Vegetation Physiognomy Research Articles

Paleoclimatic Significance of Mid-Cretaceous Floras from the Middle Clarence Valley, New Zealand

Evidence from leaf-margin analysis and vegetational physiognomy indicates that a mid-Cretaceous flora from the South Island of New Zealand grew at a mean annual temperature of 10 degrees C. This compares with an identical estimate derived from a coeval flora in northern Alaska. Both floras grew at about the same latitude and in coastal plain settings. This study provides the first direct comparison of non-marine near-polar climates in the pre-Tertiary record.

Vegetation mapping by integration of floristic analysis, GIS and remote sensing. An example from Tigray (Ethiopia)

ABSTRACT An integrated approach combining floristic analysis, Geographic Information Systems (GIS) and Remote Sensing (RS) was used to map the vegetation of a Tigray district (Ethiopia). Vegetation types were identified using a floristic method. For each type the community niche was defined on the basis of altitude and parent material. The combination of the physiognomic map obtained by RS with the niches of floristic-vegetation types was obtained with a GIS application using the Digital Elevation Model (DEM). A floristic-physionomic vegetation map combining floristic diversity and physiognomy of vegetation was produced. The map has been used for a numerical classification of administrative units (tabias), based on the percentage of cover of floristic-physiognomic vegetation types. Three main groups of tabias have been defined, each of them related to one altitudinal belt and well characterised by a floristic-physiognomic vegetation type. This characterisation and description could be very useful for the development of plans for the rehabilitation of the area based on local plant species.

Quaternary fossil fauna of South Canterbury, South Island, New Zealand

This study documents the Late Quaternary fossil fauna from 59 fossil sites in the South Canterbury downlands, South Island, New Zealand. Twenty‐seven sites were predator accumulations attributed to laughing owls, two were accumulated by falcons, two were swamp sites, and the rest were pitfalls or rockshelter deposits. A total of 60 indigenous species of birds, one bat, three rodents, one tuatara, four geckos, and two skinks were represented in the combined faunas from these sites. When the birds known historically are added, the fauna of inland South Canterbury downlands is shown to have had a minimum diversity of 87 species in the Late Holocene before human disturbance. There were no marked faunal changes associated with the climatic amelioration from the Otira Glacial to the Holocene, such as were seen in the wetter western regions. During the Holocene in South Canterbury, species characteristic of Otiran grassland – shrubland faunas survived alongside those typical of Holocene western closed‐forest communities, although the latter were not as abundant as they were in the west. The large diversity of birds, relative to other New Zealand faunas, is therefore the result of continued survival in Canterbury into the Holocene, of grassland – shrubland mosaics similar to those that characterised widespread parts of New Zealand during the glacial periods. However, the geographic feature that permitted the survival of these critical vegetation physiognomies also caused the area to be dry, and vegetation was hence most prone to destruction by fire. This complex ecosystem is now almost totally destroyed.

Fuel biomass and combustion factors associated with fires in savanna ecosystems of South Africa and Zambia

Fires are dominant factors in shaping the structure and composition of vegetation in African savanna ecosystems. Emissions such as CO2, NOx, CH4, and other compounds originating from these fires are suspected to contribute substantially to changes in global biogeochemical processes. Limited quantitative data exist detailing characteristics of biomass, burning conditions, and the postfire environment in African savannas. Fourteen test sites, differentiated by distinct burn frequency histories and land‐use patterns, were established and burned during August and September 1992 in savanna parklands of South Africa and savanna woodlands of Zambia. Vegetation physiognomy, available fuel loads, the levels of biomass consumed by fire, environmental conditions, and fire behavior are described. In the South African sites, total aboveground fuel loads ranged from 2218 to 5492 kg ha−1where fire return intervals were 1–4 years and exceeded 7000 kg ha−1at a site subjected to 38 years of fire exclusion. However, fireline intensity was only 1419 kW m−1at the fire exclusion site, while ranging from 480 to 6130 kW m−1among the frequent fire sites. In Zambia, total aboveground fuel loads ranged from 3164 kg ha−1in a hydromorphic grassland to 7343 kg ha−1in a fallow shifting cultivation site. Dormant grass and litter constituted 70–98% of the total fuel load among all sites. Although downed woody debris was a relatively minor fuel component at most sites, it constituted 43–57% of the total fuel load in the fire exclusion and shifting cultivation sites. Fire line intensity ranged between 1734 and 4061 kW m−1among all Zambian sites. Mean grass consumption generally exceeded 95%, while downed woody debris consumption ranged from 3 to 73% at all sites. In tropical savannas and savanna woodlands of southern Africa, differences in environmental conditions, land‐ use patterns, and fire regimes influence vegetation characteristics and thus influence fire behavior and biomass consumption.

