Concentrations of nitrogen and phosphorus in streams draining the mixed land-use Dee Catchment, north-east Scotland

1.  There is still some uncertainty about large-scale influences on nutrient budgets in rivers. In particular, reduced forms of nitrogen (N) in organic forms might represent a significant fraction of the soluble N present in headwater streams, but this is not well quantified. Nitrate increases in relative importance downstream within agriculturally dominated areas. Here we appraise variations in N dynamics for a representative temperate but upland river, the Dee.
2.  In the Dee catchment, the source of organic N appears to vary seasonally. During summer under low flow conditions it originates primarily from in-stream biological production, while during the winter–spring period leaching from the plant–soil system would be the major contributor.
3.  On any individual sampling day, a wide range of N : P ratios can occur in the catchment area. Generally the narrowest N : P ratios occur during the summer and early autumn, particularly for upland catchments dominated by semi-natural vegetation. It is possible that some of the tributaries and upper region of the main river may be limited by N during the summer. The interpretation of the N : P ratios depends greatly upon the potential biological availability of the organic N, which remains unknown.
4.  Together, these data further illustrate that simple ideas about the relative limiting effects of N and P in temperate freshwaters may be misleading.     

Ecological change in British broadleaved woodland since 1947

Over the last 60 years Britain's broadleaved woodlands have undergone a complex pattern of ecological change. The total extent has expanded from c . 676 000 ha in 1947 to c . 904 000 ha in 2002, but there has also been significant turnover, with losses of ancient woodland and a gain from new planting. Structural change has occurred due, in part, to change in management. In 1947 21% of the broadleaved resource was classed as coppice, 28% as scrub and only 51% of the area as high forest, compared with 97% high forest in 2002. This has been accompanied by changes in ground flora and the regeneration pattern of tree species, which will impact upon the character of the woodlands. Woods have also become more ecologically isolated because of the decline in semi-natural vegetation in the surrounding countryside, although the long-term impacts of this are poorly understood. Other factors driving ecological change are increased pollution, change in grazing pressures, climate change, alien species and game management. Overall woodland specialist species and those of open habitats tend to be doing less well than woodland generalists. Progress has been made in reversing some adverse impacts such as acid deposition, and action is being taken to reduce the impact of others such as over-grazing by deer. However, some drivers, notably climate change, will be more difficult to address. In the long term integrated management of woods as landscape components whilst meeting economic and societal needs will be required.     

Decreasing trend of Helicobacter pylori infection in children with gastrointestinal symptoms from Buenos Aires, Argentina

Background: Helicobacter pylori infection is declining in developed and developing countries. The aim of this study was to retrospectively evaluate over an 8‐year period the rate of H. pylori infection in children with gastrointestinal symptoms from Buenos Aires, Argentina. Materials and Methods: We reviewed the records of children referred from 2002 to 2009 to the gastroenterology unit of the Children Hospital “Superiora Sor Maria Ludovica” for evaluation of upper gastrointestinal signs and symptoms in which the ¹³C‐urea breath test was performed to diagnose H. pylori infection and a sociodemographic questionnaire was obtained. Results: Records of a total of 1030 children and adolescents with a mean age of 9.99 years were included in the analysis. We found an H. pylori prevalence of 41.2% (95% CI, 36.9–46.0%) for the triennium 2002–2004, dropping to 26.0% (95% CI, 20.7–31.8%) in the triennium 2007–2009. Conclusion: Our results showed a significant decrease in H. pylori infection rates from children referred for upper gastrointestinal symptoms evaluation from 2002 to 2009, following the H. pylori epidemiologic trend reported in other countries.     

Short-Term Temporal Changes in Tree Live Biomass in a Central Amazonian Forest, Brazil

We monitored seventy-two 1 ha permanent plots spread over 64 km² of terra firme forest at Reserva Ducke (Manaus, Amazonas, Brazil) over 2-yr intervals to assess the effects of a soil and topographic gradient on the rate of change in the aboveground tree live biomass (AGLB). AGLB increased significantly over the 2-yr intervals, exhibiting a mean rate of change of 1.65 Mg/ha/yr (bootstrapped 95% CI: 1.15, 2.79). The rate of change varied according to tree size class; understory and sub-canopy trees exhibited higher rates of change. Over the whole period, the rate of change was not related to soil or topographic features of the plots, but there was evidence that the relationships varied depending on the year of measurement. In the plots monitored between 2001 and 2003 we found a significant relationship between AGLB change and the soil textural gradient, but this relationship was not evident in plots monitored between 2002 and 2004. This suggests that both the temporal variation in the soil–biomass change relationship and the size structure of the forest need to be included in models of biomass change in Amazonia. We also noted that the rate of biomass change is sensitive to the equation used to estimate AGLB. Allometric models that incorporate wood-density data provide higher per plot AGLB estimates, but lower rates of change, suggesting that variations in floristic composition have important implications for carbon cycling in diverse tropical forests.
Abstract in Portuguese is available at http://www.blackwell-synergy.com/loi/btp .     

