Parallels and contrasts in the effects of drought on stream macroinvertebrate assemblages

• 1 It is axiomatic that unusually long dry periods (droughts) adversely affect aquatic biota. Recovery after drought is rapid by macroinvertebrates that possess strategies to survive drying or are highly mobile but other taxa take longer to recolonise depending on the timing, intensity, and duration of the dry phase.
• 2 Although drought acts as a sustained ‘ramp’ disturbance, impacts may be disproportionately severe when certain critical thresholds are exceeded. For example, ecological changes may be gradual while a riffle dries but cessation of flow causes abrupt loss of a specific habitat, alteration of physicochemical conditions in pools downstream, and fragmentation of the river ecosystem. Many ecological responses to drought within these habitats apparently depend on the timing and rapidity of hydrological transitions across these thresholds, exhibiting a ‘stepped’ response alternating between gradual change while a threshold is approached followed by a swift transition when a habitat disappears or is fragmented.
• 3 In two Australian intermittent streams, drought conditions eliminated or decimated several groups of macroinvertebrates, including atyid shrimps, stoneflies and free‐living caddisflies. These taxa persisted during the early stages of the drought but did not recruit successfully the following year, despite a return to higher‐than‐baseflow conditions. This ‘lag effect’ in response to drought emphasises the value of long‐term survey data. Although changes in faunal composition were inconsistent among sites, marked shifts in taxa richness, abundance and trophic organisation after the riffle habitat dried provide evidence for a stepped response.
• 4 Responses by macroinvertebrate assemblages to droughts of differing severity in English chalk streams were variable. The prolonged 1988–92 drought had a greater impact than shorter droughts in the early 1970s but recovery over the next 3 years was swift. Effects of the 1995 summer drought were buffered by sustained groundwater discharge from the previous winter. These droughts tended to reduce available riverine habitats, especially via siltation, but few taxa were eliminated because they could recolonise from perennial sections of the chalk streams.
• 5 In the contrasting environments of the intermittent streams studied in England and Australia, there are parallels in the rapid rates of recolonisation. However, recruitment by taxa that lack desiccation‐resistant stages or have limited mobility is delayed. Currently, long‐term data on these systems may be insufficient to indicate persistent effects of droughts or predict the impacts of excessive surface or groundwater abstraction or the increased frequency and duration of droughts expected with global climate change.

     

Population Reduction and Social Disorganization in Alouatta pigra Following a Hurricane

The opportunity to study the effects of a powerful hurricane on monkey populations, diet, and behavior via pre- and post-hurricane data was presented when hurricane Iris virtually destroyed the forest along Monkey River in southern Belize on October 8, 2001, including a 52-ha area where black howlers have been subjects since 1999. Before the hurricane, 8 social groups, averaging 6.37 members, had been stable in both group composition and range for q 2 years. The hurricane, which levelled much of the forest, resulted in the complete loss of the forest canopy. The trees that remained standing lost most or all branches and were 100% defoliated. The monkey population in the study area was reduced by 42% and survivers experienced a period of extended social disorganization involving transient individuals, high numbers of solitary monkeys, and small fragmentary social groups. The period of disorganization lasted 12 weeks, after which the number of solitaries reduced and stability of the large groups increased. Within the study area, 5 social groups have been more or less stable since ca. week 15; however, home ranges had yet to stabilize at week 35. The social and ranging effects are similar to what has been described for translocated primates. Post-hurricane diet was limited to fruit and leaves remaining in the deadfall for the first 2 weeks and to new leaves and leaf buds for many weeks after that. Normal fruit consumption in April and May was prevented by the failure of surviving trees to produce fruit. With the loss of forest canopy there has been increased use of low foliage and ground travel, and with the reduction in population density there has been a reduction in vocalization frequency.     

Spatio-temporal growth dynamics of a subAlpine Pinus uncinata stand in the French Alps

Natural forests are characterised by a high level of both spatial and temporal heterogeneity. The major processes involved in the creation and maintenance of forest heterogeneity in temperate climates are small-sized canopy disturbances. The resulting openings also modify the growth conditions for the remaining trees. The analysis of tree growth responses using dendrochronological techniques allows the reconstruction of the time sequence of the disturbances. The radial growth analysis is coupled here with both a spatial analysis and a demographic analysis of the forest structure. The age classes are temporally organised in cohorts and spatially distributed in spatial aggregates. The analysis of the most documented disturbance event allowed us to determine the size of the disturbance response patch (between 0.036 and 0.073 ha) and the duration of the disturbance (six years). We considered that this small-scale disturbance event may have been caused by the fall of a single tree and has freed the surrounding trees of its competition.     