Dioecy in the Cerrado vegetation of Central Brazil

Summary Occurrence of dioecy in Cerrado, Neotropical savanna region in Central Brazil, is analyzed here. Frequency of dioecious species and their phenological and morphological characteristics were studied for a cerrado woody community near Brasilia, and a broader comparative survey was done using phytosociological data for different cerrado areas and vegetation physiognomies. Nine dioecious species were found in the studied community (15% of the species in the area), most with simple open flowers visited by small insects. Flowering phenology differed from species to species including long, sparse flowering and comucopian strategies. Distance between flowering individuals varied from 13 to 70 m. Gender related differences in spatial distribution and in height or circumference were tested for three of the species and significant departure from random was found only for Rapanea guianensis . The regional survey showed frequencies of dioecy higher in denser woodlands and forest than in open cerrado areas. Most species presented small promiscuous flowers, probably associated to pollination by small insects, and fleshy fruits. Among the species occurring in the cerrado areas one could separate a group of dioecious shrubs and treelets really adapted to cerrado environment and another group of evergreen forest species which occur only in denser woodlands, the cerradao. Differences in the frequency of dioecy seems to be related to the occurrence of this latter group and the appearance of cerradao.

The physical environment and major plant communities of the Karoo National Park, South Africa

The major plant communities of the Karoo National Park are described using the methods of the Zurich-Montpellier school of phytosociology, to assist with the formulation of a management strategy for the park. The vegetation physiognomy consists of Montane Karoo grassy shrublands. Karoo grassy dwarf shrublands. Karoo succulent dwarf shrublands and riparian thicket. Steep elevation and precipitation gradients within the study area have a direct impact on gradients in the vegetation. High elevation (1 800 m), and relatively high rainfall (406 mm) montane grasslands occupy communities dominated by grasses (Merxmuellera disticha, Themeda triandra) and woody species (Diospyros austro-africana, Elytropappus rhinocerotis, Euryops annae, Passerina montana). The increasing aridity away from the escarpment edge in a northerly direction is steep, and Montane Karoo dwarf shrublands replace these mesic communities. Species such as Eriocephalus ericoides, Rosenia oppositifolia and Pteronia tricephala dominate. At lower elevation (800 m) the precipitation is very low (175 mm) and uncertain (coefficient of variation of 78 ). The substrata influence the vegetation, with the sandy substrata of the drainage lines supporting more woody taxa (Acacia karroo, Lycium cinereum) and grasses (Hyparrhenia hirta, Stipagrostis namaquensis, Cenchrus ciliaris). Moving away from the mesic environment of the riparian zone, rapid desiccation occurs and the most xeric communities are encountered, dominated by Stipagrostis obtusa, S. ciliata and Pent-da incana. This document provides descriptions of the general communities and their associated landscape, lithology and soils.

A Functional Analysis of New Zealand Alpine Vegetation: Variation in Canopy Roughness and Functional Diversity in Response to an Experimental Wind Barrier

There is an increasing interest in describing species assemblages in terms of function or texture, as an alternative to floristics. However, the use of objective criteria for evaluating plant function remains uncommon. A procedure for determining plant fractal dimensions was applied to the measurement of canopy roughness of alpine plants, as a major functional character in high-alpine cushionfield vegetation in southern New Zealand. The study was carried out within a 50 x 30 m area located around an experimental snow fence. The snow fence was erected on an originally uniform site 32 years prior to this study. Since its construction it has caused visible changes in snow lie and in the composition of vegetation on its leeward side, apparently because of its ameliorating effect on microenvironmental parameters. Floristic patterns across the study area, revealed by a DCA ordination of species frequency data, showed a strong association with snow depth, corresponding to the sheltering effect of the snow fence. Vegetation canopy roughness increased along a gradient of increasing snow depth, consistent with the expectation that plants growing in the most exposed alpine environments would possess smooth canopies that promote laminar air flow over their surface and thus enhance plant/air temperature differentials. Functional diversity in terms of canopy roughness similarly increased with increasing snow depth, consistent with expectations of community structure theory. Species richness and Shannon-Wiener species diversity showed similar relations. It is concluded that plant canopy roughness, measured by means of fractal dimensions of canopy profiles, has value for predicting the physiognomy of alpine vegetation from the microenvironment. It is also practical to measure. It should be considered for future studies examining vegetation texture in the alpine zone.

