries

We introduce a new index for measuring perpendicularity of animal movements with respect to a boundary (e.g., a habitat patch edge), and provide a computer algorithm for its calculation. Our index, η, improves on an approach that measures perpendicularity with respect to a fixed boundary direction. This is because η accounts for moment-to-moment trajectories relative to nearest-neighbor boundary attributes at the scale of an animal's movement. Our algorithm prp calculates η efficiently and accurately with both synthetic data and large telemetry datasets. In addition, we have included routines in prp which account for scenarios inherently problematic to perpendicularity estimators.     

Use of stochastic patch occupancy models in the California red-legged frog for Bayesian inference regarding past events and future persistence

Assessing causes of population decline is critically important to management of threatened species. Stochastic patch occupancy models (SPOMs) are popular tools for examining spatial and temporal dynamics of populations when presence–absence data in multiple habitat patches are available. We developed a Bayesian Markov chain method that extends existing SPOMs by focusing on past environmental changes that may have altered occupancy patterns prior to the beginning of data collection. Using occupancy data from 3 creeks, we applied the method to assess 2 hypothesized causes of population decline—in situ die-off and residual impact of past source population loss—in the California red-legged frog. Despite having no data for the 20–30 years between the hypothetical event leading to population decline and the first data collected, we were able to discriminate among hypotheses, finding evidence that in situ die-off increased in 2 of the creeks. Although the creeks had comparable numbers of occupied segments, owing to different extinction–colonization dynamics, our model predicted an 8-fold difference in persistence probabilities of their populations to 2030. Adding a source population led to a greater predicted persistence probability than did decreasing the in situ die-off, emphasizing that reversing the deleterious impacts of a disturbance may not be the most efficient management strategy. We expect our method will be useful for studying dynamics and evaluating management strategies of many species.     

Old endemics and new invaders: alternative strategies of passerines for living in the Australian environment

Life history parameters of ”old endemics” and ”new invaders” among the Australian passerines were compared, controlling for phylogeny by using current, phylogenetic research methods. Old endemics were found to have significantly smaller clutches and demonstrated significantly more cooperative breeding than the new invaders, whose fledging periods are significantly longer than those of the old endemics. We argue that breeding under conditions of a small annual food increment during the breeding season, as is probably the situtation in Australia, is possible only when clutch size is small, or while breeding cooperatively, or by extending the fledging period. Our data show that the old endemics use the first two options, while the new invaders employ the third. Received: 10 February 1999 / Received in revised form: 10 November 1999 / Accepted: 20 November 1999     

Contrasting diel patterns in vertical movement and locomotor activity of whale sharks at Ningaloo Reef

Activity patterns of animals often relate to environmental variables such as food availability and predation pressure. Technological advances are providing us with new tools to monitor and better understand these activity patterns. We used animal-attached data loggers recording acceleration and depth to compare activity patterns and vertical habitat use of whale sharks (Rhincodon typus) at Ningaloo Reef, Western Australia. Whale sharks showed a moderate reverse diel vertical migration but exhibited a clear crepuscular pattern in locomotory activity. Peak activity occurred at sunset, whereas vertical movement peaked prior to this. Typical ram surface filter feeding could be identified and occurred primarily during sunset and the first hours of night. At such times, direct observations indicated whale sharks were feeding on tropical krill swarms. Kinematic analysis of postural data and data from vertical movement suggests that whale sharks at Ningaloo spend ~8 min per day actively ram surface filter feeding. Considering the high biomass present in krill schools, it is estimated that whale sharks at Ningaloo have a similar energy intake as those at other aggregation sites. Diel patterns in activity and diving behaviour suggest that whale sharks have tuned their diving behaviour in anticipation of the formation of these high-density patches which appear to only be periodically, but predictably available at sunset. Our results confirm that diel patterns in vertical habitat selection and vertical movements do not necessarily reflect patterns in activity and foraging behaviour. Direct quantification of activity and behaviour is required in gaining accurate representation of diel activity patterns.     

Further studies on the pressure tolerance of deep-sea crustacea,with observations using a new high-pressure trap

A deep-sea benthic trap is described with which amphipods (Tmetonyx cicada) were both collected and observed at their ambient pressure of 134 atm. T. cicada collected with decompression from the same depth and locality were more sensitive to subsequent recompression than those amphipods brought to the surface at their ambient pressure. T. cicada from 2700 m experience irreversible injury during decompression to atmospheric pressure. The pressure tolerance of the deep-sea mysid Gnathophausia zoea was measured and compared with the tolerance of mid-water decapods and the shallow-water Crangon crangon. G. zoea became more sensitive to high pressure with prolonged exposure to atmospheric pressure. Deep sea animals exhibit a tolerance to high pressure related to their normal ambient pressure; sensitivity to decompression is also related to normal ambient pressure.     

