ABNORMAL MIGRATION AND HYBRIDIZATION OF LARUS ARGENTATUS AND L. FUSCUS AFTER INTERSPECIES FOSTERING EXPERIMENTS

Between 1962 and 1966 eggs of Larus argentatus and L. fuscus were interchanged andalmost 900 young were reared by the wrong species. Many of these cross-fostered young were later recovered or retrapped on and away from the colonies.
The British population of argentatus is sedentary and ringed birds are not recovered outside Britain, whereas fuscus normally migrates. Many cross-fostered argentatus migrated to France, Spain and Portugal, areas where fuscus is common, but they did not migrate as far as the control fuscus . It is possible that these cross-fostered argentatus had followed their foster parents when these migrated, but this is unlikely as the cross-fostered fuscus also migrated although their foster parents would have remained in Britain.
Despite wide ecological and behavioural overlaps, interbreeding between L. argentatus and L.fuscus is exceedingly rare. However, as a result of cross-fostering experiments, 31 and 40 mixed pairs were found on Skokholm in 1968 and 1969 respectively. Although some of the birds involved were unringed it is probable that all the adults in mixed pairs had been cross-fostered. Other cross-fostered birds were found mated with their own species and it appears that the sex of the imprinted birds was important. Female gulls will usually only mate with males of their own species, or in the case of the cross-fostered birds, with males of their foster species. Males will mate with either species.
Evidence is given that suggests that the colour of the mantle and wings is important in species recognition at long range, and the colour of eye-ring and join of the mandibles for recognition at short range. The role of voice is uncertain but general behaviour is probably unimportant.     

The sympatric breeding of Larus fuscus, L. cachinnans and L. argentatus in western France

An opportunity to study the validity of the species Larus cachinnans , considered by some authors as merely a group of races related to either L. fuscus or L. argentatus , is given by its sympatric breeding with these species. Both of these latter species have recently extended their breeding range southwards, while Mediterranean representatives of the cachinnans group have spread northwards, and the three forms now breed side by side over c. 350 km along the Atlantic coast of France, where their breeding biology was studied from 1983 to 1990. L. cachinnans is very territorial, some pairs holding territories almost all year round; argentatus pairs may defend territories from mid-winter; and the supposedly migratory fuse us are on their territories for only a short period before laying. L. cachinnans is an early breeder, laying from late March to early May, while fuscus lay from May into June, and argentatus has a protracted breeding season overlapping the other two. Interbreeding occasionally occurs; it can be productive in the case of fuscus x argentatus whereas fuscus x cachinnans attempts did not produce any young. No cachinnans x argentatus pair has been observed. This indicates strong reproductive isolation, evidence that L. cachinnans must be considered a good species.     

Diet dimorphism' within an Oystercatcher Haematopus ostralegus population– adaptive significance and effects on recent distribution dynamics

Of the 50 or so Oystercatcher pairs breeding on Skokholm in 1965 and 1966, 15% bred near the shore and 85% bred inland; 28% fed their young on limpets, and 72% on terrestrial arthropods and earthworms. 'Terrestrial-feeders' were three times more successful than limpet-feeders in fledging young, irrespective of variation in initial brood size and of hatching date but depending on differential losses to gull predation. This is because parents' vigilance is relatively low in limpet-feeders' territories, and limpet-fed young are on average undernourished and hence behave relatively carelessly. Since the 1930s the population of shore-breeding gulls has increased considerably on Skokholm. A certain increase of the Oystercatcher population during this period seems to have been associated with a shift inland, until the island became saturated at the late 40s. It is proposed that the increase in shore-breeding gulls has triggered a genetically based behavioural change in Oystercatchers, which has been responsible for the shift to inland habitats and to terrestrial food, which promotes a population increase. This interpretation for events on Skokholm may be applicable for the general trend of habitat shift and population increase of other British populations.     