Carbon dioxide exchange over multiple temporal scales in an arid shrub ecosystem near La Paz,Baja California Sur,Mexico

Arid environments represent 30% of the global terrestrial surface, but are largely under‐represented in studies of ecosystem carbon flux. Less than 2% of all FLUXNET eddy covariance sites exist in a hot desert climate. Long‐term datasets of these regions are vital for capturing the seasonal and interannual variability that occur due to episodic precipitation events and climate change, which drive fluctuations in soil moisture and temperature patterns. The objectives of this study were to determine the meteorological variables that drive carbon flux on diel, seasonal, and annual scales and to determine how precipitation events control annual net ecosystem exchange (NEE). Patterns of NEE from 2002 to 2008 were investigated, providing a record with multiple replicates of seasons and conditions. Precipitation was extremely variable (55–339 mm) during the study period, and reduced precipitation in later years (2004–2008) appears to have resulted in annual moderate to large carbon sources (62–258 g C m^?2 yr^?1) in contrast to the previously reported sink (2002–2003). Variations in photosynthetically active radiation were found to principally drive variations in carbon uptake during the wet growing season while increased soil temperatures at a 5 cm depth stimulated carbon loss during the dry dormant season. Monthly NEE was primarily driven by soil moisture at a 5 cm depth, and years with a higher magnitude of precipitation events showed a longer growing season with annual net carbon uptake, whereas years with lower magnitude had drier soils and displayed short growing seasons with annual net carbon loss. Increased precipitation frequency was associated with increased annual NEE, which may be a function of increased microbial respiration to more small precipitation events. Annual precipitation frequency and magnitude were found to have effects on the interannual variability of NEE for up to 2 years.     

Helicobacter pylori Seroprevalence in Symptomatic Veterans: A Study of 7310 Patients Over 11 Years

Background and Aims:  The prevalence of Helicobacter pylori infection has been decreasing in the USA, but recent data are lacking. This study evaluates the seroprevalence for anti- H. pylori antibodies in symptomatic veterans tested over the past 11 years.
Materials and Methods:  The same serum anti- H. pylori IgG detection system has been used at a tertiary care Veterans Affairs hospital since late 1996. Results of all tests performed from 1997 to 2007 were analyzed.
Results:  Of 7310 unique patients tested, 3982 (54.5%) were positive. Seropositivity declined from 70.8% in 1997 to 48.6% in 2002, then reached a plateau around 50%. A strong birth cohort effect was present, from a seropositivity of 72.7% for the veterans born before 1920 to 22% for those born between after 1980.
Conclusions:  Despite a constant birth cohort effect, H. pylori seropositivity among symptomatic veterans leveled down at ~50% after declining steadily from 1997 to 2002.     

Mass aerial migration in the carabid beetle Notiophilus biguttatus

Abstract.  1. Catches in traps at 12 and 200 m above ground, and observations with entomological radar, were used to document the high-altitude windborne movements of carabid beetles over agricultural land in south-east U.K. during July 1999, 2000, 2002, and 2004.
2.  Notiophilus biguttatus (Fabricius) was found to be the most abundant species at altitude, with flights by reproductively immature adults resulting in millions of beetles passing through a 1 km² window during the brief migration period in July 2002.
3. This result was unexpected in view of the conclusions of an earlier, classic Dutch study by den Boer and colleagues that classified N. biguttatus as a poorly dispersing species that tended to become isolated in remnants of natural habitat.
4. Reasons for the discrepancy between the two studies are discussed. It is hypothesised that the high mobility seen in the N. biguttatus populations results from a recent adaptation of the beetle's migration syndrome allowing it to exploit the mosaic of temporary habitat patches making up the increasingly intensive agroecosystems of northern Europe.     