The Relationship of Vegetation to Methane Emission and Hydrochemical Gradients in Northern Peatlands

1 The bryophyte and vascular flora were described for a range of forested and open peatlands in the mid-boreal Clay Belt region of Canada, and in the subarctic region of the Labrador Trough, Quebec. The floristic patterns and their relationships to methane (CH4) emission, hydrology and water chemistry were analysed with classification (TWINSPAN), detrended correspondence analysis (DCA) and canonical correspondence analysis (CCA). 2 Despite differences in vegetation physiognomy, chemistry, and geology, water chemistry (pH, Ca, Mg, Kcorr) and hydrology explain most of the species distribution in both peatland regions. In the Clay Belt, bryophytes respond primarily to water table position, and secondarily to water chemistry; the reverse is true for the vascular flora. The bryophyte and vascular ordinations are more similar to each other in the Labrador Trough than in boreal sites. 3 Although CH4 emissions are lower in subarctic than in boreal sites, mean CH4 flux is strongly correlated with mean water table position (r2 = 0.73) in both regions, but is not correlated with water chemistry. The highest CH4 emissions (seasonal mean > 100mg m-2 day-') occur in raised bog and patterned poor fen pools where the peat mat is degrading. 4 The relationships among CH4 flux, mean water table position, and species distribution are similar in both boreal and subarctic regions. Bryophytes are generally better predictors of CH4 flux than vascular plants, except for certain species in flooded sites (e.g. Carex spp.). Bryophytes are also better indicators of the long-term average position of the water table, reflecting relative zones of methane production and oxidation over the season. In the vascular plant analyses, shrub cover indicates dry conditions and low CH4 flux in the sharply differentiated string-flark-pool topography of the subarctic fens, but is less predictive of CH4 emission in the drier hummockhollow microtopography at boreal sites where shrubs and trees are more pervasive. Overall herbaceous or sedge cover is not indicative of CH4 flux in either region.

Woodland pattern and structure in a peasant farming area of Zimbabwe: ecological determinants and present and past use

This article relates the species pattern and structure of miombo woodland in a deforested peasant farming area of central Zimbabwe to its ecological determinants and its present and past use. Data on species composition, basal area, height and physiognomy of the woody vegetation are presented for different parts of the landscape. The nature and degree of disturbance from harvesting for fuel and timber is quantified. Woodland in the arable areas and close to village lines is shown to be dominated by fruit trees, other trees with cultural controls on their cutting and species which quickly invade disturbed ground. The latter may be highly productive of woody biomass. In the grazing area, deeper soils which were cultivated in the past have retained an open structure and tend to be dominated by heavily coppiced Brachystegia spiciformis and Combretum molle (also dominants of less disturbed miombo woodland). Lithic soils which have not been cleared support denser, coppiced woodlands also dominated by climax species. Woody vegetation on kopjes and along riverine fringes is less disturbed in terms of its distribution, species composition, density and height than woodland in other parts of the landscape. The spatial pattern and nature of cutting is shown to vary between land use categories, between different species and according to the dimensions of individual stems. Miombo woodland shows a relatively high degree of stability in species composition under disturbance by cutting: of the 94 species included in the analysis, relatively few were shown to be significantly associated with a particular soil type, catenal position or land use category.

THE IMPORTANCE OF HABITAT STRUCTURE AND FOOD SUPPLY FOR CARABID BEETLES (COLEOPTERA, CARABIDAE) IN PEAT BOGS

AbstractDuring the period 1985–1989, the carabid assemblages of a raised peat bog in eastern central Sweden and of six peatlands in New York State and Maine were studied. The principal methods of investigation were pitfall trapping in summer, and quadrat sampling in winter. In addition to carabids, potential food organisms were evaluated from the pitfall catches.The bogs, although of different types and located on different continents, show similar physiognomic vegetation units. Eight of these units are considered relevant as carabid microhabitats. The carabid assemblages of open sunny, and wooded shaded habitats are clearly distinguished. Only a few species are adapted to the extreme environment of open Sphagnum L. habitats. Among them, stenotopic species are dominant. The seasonal distribution of two species on Swedish Ryggmossen [Agonum ericeti (Panzer), Pterostichus rhaeticus Heer] suggests that the carabids of open bog plateau habitats use damp Sphagnum lawns for their summer activities and migrate to drier hummocks for overwintering.On bog plateaus, larger populations of ground beetles were only found where a pronounced pattern of hummocks and hollows exists. In bogs with a prevailing uniform vegetation of Sphagnum mats, carabids were conspicuously scarce. Thus, the development of a bog indigenous carabid fauna is closely related to the presence of a hummock–hollow mosaic. The data indicate that in most bog habitats food is not a limited resource and does not play a key role as a population controlling factor.