Components of spatial patterning in a serpentine grassland

We analyzed the components of spatial patterning in species abundance in a diverse grassland dominated by early season, annual forbs. Species abundance, soil depth and gopher disturbances were measured by means of a nested spatial design on two 8m×20m plots that differed in the amount of larger scale variability in soil depth. Species distributions at the scale of the 10cm×10cm quadrat were highly clumped, with a decline in clumping with mean abundance. Even with the effect of abundance on clumping removed, species showed differences in the degree of clumping and these differences were consistent between the two plots. The abundance of the species and the frequency of disturbance correlated weakly to moderately with soil depth. Semivariance analyses indicated that the most common species all showed complex spatial patterning at a range of scales. Some of this variation corresponded to patterns of soil depth variation and patterns of gopher disturbance; however, a large amount of the spatial patterning in species abundance remains unexplained.     

Models of central pattern generators for quadruped locomotion I. Primary gaits

In this paper we continue the analysis of a network of symmetrically coupled cells modeling central pattern generators for quadruped locomotion proposed by Golubitsky, Stewart, Buono, and Collins. By a cell we mean a system of ordinary differential equations and by a coupled cell system we mean a network of identical cells with coupling terms. We have three main results in this paper. First, we show that the proposed network is the simplest one modeling the common quadruped gaits of walk, trot, and pace. In doing so we prove a general theorem classifying spatio-temporal symmetries of periodic solutions to equivariant systems of differential equations. We also specialize this theorem to coupled cell systems. Second, this paper focuses on primary gaits; that is, gaits that are modeled by output signals from the central pattern generator where each cell emits the same waveform along with exact phase shifts between cells. Our previous work showed that the network is capable of producing six primary gaits. Here, we show that under mild assumptions on the cells and the coupling of the network, primary gaits can be produced from Hopf bifurcation by varying only coupling strengths of the network. Third, we discuss the stability of primary gaits and exhibit these solutions by performing numerical simulations using the dimensionless Morris-Lecar equations for the cell dynamics.     

Predicting extinction risks for plants: environmental stochasticity can save declining populations

An emerging generalization from theoretical and empirical studies on conservation biology is that high levels of environmental stochasticity increase the likelihood of population extinction. However, coexistence theory has illustrated that there are circumstances under which environmental stochasticity can increase the chance of population persistence. These theoretical studies have shown that the sign of the effect of environmental stochasticity on population persistence is determined by interactions between life history and environmental stochasticity. These interactions mean that the stochastic and deterministic rates of population growth might differ fundamentally. Although difficult to demonstrate in real systems, observed life histories and variance in the vital rates of populations suggest that this phenomenon is likely to be common, and is therefore of much relevance to conservation biologists.     

A ‘minimum stress inflexion’ in relation to environmental and biotic influences on the dynamics of subtidal encrusting communities?

The sedentary fauna of sublittoral boulders was studied at five sites with very different environmental conditions within Lough Hyne Marine Nature Reserve (51° 30 N, 9° 18 W). The degree of competition was assessed through the construction of competitive hierarchies built up from the results of nearly 3000 interactions between organisms. Communities at all sites were predominantly hierarchically organised with few stand-offs recorded. Density of interactions was correlated with total space occupation. Flow speed at all sites was logged over 24 h and this factor also correlated with space occupation. Degree of disturbance was assessed through the use of marked boulders at each site. Species diversity peaked where all factors were moderate and was lowest at extreme sites. The effects of these environmental variables and biotic factors in determining community composition are discussed, and a working model detailing a Minimum Stress Inflexion is described. This represents an extension of the intermediate disturbance hypothesis in that it is proposed that disturbance in this system may serve to reduce sediment load as opposed to preventing space monopolisation in the classical model.     