A census of the Manx Shearwater Puffinus puffinus on Skomer,Skokholm and Middleholm,west Wales

In 1997/98 a census was made of the Manx Shearwater populations on three Welsh Islands: Skomer, Skokholm and Middleholm. The census was made by counting all the burrows in the winter, playing a tape-recording of a male call (to which only males respond) down a sample of the burrows during the incubation period and applying a correction for the response rate (the proportion of males which respond). The response rate was determined by playing the tape-recordings down a set of burrows in which eggs had been laid and where the incubating birds could be examined. Knowing the total number of burrows, the number of burrows sampled, the number of responses and the response rate enabled us to make an estimate of the population size on each island. Skomer, with an estimated population of some 102000 breeding pairs is probably the largest colony of Manx Shearwaters in the world; the estimates for Skokholm and Middleholm were 46000 and 3000 respectively. These figures are compared with earlier censuses undertaken using traditional methods.     

The behaviour of seabirds foraging at fishing boats around Shetland

Among the different types of fishing vessels around Shetland, whitefish trawlers attract the largest numbers of scavenging seabirds and provide the most food. Offal was almost all consumed by seabirds, predominantly by Fulmars Fulmarus glacialis , which excluded other species by their aggression. Fulmars generally ignored discarded whole fish, which were mainly taken by Great Black-backed Gulls Larus marinus , Gannets Sula bassana and Great Skuas Catharacta skua . Although flatfish were usually ignored because seabirds found them difficult to swallow and they sank faster, most discarded roundfish were consumed. Herring Gulls L. argentatus , Lesser Black-backed Gulls L. fuscus and Kittiwakes Rissa tridactyla were rarely able to obtain offal or discards. Herring Gulls and Lesser Black-backed Gulls spent much time on the periphery of feeding flocks while Kittiwakes rarely attempted even to join these. Most of the birds at trawlers were in adult plumage, and it is suggested that the low proportion of immature birds present was a further reflection of the highly competitive feeding conditions at trawlers. We suggest that likely changes in fishing practice and seabird population sizes in the immediate future may result in Herring Gulls, Lesser Black-backed Gulls and Great Skuas finding feeding on waste around trawlers increasingly difficult, so they may be further displaced by Fulmars, Gannets and Great Black-backed Gulls.     

Seasonal grazing rates and food processing by tropical herbivorous fishes

Seasonal variability in grazing rates and food-processing characters were assessed for three abundant fishes in a tropical rocky shore: the damselfish Stegastes fuscus , the parrotfish Sparisoma atomarium , and the surgeonfish Acanthurus bahianus . Significant differences were found in grazing rates among hour of day and seasons, and in food-processing characters among seasons for the three fishes. Grazing rates for S. atomarium and A. bahianus peaked at 1300 and 1400 hours for S. fuscus . Three main periods of different intensity in bite rates, low, intermediate and intense, were identified for all fishes. As expected, total bite rates, ingestion rates and gut fullness were highest in A. bahianus , the largest species studied, followed by S. atomarium and S. fuscus. S. atomarium with fused jaw teeth, holds the highest bite size and S. fuscus the lowest one. Gut turnover was high for S. fuscus and similar for S. atomarium and A. bahianus . Grazing rates and food-processing characters seem to vary between seasons not only due to changes in temperature, but also affected by other factors. Fishes employ different food-processing mechanisms and bear some differences in food-processing characters to achieve optimal energetic supplies from a poor nutrient food resource.     

Effects of an alien invertebrate species and wave action on prey selection by African black oystercatchers (Haematopus moquini)

Abstract Shorebirds foraging in the intertidal have been shown to exert a significant effect on assemblage level processes; this is particularly true of the oystercatcher–limpet–algae system. The African black oystercatcher (Haematopus moquini) is endemic to the southern African coastline, where it plays a significant role in ecosystem processes as a rocky‐shore predator, especially of mussels and limpets. This understanding was based on studies of a rocky shore environment that has since been considerably modified following invasion of an alien mussel (Mytilus galloprovincialis). This invasion has not only changed the relative proportions of different food types on the shore, but has also greatly increased overall food biomass. We tested the previous model that food selection by oystercatchers reflected prey abundance and that intake by male and female oystercatchers differed owing to bill morphology. We predicted that this difference would persist despite the changed nature of the food base. We also predicted that wave action would modify prey selection as a result of both its influence on prey behaviour and its impact on searching and handling times of the birds. Overall, both sexes consumed more limpets than expected by encounter rate alone, but contrary to prediction, the relative proportions of different prey types taken post invasion did not differ between the sexes. Dietary convergence is interpreted as a result of greatly increased food biomass on the shore, which is also reflected in increased oystercatcher densities since the invasion. Also contrary to prediction there was no evidence that waves acted as indirect modifiers of the interaction between oystercatchers and their prey. The results of this study indicate that models of trophic cascades will need to be altered in the event of a significant change in a trophic level, which then effects behavioural changes in the key predator.     