Initial observations of the zooplankton microdistribution on the fringing coral reef at Aqaba (Jordan)

Diurnal and nocturnal zooplankton was sampled simultaneously at the surface and at 35 m depth on the outer slope of the fringing reef off the Jordanian coast in the Guff of Aqaba, Red Sea, over a period of 8 d during July 1982. Complementary surface sampling was carried out above the reef and in the open sea in the morning and afternoon for a period of 5 d. Five groups of organisms were classified according to changes in their vertical distribution in the water column which is analyzed and discussed. Zooplankters of the first group appear to be positively phototactic (Calanus robustior, Mecynocera clausi, Oithona helgolandica, Corycella rostrata). Those of the second group also seem to be positively phototactic, but only as regards average and low light (moonlight) intensities (Nannocalanus minor, Acartia negligens, Sagitta enflata, S. hexaptera, Thalia democratica, Oikopleura fusiformis, O. longicauda). The zooplankton of the third group displays a negative phototaxis (Oithona nana, calyptopis and furcilia euphausiid stages, pelecypods and other mollusc veligers). Species exhibiting uncertain behaviour or caught in low numbers and exhibiting no particular distribution are relegated to the fourth and the fifth groups. The horizontal microdistribution of the zooplankton community is not easily discernible: certain neritic or oceanic affinities can be identified, however, by comparing the WP2 standard net surface samples collected above the reef with those from the open sea. The importance of zooplankton communities is emphasized, and the necessity of elucidating the temporal and spatial migrations and exchanges between reef and planktonic organisms in order to gain more insight into their trophic relationship with the reef system.     

A new algorithm for estimating the parameters of the spatial generalized linear mixed models

Non-Gaussian spatial responses are usually modeled using a spatial generalized linear mixed model with location specific latent variables. The likelihood function of this model cannot usually be given in a closed form, thus the maximum likelihood approach is very challenging. So far, several numerical algorithms to solve the problem of calculating maximum likelihood estimates of this model have been presented. In this paper to estimate the parameters an approximate method is considered and a new algorithm is introduced that is much faster than existing algorithms but just as accurate. This is called the Approximate Expectation Maximization Gradient algorithm. The performance of the proposed algorithm and is illustrated with a simulation study and on a real data set.     

新时期我国区域经济地理学教学新体系建设

21世纪知识经济时代对高素质专业人才需求的迫切性、经济地域系统及区域开发问题的日益复杂性对区域经济地理学教学体系建设提出了新的要求.本文从课程体系的设置、教材的建设和教学改革几个方面,探讨了新时期我国区域经济地理学教学体系的建设问题.     

The effects of sand movement on intertidal seaweeds and selected invertebrates at Bound Rock,New Hampshire,USA

A seasonal study of sand movement and the benthic intertidal organisms at Bound Rock, New Hampshire, USA was conducted between November, 1973 and February, 1975. The site is subjected to irregular sand fluctuations, as well as diurnal, neapspring and major summer sand intrusions. The abundance and distribution of intertidal species was interrelated with the historical sand fluctuations at the area. For example, the lower limits of Mytilus edulis, Balanus balanoides and Porphyra umbilicalis approximated the highest summer sand elevations. Highly abraded rock surfaces in the lower intertidal zone were dominated by opportunistic annuals (e.g. Enteromorpha spp.) and perennial psammophytic or sand-loving seaweeds (e.g. Ahnfeltia plicata and Sphacelaria radicans). Overall, the intertidal seaweed populations at Bound Rock showed a lower number of perennials and fewer species than adjacent rocky shores. The low species diversity of seaweeds at the study site is attributable to unstable environmental conditions and a limited number of habitats. Even so, several psammophytic microhabitats are evident at Bound Rock depending upon the extent of sand burial and abrasion. The morphological and reproductive adaptations of several psammophytic species are discussed.Published with the approval of the Director of the New Hampshire Agricultural Experiment Station as Scientific Contribution No. 812.Scientific Contribution No. 53 of the Jackson Estuarine Laboratory.     