Land cover change 2002–2005 in Borneo and the role of fire derived from MODIS imagery

Borneo has experienced heavy deforestation and forest degradation during the past two decades. In this study the Moderate Resolution Imaging Spectroradiometer was used to monitor land cover change in Borneo between 2002 and 2005 in order to assess the current extent of the forest cover, the deforestation rate and the role of fire. Using Landsat and ground observation for validation it was possible to discriminate 11 land cover classes. In 2002 57% of the land surface of Borneo was covered with forest of which 74% was dipterocarp and more than 23% peat swamp forest. The average deforestation rate between 2002 and 2005 was 1.7% yr^{− 1}. The carbon-rich ecosystem of peat swamp forests showed a deforestation rate of 2.2%. Almost 98% of all deforestation occurred within a range of 5 km to the forest edge. Fire is highly correlated with land cover changes. Most fires were detected in degraded forests. Ninety-eight per cent of all forest fires were detected in the 5 km buffer zone, underlining that fire is the major driver for forest degradation and deforestation.     

Climatic effects on arrival and laying dates in a long-distance migrant, the Collared Flycatcher Ficedula albicollis

Long‐distance migrants may respond to climate change in breeding, wintering or staging area by changing their phenology. The geographical variation in such responses (e.g. coastal vs. continental Europe) and the relative importance of climate at different spatial scales remain unclear. Here we analysed variation in first arrival dates (FADs) and laying dates of the Collared Flycatcher Ficedula albicollis in a central European population, from 1973 to 2002. The North Atlantic Oscillation (NAO) index correlated weakly with local temperature during the laying period. Decreasing spring temperatures until 1980 were associated with a trend towards later laying. The rate of warming (0.2 °C per year) and laying advancement (0.4 days per year) since 1980 are amongst the highest values reported elsewhere. This long‐term trend in laying date was largely explained by the change in climatic factors. The negative effect of local spring temperature on laying was relatively stronger than that of NAO. The number of clutches initiated on a particular day was marginally affected by the temperature 3 days prior to laying and the response of females to daily variation in temperature did not change over years. Correspondence between the average population‐level and the individual‐level responses of laying date to climate variation suggests that the advancement of laying was due to phenotypic plasticity. Despite warmer springs and advanced laying, FADs did not change over years and were not correlated with local spring temperature. Marginal evidence suggests later departure from wintering grounds and faster migration across staging areas in warmer conditions. Advancement of arrival was probably constrained by low local temperatures in early spring just before arrival that have not changed over years. The interval between first arrival and laying has declined since 1980 (0.5 days per year), but the increasing temperature during that period may have kept the food supply approximately unchanged.     

Changes in terrestrial carbon storage in the United States. 1: The roles of agriculture and forestry

• 1 Changes in the areas of croplands and pastures, and rates of wood harvest in seven regions of the United States, including Alaska, were derived from historical statistics for the period 1700–1990. These rates of land‐use change were used in a cohort model, together with equations defining the changes in live vegetation, slash, wood products and soil that follow a change in land use, to calculate the annual flux of carbon to the atmosphere from changes in land use.
• 2 The calculated flux increased from less than 10 TgC/yr in 1700 to a maximum of about 400 TgC/yr around 1880 and then decreased to approximately zero by 1950. The total flux for the 290‐year period was a release of 32.6 PgC. The area of forests and woodlands declined by 42% (160 × 10⁶ ha), releasing 29 PgC, or 90% of the total flux. Cultivation of soils accounted for about 25% of the carbon loss. Between 1950 and 1990 the annual flux of carbon was approximately zero, although eastern forests were accumulating carbon.
• 3 When the effects of fire and fire exclusion (reported in a companion paper) were added to this analysis of land‐use change, the uptake of carbon calculated for forests was similar in magnitude to the uptake measured in forest inventories, suggesting that past harvests account for a significant fraction of the observed carbon sink in forests.
• 4 Changes in the management of croplands between 1965 and 1990 may have led to an additional accumulation of carbon, not included in the 32.6 PgC release, but even with this additional non‐forest sink, the calculated accumulation of carbon in the United States was an order of magnitude smaller than the North American carbon sink inferred recently from atmospheric data and models.