Experimental studies on homing in the intertidal patellid limpetCellana tramoserica (Sowerby)

Summary A model of two-dimensional random walk was developed to allow statistical tests for the presence of homing behaviour in intertidal populations of the limpetCellana tramoserica. Not all limpets return to a home-site after feeding excursions. Some move around at random. The latter pattern of movement was tested to justify the assumptions of the model. Limpets can stop homing and begin to move at random; moving limpets can become homers. Thus, random movement and consistent homing behaviour are not discrete patterns of behaviour in a population.Controlled field experiments showed that the proportion of animals which home is not affected by the height on the shore, force of wave action, the cover of macroalga, nor the irregularity of the substratum. Small limpets, however, home more than large ones.Close proximity to other homing limpets causes individuals to stop homing and move away. Alterations of density demonstrated thatCellana shows density-dependent dispersal. More limpets emigrate from areas of increased density and more immigrate into areas of decreased density, when compared with control areas. Less animals immigrate into, and remain in, areas from which microalgal food was experimentally removed. More animals emigrate from these areas.These experiments support the hypothesis that homing behaviour is an adaptation which regulates local density and dispersion to maximize utilization of food resources and, thus, to reduce intraspecific competition for food at high densities of limpets.     

Predation by small mobile aquatic predators regulates populations of the intertidal limpet Patella vulgata (L.)

Highly mobile aquatic predators are known to forage in the intertidal during periods of immersion. There is limited quantitative information, however, on the extent to which these predators influence the abundance of grazing molluscs which are known to have a key role in structuring intertidal assemblages. Our preliminary video observations revealed that crabs and small fish were abundant on shores in southwest England during high-tide. We then used manipulative experiments to quantify the effect of small mobile aquatic predators on the abundance of limpets (Patella vulgata L.). On the lower shore at two moderately sheltered rocky shores three treatments were established: complete cage, partial cage (cage control) and uncaged (natural condition). The complete cages excluded all predators. The partial cage treatment allowed full access to small predators and the uncaged treatment allowed access to all predators. After two months, limpet abundance in uncaged and partial cage treatments had declined by around 50% compared to the complete cage treatment. Population structure also changed with survival of larger individuals being greater than smaller individuals in the open and partial cage treatments compared to the complete cage treatment. The effects of excluding predators were consistent at small (meters) and large spatial scales (kilometres) and hence, it would appear that the outcomes of our research are generally applicable to similar shores in the region.To explore the mechanism behind the differential effects of predators according to prey size, we compared the detachment force required to remove limpets of differing sizes from the shore. This was around four times greater for larger individuals than for smaller ones indicating that smaller limpets were more vulnerable to predation. These effects were also consistent between locations. Subsequent laboratory observations showed that the crabs Carcinus maenas (L.), Necora puber (L.) and Cancer pagurus (L.) which are locally abundant predators of limpets, had differing handling behaviour but were all highly efficient at removing limpets from substratum. Hence, shell width and attachment force appeared to be critical factors influencing the vulnerability of limpets to predation by these crabs. Limpets are known to control the abundance of macroalgae on shores in the North-east Atlantic and so our conclusions about the role of mobile predators in regulating the abundance of these grazers are important to our broader understanding of the ecology of these shores.     

The grazing effect of limpets on the macroalgal community of a rocky intertidal shore

Abstract Abundance of macro-algae in the mid-littoral zone on a Victorian intertidal rocky shore varied seasonally, algae being rare in Summer and common in Winter and Spring. Field experiments demonstrated that, of two species of grazing limpets (Cellana tramoserica and Siphonana diemenensis), only Siphonaria had a major effect on the abundance of foliose algae (e.g. Scytopsiphon lomeniarid) and neither species had a great effect on the encrusting algae (Ralfsia spp.). There was no evidence of competition for food between the two species of limpets, in contrast with results found for similar species in New South Wales.     