Assessment of metals in soil extracts and their uptake and movement within Tamarix nilotica at Lake Nasser banks,Egypt

This study aims to determine heavy-metal levels in soil from the banks of Lake Nasser, the ability of Tamarix nilotica to accumulate such metals from soil and hence its potential for phytoextraction. Soil and Tamarix samples were collected from the banks of four bights around Lake Nasser and analysed for Fe, Mn, Ca, Mg, Cr, Cu, Ni, Zn, Cd and Pb by atomic absorption spectrometry, whereas Na and K were measured by atomic emission spectrophotometry. Three different methods of extraction were used for the soil samples. Lead, copper and zinc were equally distributed between the exchangeable phase and Fe/Mn oxide-bound form, while other measured metals were mainly present in the Fe/Mn oxide fraction. With the exception of iron, all metals studied showed total concentrations within the geochemical background values. T. nilotica exhibited elevated concentrations of Na (36.2–48.5?mg?g^?1) and K (2.74–4.33?mg?g^?1) in stems, and relatively high concentrations of Pb, Cd and Co (0.39–1.03?µg?g^?1, 0.24–1.3?µg?g^?1 and 1.94–5.3?µg?g^?1, respectively) are found in plant leaves. Bioaccumulation factors of Na and K (9.3 and 12.63, respectively) were high in T. nilotica stems. While the bioaccumulation of Pb, Cd, Co and Ni (2870.1, 2035.4, 10.5 and 5313.2, respectively) was high in plant leaves, Fe, Mn, Ca and Mg were accumulated relatively equally in plant stems and leaves. T. nilotica was found to secrete high amounts of Na, Ca and K, in addition to small amounts of all accumulated metals except Cd and Cu. These secreted metals appeared as salt crystals (67.5% Na; 25.8% Ca; 5% Mg; 1.5% K and 0.16% trace and minor elements) on the plant surface. The concentrations of all the metals studied in T. nilotica were higher than in the salt crystals. Statistical analysis of the database suggests bioaccumulation of these metals from soil to T. nilotica. This reflects the importance of using T. nilotica as a model in the phytoremediation process as an established environmental clean-up technology.     

Assessment of metals in soil extracts and their uptake and movement within Tamarix nilotica at Lake Nasser banks, Egypt

This study aims to determine heavy-metal levels in soil from the banks of Lake Nasser, the ability of Tamarix nilotica to accumulate such metals from soil and hence its potential for phytoextraction. Soil and Tamarix samples were collected from the banks of four bights around Lake Nasser and analysed for Fe, Mn, Ca, Mg, Cr, Cu, Ni, Zn, Cd and Pb by atomic absorption spectrometry, whereas Na and K were measured by atomic emission spectrophotometry. Three different methods of extraction were used for the soil samples. Lead, copper and zinc were equally distributed between the exchangeable phase and Fe/Mn oxide-bound form, while other measured metals were mainly present in the Fe/Mn oxide fraction. With the exception of iron, all metals studied showed total concentrations within the geochemical background values. T. nilotica exhibited elevated concentrations of Na (36.2-48.5 mg g^-1) and K (2.74-4.33 mg g^-1) in stems, and relatively high concentrations of Pb, Cd and Co (0.39-1.03 µg g^-1, 0.24-1.3 µg g^-1 and 1.94-5.3 µg g^-1, respectively) are found in plant leaves. Bioaccumulation factors of Na and K (9.3 and 12.63, respectively) were high in T. nilotica stems. While the bioaccumulation of Pb, Cd, Co and Ni (2870.1, 2035.4, 10.5 and 5313.2, respectively) was high in plant leaves, Fe, Mn, Ca and Mg were accumulated relatively equally in plant stems and leaves. T. nilotica was found to secrete high amounts of Na, Ca and K, in addition to small amounts of all accumulated metals except Cd and Cu. These secreted metals appeared as salt crystals (67.5% Na; 25.8% Ca; 5% Mg; 1.5% K and 0.16% trace and minor elements) on the plant surface. The concentrations of all the metals studied in T. nilotica were higher than in the salt crystals. Statistical analysis of the database suggests bioaccumulation of these metals from soil to T. nilotica. This reflects the importance of using T. nilotica as a model in the phytoremediation process as an established environmental clean-up technology.     

Chlorophyll blooms and the spring/neap tidal cycle: observations at two stations on the coast of Connemara,Ireland

Water temperature, chlorophyll a and phaeophytin were recorded every 2 to 4 d, in order to describe short-term variability in phytoplankton blooms. Data were collected from May to September 1984 and March to September 1985 at shore stations on the coast of Connemara, western Ireland. Comparisons with offshore data suggest that the shore data is representative of coastal water. Chlorophyll a was highest at neap tides in spring, and at spring tides in early summer. In late summer, transient chlorophyll blooms occurred at neap tides, possibly due to a localized accumulation of flagellates in stable conditions.