     

Photoperiodic and temperature control of diapause induction and colour change in the southern green stink bug Nezara viridula

Abstract. The effect of photoperiod and temperature on the duration of the nymphal period, diapause induction and colour change in adults of Nezara viridula (L.) (Heteroptera: Pentatomidae) from Japan was studied in the laboratory. At 20 °C, the developmental period for nymphs was significantly shorter under LD 10 : 14 h (short day) and LD 16 : 8 h (long day) than under intermediate photoperiods, whereas at 25 °C it was slightly shorter under intermediate than short- and long-day conditions. It is assumed that photoperiod-mediated acceleration of nymphal growth takes place in autumn when day-length is short and it is unlikely that nymphal development is affected by day-length under summer long-day and hot conditions. Nezara viridula has an adult diapause controlled by a long-day photoperiodic response. At 20 °C and 25 °C in both sexes, photoperiodic responses were similar and had thresholds close to 12.5 h, thus suggesting that the response is thermostable within this range of temperatures and day-length plays a leading role in diapause induction. Precopulation and preoviposition periods were significantly longer under near-critical regimes than under long-day ones. Short-day and near-critical photoperiods induced a gradual change of adult colour from green to brown/russet. The rate of colour change was significantly higher under LD 10 : 14 h than under LD 13 : 11 h, suggesting that the colour change is strongly associated with diapause induction. The incidences of diapause or dark colour did not vary among genetically determined colour morphs, indicating that these morphs have a similar tendency to enter diapause and change colour in response to short-day conditions.     

Lipid content and fatty acid composition of target tissues in wild Perca fluviatilis females in relation to hepatic status and gonad maturation

Variation in liver ultra‐structure and composition in relation to energy mobilization was investigated in female perch Perca fluviatilis from the Meuse River between August 2001 and June 2002. In April, just before spawning, the lipo‐somatic index ( I _F) was 0·3%, the gonado‐somatic index ( I _G) was 28% and the total lipid content of the liver was 2·53%. The average areas of lipid droplets and mitochondria were 0·05 and 0·06 μm², respectively. Glycogen supply reached 7·9% of the total area of the hepatocyte. During the sexual resting period, females accumulated energy in perivisceral fat and in the liver to reach 1·6% I _F and 4·85% of liver lipid content in August with lipid droplets average size of 0·09 μm² and glycogen average area of 15%. Liver cells contained a weakly developed rough endoplasmic reticulum (RER) and a great number of small mitochondria (average size 0·02 μm²). The I _G was 0·6% at this time. During the whole annual cycle, the average lipid content of female liver never exceeded 3·9 ± 1·9%. The concentration of docosahexaenoic (DHA), linolenic and linoleic acids increased in mature gonads while linolenic and linoleic acids decreased in the liver during the same period. Fatty acid composition of muscles of perch was characterized by a high content of DHA.     

Long-term changes of aphid vectors of Barley yellow dwarf viruses in north-eastern Italy (Friuli-Venezia Giulia)

Migrations of aphid vectors of Barley yellow dwarf viruses (BYDV) were monitored using a Rothamsted Insect Survey suction trap in Friuli-Venezia Giulia (north-eastern Italy). Catches from 1983 to 2002 were studied for trends, correlations of total catches of each year with those of previous years, correlations between the autumn and the spring + summer catches of the same year and between spring + summer catches of one year with catches of the previous autumn. Infectivity of autumn alates was studied using biological tests, and infectivity indexes were calculated for all vector species and for Rhopalosiphum padi alone. Colonisation of barley and proportion of infected plants were checked in a field close to the suction trap from 1992 to 2002 and related to trap catches. Catches were also correlated to acreage dedicated to cereal and fodder crops in the region. During the 20 years, 15 BYDV vector species were caught in the trap, but only five species were found consistently colonising barley plants during autumn. R. padi was the most numerous species in catches, while Sitobion avenae was the predominant colonising species in the barley field. Relatively to R. padi , S. avenae colonies were about six times more numerous than expected from catches. The yearly abundance of catches of most species did not change significantly during the 20 years, with a few exceptions, significantly correlated to changes in the acreage dedicated to cereal and fodder crops. There was a significant decrease of the autumn catches of both R. padi and the total of BYDV vectors.     

Effect of sowing date and straw mulch on virus incidence and aphid infestation in organically grown faba beans (Vicia faba)