The interplay of physical and biological factors in maintaining mid-shore and low-shore assemblages on rocky coasts in the north-west Mediterranean

This study examined the interactive effects of grazing by limpets and inclination of the substratum in maintaining differences between mid-shore and low-shore assemblages of algae in the northwest Mediterranean, at different scales of space and through time. Alternative models leading to different predictions about these effects were proposed and tested. Limpets were excluded by fences from areas of the substratum at mid levels on the shore. The response of algal assemblages to this manipulation was compared with control and enclosure plots at the same level, and with unmanipulated plots in the low shore where limpets are less abundant. The effects of limpets were examined at several replicated sites (0.1–4 km apart) for each slope of the substratum (nearly horizontal vs vertical), at different locations (hundreds of kilometres apart) and at different times. Individual taxa responded differently to limpet exclusion. The percentage cover of the coarsely branched and filamentous algae increased significantly in exclosure plots, in some loser reaching values found on the low shore. These patterns, however, varied greatly from shore to shore and significant effects were found both on horizontal and vertical substrata. Multivariate analyses indicated that grazing by limpets accounted for about 20% of the differences between mid-shore and low-shore assemblages. This effect was independent of substratum inclination and was consistent in space and time, suggesting that physical conditions were not as stressful for macroalgae on vertical substrata as initially supposed. Variable recruitment of algae is proposed as a possible explanation for the lack of consistency in the effects of limpets at the scale of the shore. The results of this study emphasize the need for multiple-scale analyses of the interactive effects of physical and biological factors to understand the organization of natural assemblages. Received: 22 June 1999 / Accepted: 15 November 1999     

世界主要国家耕地动态变化及其影响因素

在世界人口持续攀升、全球耕地面积不断减少的背景下,探讨世界主要国家耕地变化其影响因素,对于分析预测未来世界耕地变化趋势、研究世界粮食安全具有积极意义。选择2050年人口过亿的17个国家和耕地面积排名前10的国家为研究对象,在分析1961—2007年耕地总量变化、人均耕地变化的基础上,探讨了耕地变化影响因素。研究结果表明,从20世纪60年代到2007年间的不同时期内,有越来越多的国家表现出耕地减少趋势,而人均耕地面积减少的国家个数高达90%以上。满足人口消费需求、城市化与经济发展是大多数国家耕地总量变化的主要动力;而人口快速增长、城市化则是导致许多国家人均耕地显著减少的重要原因。     

Arrival and habitat use by Lapwings Vanellus vanellus in the early breeding season

The arrival on the breeding grounds and habitat use in relation to agricultural land use by Lapwings Vanellus vanellus was compared in rough grazing and arable study areas in the Midland Valley of Scotland. First arrivals took place in early February. Males preceded females and territory occupation by the males began soon after first arrival. During the pre-breeding period birds remained in flocks and day time habitat use and activity was strongly influenced by the lunar cycle. Birds fed mainly in those habitats in which prey (leatherjackets and earthworms) were most plentiful, and in which their feeding success was high. The choice of nesting habitat was not influenced by food availability in the immediate vicinity of the nest site but by the crypticity of the clutch and incubating adults. On rough grazing birds preferred unimproved land while on arable land spring cereal was the preferred habitat. The choice of actual nesting field on the arable land was influenced by the risk of predation and the proximity of suitable feeding fields for the adults and chicks.     

An analysis of distribution and abundance of populations of the high-shore limpet,Notoacmea petterdi (tenison-woods)

Summary Notoacmea petterdi occurs only on vertical surfaces at the highest levels on the shore. On a given surface, density of limpets decreases with increasing height. Mean and maximum sizes of limpets increase with increasing height, due to faster rates of growth at higher levels where densities are lower. Rates of mortality are also negatively correlated with height on the shore, and are therefore generally less at higher levels. Both adult and juvenile limpets exhibit well-developed homing behaviour. There is no evidence of seasonal migrations, density-dependent migrations, nor gradual upward migrations as limpets grow older.Hypotheses are formed on the basis of these preliminary observations and measurements. Results of subsequent experimental manipulations of densities support the hypothesis that density-dependent mortality, due to intraspecific competition, plays a major role in regulating population densities of this limpet. It is also suggested that the pattern of settlement of juvenile limpets is the major factor that determines the limits to the vertical distribution of this species.     