Competitive abilities of three narrowly endemic plant species in experimental neighborhoods along a fire gradient

We conducted field experiments manipulating lichens, shrubs, and herbs along a time-since-fire gradient and assessing effects on three endemic herbaceous species of Florida scrub: Eryngium cuneifolium, Hypericum cumulicola, and Polygonella basiramia. Responses included seed germination, survival, biomass, and fecundity. Transplants into recently burned patches generally had higher survival, larger biomass, and greater reproductive output than transplants into long-unburned patches. Open areas and sites near oaks frequently were more favorable than sites near Florida rosemary. Ground lichens did not affect germination but increased mortality rate of seedlings. Neighboring small shrubby and herbaceous species did not affect the performance of these species. Of the three species, naturally occurring E. cuneifolium were farthest from large shrubs, and their microhabitats had the least ground lichens and shrubs. Eryngium cuneifolium and H. cumulicola are capable of forming persistent seed banks and their recruitment after fire depends mostly on these dormant seeds. Polygonella basiramia relies on seed dispersal and immediate seed germination to colonize recently burned patches. Management for these species should involve variable fire regimes to allow all three species to persist along with many other scrub endemics.     

RESPONSE OF PACIFIC WALRUSES TO DISTURBANCES FROM CAPTURE AND HANDLING ACTIVITIES AT A HAUL-OUT IN BRISTOL BAY, ALASKA

Observations were made on herds of the Pacific walrus ( Odobenus rosmarus divergens ) to study their response during the capturing and handling of adult males in summer 1995 at a haul-out at Cape Peirce in southwestern Alaska. Three behaviors (alertness, displacement, and dispersal) were quantified from 16 capture sessions. Herd sizes ranged from 622 to 5,289 walruses. Handling of an immobilized walrus consisted of attempts to attach telemetry devices to the tusks and collect various biological samples. Handling activities resulted in an average of about 10-fold or greater levels of behavior in alertness, displacement, and dispersal than during precapture and darting periods. High levels of behavior usually occurred within the first 45 min of handling. In 8 of 10 capture sessions, walruses returned to predisturbance levels of behavior within 40 min of cessation of the handling disturbance. Alertness and displacement were moderately and negatively correlated with herd size during the handling period, which may reflect an effect of a threshold distance from the point of disturbance to responding individuals. Observations of walruses tagged with VHF radio transmitters indicated that the activities from a given capture session did not preclude tagged walruses from using the haul-out over a subsequent 11 -wk monitoring period. Moreover, non-tagged walruses continued to extensively use the haul-out during and after the period in which capture sessions were conducted.     

Material Spiraling in Stream Corridors: A Telescoping Ecosystem Model

Stream ecosystems consist of several subsystems that are spatially distributed concentrically, analogous to the elements of a simple telescope. Subsystems include the central surface stream, vertically and laterally arrayed saturated sediments (hyporheic and parafluvial zones), and the most distal element, the riparian zone. These zones are hydrologically connected; thus water and its dissolved and suspended load move through all of these subsystems as it flows downstream. In any given subsystem, chemical transformations result in a change in the quantity of materials in transport. Processing length is the length of subsystem required to “process” an amount of substrate equal to advective input. Long processing lengths reflect low rates of material cycling. Processing length provides the length dimension of each cylindrical element of the telescope and is specific to subsystem (for example, the surface stream), substrate (for instance, nitrate), and process (denitrification, for example). Disturbance causes processing length to increase. Processing length decreases during succession following disturbance. The whole stream-corridor ecosystem consists of several nested cylindrical elements that extend and retract, much as would a telescope, in response to disturbance regime. This telescoping ecosystem model (TEM) can improve understanding of material retention in running water systems; that is, their “nutrient filtration” capacity. We hypothesize that disturbance by flooding alters this capacity in proportion to both intensity of disturbance and to the relative effect of disturbance on each subsystem. We would expect more distal subsystems (for example, the riparian zone) to show the highest resistance to floods. In contrast, we predict that postflood recovery of functions such as material processing (that is, resilience) will be highest in central elements and decrease laterally. Resistance and resilience of subsystems are thus both inversely correlated and spatially separated. We further hypothesize that cross-linkages between adjacent subsystems will enhance resilience of the system as a whole. Whole-ecosystem retention, transformation, and transport are thus viewed as a function of subsystem extent, lateral and vertical linkage, and disturbance regime. Received 15 April 1997; accepted 1 September 1997.