The effect of sowing date on aphid infestation and the incidence of aphid-transmitted viruses were investigated in organically managed, small-scale field experiments with two faba bean cultivars over 3 years (2002–04). As an additional factor, straw mulch was applied in 2 of the 3 years shortly before the start of vector activity in May. Virus incidence was determined using enzyme-linked immunosorbent assay and immunoelectron microscopy. Aphid flight activity was monitored using standard yellow water traps. Bean colonising aphids were assessed throughout the vegetation period by counting the number of plants infested with Acyrthosiphon pisum , Megoura viciae and Aphis fabae . Pea enation mosaic virus and bean yellow mosaic virus were the most abundant aphid-transmitted viruses, being detected in 22–54% and 9–69%, respectively, of the total number of virus-infected plants analysed per year. Further aphid-transmitted viruses found in faba bean were bean leaf roll virus, beet western yellows virus, clover yellow vein virus (in 2002) and soybean dwarf virus (in 2004). A. pisum was the predominant aphid species colonising faba bean plants. Early sowing compared with late sowing led to a significant reduction of the total virus incidence in faba bean in all 3 years. However, significantly decreased levels of A. pisum colonisation as a result of early sowing were observed only in 1 year and one cultivar. Irrespective of sowing date, straw mulching had no significant effects on virus incidence and aphid colonisation. Compared with late sowing, early sowing significantly increased bean yield in all 3 years and kernel weight in 2 years, whereas straw mulching had no effect on yield.     

Long-term effects of prescribed early fire, grazing and selective tree cutting on seedling populations in the Sudanian savanna of Burkina Faso

Annual early fire, selective tree cutting and exclusion of grazing are currently used as management tools in the Sudanian savanna of Burkina Faso although their long-term effects on seedling recruitment are poorly documented. A factorial experiment involving fire, grazing and cutting, each with two levels, was established in 1992 to study the effects of these management regimes and their interactions on the regeneration of woody species, and examine whether their effects varied temporally. Species richness, density and the morphology of seedlings were assessed in 1997 and 2002, and their relative changes were determined. The change in species richness of multi-stemmed individuals was significantly higher ( P  = 0.018) on plots that received fire × cutting treatment than the control plots. Significantly more species with single-stem were found on unburnt than on burnt plots ( P  < 0.001). Grazing tended to reduce the change in total density while fire ( P  < 0.001) and grazing ( P  = 0.029) significantly reduced the change in density of single-stemmed individuals. Selective cutting did not affect the total seedling density, but tended to reduce the change in single-stemmed seedling density. Principal component analysis revealed species-specific responses to treatments, particularly the relatively high abundance of lianas compared with other species.     

Quantifying small‐scale deforestation and forest degradation in African woodlands using radar imagery

Carbon emissions from tropical land‐use change are a major uncertainty in the global carbon cycle. In African woodlands, small‐scale farming and the need for fuel are thought to be reducing vegetation carbon stocks, but quantification of these processes is hindered by the limitations of optical remote sensing and a lack of ground data. Here, we present a method for mapping vegetation carbon stocks and their changes over a 3‐year period in a > 1000 km² region in central Mozambique at 0.06 ha resolution. L‐band synthetic aperture radar imagery and an inventory of 96 plots are combined using regression and bootstrapping to generate biomass maps with known uncertainties. The resultant maps have sufficient accuracy to be capable of detecting changes in forest carbon stocks of as little as 12 MgC ha^?1 over 3 years with 95% confidence. This allows characterization of biomass loss from deforestation and forest degradation at a new level of detail. Total aboveground biomass in the study area was reduced by 6.9 ± 4.6% over 3 years: from 2.13 ± 0.12 TgC in 2007 to 1.98 ± 0.11 TgC in 2010, a loss of 0.15 ± 0.10 TgC. Degradation probably contributed 67% (96.9 ± 91.0 GgC) of the net loss of biomass, but is associated with high uncertainty. The detailed mapping of carbon stock changes quantifies the nature of small‐scale farming. New clearances were on average small (median 0.2 ha) and were often additions to already cleared land. Deforestation events reduced biomass from 33.5 to 11.9 MgC ha^?1 on average_. Contrary to expectations, we did not find evidence that clearances were targeted towards areas of high biomass. Our method is scalable and suitable for monitoring land cover change and vegetation carbon stocks in woodland ecosystems, and can support policy approaches towards reducing emissions from deforestation and degradation (REDD).     

Ecosystem service values and land use change in the opium poppy cultivation region in Northern Part of Lao PDR

Land use change and land-cover impacts ecosystem services and functions. In this paper according to the study area’s land use characteristic and ecosystem type, the Land use category of the study area was divided into seven categories, including Forest, Grassland, Farmland, Water, Wetlands, Urban land and Barren land. The dynamic information of the forest Land use change during 10 years was calculated by the map algebra in ArcGIS 9.2. Both in 1992 and in 2002, Forest and Grassland were two largest Land use category in the study area. Forest took up 44.7% and 39.4% of the total area, and Grassland was 50.13% and 50.72% of the total area in 1992 and 2002. Finally, we valued change in ecosystem services delivered by each land use category using coefficients published by Costanza et al. [5]. Ecosystem services value of study area, the total ecosystem services value of 10.6 million hectares of this study area decreased by 11.74%. From the coefficient of sensitivity (CS) was less than unity in all case, it indicated that the total ecosystem services values was relatively inelastic and the results suggest that we have to pay attention more on land use change and finally, policy for driving forces of land use change were developed.     