Distribution and abundance of the acmaeid limpet,Patelloida latistrigata,and its interaction with barnacles

Summary At Cape Banks, New South Wales, adults of the small intertidal limpet, Patelloida latistrigata occur exclusively in the barnacle zone, and are primarily associated with the barnacle, Tesseropora rosea. Limpet density increases with barnacle density. Juvenile limpets can be found throughout the barnacle zone, and on patches of bare rock that may be temporarily available at lower levels on the shore. The failure of juveniles to survive and grow in places other than among Tesseropora is due to a combination of factors. These include desication at high levels on the shore, smothering by rapidly growing algae low on the shore, and the grazing activities of the larger limpet, Cellana tramoserica. These latter two factors also reduce the survival of experimentally transplanted adult Patelloida: algae by covering the substratum and smothering the limpets, and Cellana by outcompeting them for food. The density of Cellana is greater on patches of bare rock than among barnacles, and these large limpets may be unable to move and feed effectively over the irregular surface created by Tesseropora. Patelloida, however, is small enough to feed over and among these barnacles, and hence has a refuge from competition with Cellana. Barnacles may also provide shelter from the effects of desiccation and strong wave action, and thus increase the survival of juvenile Patelloida. By being associated with barnacles, however, Patelloida becomes vulnerable to intermittent predation by the whelk, Morula marginalba. This effect may be serious enough to eliminate small local populations of limpets, either by direct predation or by removing the refuge-providing barnacles. The association with barnacles may also limit the maximum size to which Patelloida can grow. Patelloida is not always found with Tesseropora, and adults in different localities can be found in association with other sessile organisms. It may be argued that small species of limpets require a spatial refuge from physical and/or biological pressures. To examine this hypothesis, the relationship between Patelloida and Tesseropora is compared to other published accounts of limpets with specialised modes of life.     

Adaptive significance of facultative paedomorphosis in Triturus alpestris (Amphibia, Caudata): resource partitioning in an alpine lake

1. Facultative paedomorphosis is a polymorphism that has important evolutionary implications in promoting morphological differentiation and variation in habitat use. It occurs in several urodele species throughout the world. Several hypotheses based on life-history theory have been proposed to explain the wide range of habitats in which facultative paedomorphosis occurs, suggesting multifactorial causes.
2. In harsh habitats, such as mountain lakes, paedomorphosis might promote niche partitioning between morphs. This hypothesis was tested by comparing micro-habitat use and diet of two coexisting morphs in an alpine lake.
3. Paedomorphs occupied all microhabitats in the lake while metamorphs occurred only along the shoreline or at the water surface. Paedomorphic newts were frequent in deep water, where they foraged mainly on plankton. Plankton was poorly exploited by metamorphic newts, which fed mainly on terrestrial insects. Difference between morphs in microhabitat use, and consequently in the diet, were consistent in both sexes and in juveniles.
4. In adults, the mass and energy value of stomach contents did not differ between morphs, suggesting a similar food availability in the habitats occupied.
5. This study indicates habitat partitioning between morphs involving dietary differences. Specific benefits and costs of each ontogenetic pathway may allow their coexistence in this deep and fishless lake. Paedomorphosis benefits individual newts by making new food resources available and presumably reducing competition at the shore of the lake. However, the proximate causes of such an ontogenetic switch remain unclear.     