Age, growth and food composition of Scorpaena porcus (Linnaeus, 1758) in the southeastern Black Sea

Samples of 525 Scorpaena porcus (Linnaeus, 1758) were collected from the southeastern Black Sea between January 2002 and May 2003 for information on age, growth, length–weight relationships and stomach contents. Total length of sampled fish ranged from 4.6 to 22.9 cm and total weight from 1.3 to 220 g. The sex ratio (1 : 1.61) was biased toward females (P < 0.05). Positive allometric growth was determined in the collected samples. The length–weight relationship for all individuals was described by the parameters a  = 0.0149 and b  = 3.09, with r ² = 0.99. The sample was composed of five age-classes (0–7 years). Three growth models were used to identify the growth characteristics: von Bertalanffy,     ; logistic,     ; and Schnute and Richards,     . Among these, the Schnute and Richards model was best fitted to the data as     The diet was composed of Carcinus mediterraneus (18.2%), Crangon crangon (12.1%), unidentified crab species (10.6%), Gobius sp. (4.6%), Palaemon sp. (4.5%), Macropipus sp. (4.5%), Engraulis encrasicholus (1.5%), Gastropoda (1.5%), and unidentified remains (34.9%).     

Climate-induced changes in high elevation stream nitrate dynamics

Mountain terrestrial and aquatic ecosystems are responsive to external drivers of change, especially climate change and atmospheric deposition of nitrogen (N). We explored the consequences of a temperature-warming trend on stream nitrate in an alpine and subalpine watershed in the Colorado Front Range that has long been the recipient of elevated atmospheric N deposition. Mean annual stream nitrate concentrations since 2000 are higher by 50% than an earlier monitoring period of 1991–1999. Mean annual N export increased by 28% from 2.03 kg N ha⁻¹ yr⁻¹ before 2000 to 2.84 kg N ha⁻¹ yr⁻¹ in Loch Vale watershed since 2000. The substantial increase in N export comes as a surprise, since mean wet atmospheric N deposition from 1991 to 2006 (3.06 kg N ha⁻¹ yr⁻¹) did not increase. There has been a period of below average precipitation from 2000 to 2006 and a steady increase in summer and fall temperatures of 0.12 °C yr⁻¹ in both seasons since 1991. Nitrate concentrations, as well as the weathering products calcium and sulfate, were higher for the period 2000–2006 in rock glacier meltwater at the top of the watershed above the influence of alpine and subalpine vegetation and soils. We conclude the observed recent N increases in Loch Vale are the result of warmer summer and fall mean temperatures that are melting ice in glaciers and rock glaciers. This, in turn, has exposed sediments from which N produced by nitrification can be flushed. We suggest a water quality threshold may have been crossed around 2000. The phenomenon observed in Loch Vale may be indicative of N release from ice features such as rock glaciers worldwide as mountain glaciers retreat.     

Support polygons and symmetricalgaits in mammals

The symmetrical gaits of quadrupedal mammals are oftendescribed in terms of two variables: duty factor (S = the stanceperiod of one foot, as a percentage of the gait cycle) and diagonality(D = the percentage of the cycle period by whichthe left hind footfall precedes the left fore footfall). We showthat support polygons are optimized during walking (i.e.the percentage of the locomotor cycle spent standing on only twofeet is minimized) for: (1) the diagonal-sequence, diagonal-coupletswalks characteristic of primates (50 < D < 75)when D = [hindlimb S]; (2) lateral-sequence,lateral-couplets walks (0 < D < 25)when D = [hindlimb S]− 50;(3) lateral-sequence, diagonal-couplets walks (25 < D < 50)when D = 100 −[forelimb S].To determine whether animal behaviour is optimal in this sense,we examined 346 symmetrical gait cycles in 45 mammal species. Ourempirical data show that mammalian locomotor behaviour approximatesthe theoretical optima. We suggest that diagonal-sequence walkingmay be adopted by ­primates as a means of ensuring thata grasping hindfoot is placed in a protracted position on a testedsupport at the moment when the contralateral forefoot strikes downon an untested support.  © 2002 The Linnean Societyof London, Zoological Journal of the Linnean Society , 2002, 136 ,401−420