江苏省土地利用变化的图谱特征

基于地学信息图谱理论与方法,运用ArcGIS 10.0软件,以江苏省1990、2000和2010年3期遥感解译数据为空间数据源合成江苏省1990—2000和2000—2010年两个10年阶段的土地利用变化图谱以及耕地利用变化模式图谱,据此揭示江苏省土地利用变化时空特征及规律.结果表明: 1990—2000年,江苏省土地利用变化以耕地与城乡建设用地、耕地与水域以及耕地与林草地的图谱单元互换为主要特征,尤其是以“耕地→城乡建设用地”、“耕地→水域”最为显著,耕地数量减少、空间分布集中.2000—2010年,江苏省土地利用变化最显著的仍是“耕地→城乡建设用地”图谱单元;同时,“草地→水域”、“城乡建设用地→水域”图谱类型分布集中且扩大趋势明显;与1990—2000年相比,空间图谱变化总面积和空间分离度均有所增大.江苏省耕地利用变化模式以后期变化型为主,前期变化型次之,反复和持续变化型耕地所占比例较小.     

Physical and biological conditions on a steep intertidal gradient at Rottnest Island,Western Australia

Seven species of grazing molluscs, two littorinids, one nerite, three limpets and one chiton, lived on the vertical intertidal rock wall at the landward edge of the coastal limestone platforms at Rottnest Island, Western Australia. On the average, the vertical ranges of these species overlapped broadly, although a consistent zonation pattern was conspicuous from the platform surface upwards for 2 m. On these vertical intertidal shores, physical and biological conditions were predicted quantitatively from easily made measurements of vertical height on the shore; the percentage of time any shore level was immersed in seawater, the percent weight loss of plaster clods, the standing crop of algae, and the growth rate of the limpet, Notoacmea onychitis, all decreased linearly or semi-logarithmically with increasing height on the shore. The standing crop of animals was greatest, largely due to the presence of the chiton, between 40 and 70 cm from the platform surface and decreased rapidly down the shore and more gradually towards higher shore levels. We interpret this information and the positive correlation between algal production rate, and egestion rate of the animal community at various levels of the shore as evidence supporting the idea that food may be in short supply on these shores.     

Defining a land boundary for sustainable livestock consumption

The need for more sustainable production and consumption of animal source food (ASF) is central to the achievement of the sustainable development goals: within this context, wise use of land is a core challenge and concern. A key question in feeding the future world is: how much ASF should we eat? We demonstrate that livestock raised under the circular economy concept could provide a significant, nonnegligible part (9–23 g/per capita) of our daily protein needs (~50–60 g/per capita). This livestock then would not consume human‐edible biomass, such as grains, but mainly convert leftovers from arable land and grass resources into valuable food, implying that production of livestock feed is largely decoupled from arable land. The availability of these biomass streams for livestock then determines the boundaries for livestock production and consumption. Under this concept, the competition for land for feed or food would be minimized and compared to no ASF, including some ASF in the human diet could free up about one quarter of global arable land. Our results also demonstrate that restricted growth in consumption of ASF in Africa and Asia would be feasible under these boundary conditions, while reductions in the rest of the world would be necessary to meet land use sustainability criteria. Managing this expansion and contraction of future consumption of ASF is essential for achieving sustainable nutrition security.     

Morphological and behavioral evidence for adaptive diversification of sympatric Hawaiian limpets (Cellana spp.)

The endemic Hawaiian limpets (Cellana exarata, Cellana sandwicensis, and Cellana talcosa), reside at different elevations on wave-exposed rocky shores and comprise a monophyletic lineage that diversified within Hawai'i. Here, I report phenotypic differences in shell, soft tissue, and behavioral characters among these limpets and discuss their potential utility in exploiting their respective niches. The high-shore limpet, C. exarata, is characterized by a tall round shell, short mantle tentacles, and long evasion distance when confronted by a predatory gastropod. The mid-shore limpet, C. sandwicensis, is characterized by a shorter oblong shell, long mantle tentacles, and a short evasion distance when confronted by a predatory snail. The low-shore, shallow-subtidal limpet, C. talcosa, is characterized by a flat shell that is thin in juveniles and disproportionately massive in large adults (relative to the other two species), and mantle tentacles of varying lengths (some individuals exhibit short tentacles, some long). These species-specific suites of characters are likely to confer specific fitness advantages on the high shore (C. exarata) where thermal and desiccation stress is severe, on the mid shore (C. sandwicensis) where hydrodynamic forces are severe, and on the low-shallow subtidal shore (C. talcosa) where pelagic predators have free access to the limpets. These data add to the growing body of evidence for adaptive diversification and speciation in the Hawaiian Cellana, and in marine species in general.