Landscape and Climatic Control of Woody Vegetation in a Dry Tropical Ecosystem: Turkana District, Kenya

The spatial organization of a dry woodland/ savanna/shrub-steppe ecosystem in a 9000 km2 region of arid Northern Kenya was explored by analysing the abundance and distribution of woody vegetation in relation to landscape gradients and gradients in rainfall. Woody species assemblages were clustered into four major groups. Three of these (and most of the sites) were dominated by species of Acacia. Acacia tortilis was the community dominant in riparian and riverine zones, A. senegal on hilly and rocky sites, and A. reficiens on non-riparian sites with fine soils. The fourth group, found at highest elevations, was distinguished by a very low abundance of Acacia. These soil/landform associations were systematically distributed along land system gradients (mountains, pediment, valley, bajada), thus relating woody species groups to large-scale landscape characteristics. Tree woody canopy cover ranged from < 1.0% to 100% over the region as a whole. Cover was directly related to precipitation when the effects of water concentration in riparian zones were removed. However, canopy cover was not greatly influenced by either species composition or landscape characteristics. Fire reduced woody canopy cover, however, fires appeared to affect a relatively small portion of the ecosystem. Recent anthropogenic disturbances such as wood-cutting and livestock corralling were encountered in patches, but the cumulative long-term effects of patch scale disturbances could not be discerned in vegetation patterns at the regional scale. Vegetation physiognomy (woodland, bush, bushed grassland, etc.) was controlled by both water availability and landscape pattern. Woodland and forest occurred almost exclusively in riparian and riverine situations while the driest parts of the region supported dwarf shrub grassland with few trees. Over the region as a whole, climate and landscape gradients converged in diverse ways, giving rise to structurally variable associations of woody plant species. Vegetation structure in tropical savannas and dry woodlands is often interpreted in terms of competition between woody and herbaceous life forms for soil moisture. The outcome of this competitive interaction is thought to be influenced by disturbances that shift the system from one stable state to another. However, our findings suggest that dry tropical ecosystem structure is hierarchically constrained by physical factors: by climate at regional to continental scales; by topographic effects on rainfall and landscape water redistribution, and geomorphic effects on soil and plant available water at the landscape to regional scales; and finally by water redistribution and disturbance at local and patch scales.

Distribution and dynamics of shrub roots in recent coastal dune valley ecosystems of Belgium

A chronosequence, up to 60 years old, was investigated in the Westhoek Nature Reserve. Root growth of the shrubs in dune valleys was found to be restricted to the surface Albi and Bbi horizons which often did not exceed 25 cm. The thickness of these horizons and the amount of litter and humus could be explained by the position along secondary topo-hydrosequences within the chronosequence. The mostly stratified sands underlying the Albi and Bbi horizons had a higher bulk density. In these relatively dense Cd horizons root growth of new colonizing shrubs seems to be possible only along old root or stem channels of the previous vegetation of dune building plants such as Ammophila and Salix which grew in pace with the accumulating dune. It is concluded that root growth in these sandy soils is restricted by the high penetration resistance of the closely packed sands in the subsurface horizons. The root penetration problem in these sandy soils explains the uniform vegetation physiognomy better than other climatic or edaphic factors.

Special Paper: A Global Biome Model Based on Plant Physiology and Dominance, Soil Properties and Climate

A model to predict global patterns in vegetation physiognomy was developed from physiological considera- tions influencing the distributions of different functional types of plant. Primary driving variables are mean coldest- month temperature, annual accumulated temeprature over 5C, and a drought index incorporating the seasonality of precipitation and the available water capacity of the soil. The model predicts which plant types can occur in a given environment, and selects the potentially dominant types from among them. Biomes arise as combinations of domi- nant types. Global environmental data were supplied as monthly means of temperature, precipitation and sunshine (interpolated to a global 0.5 grid, with a lapse-rate correc-

Evidence of Incidental Nest Predation and Its Effects on Nests of Threatened Grassland Birds

In this study we document the effect of nest predation on a grassland bird community that includes habitat for regionally endangered or rare bird species. Predation on bird eggs and nestlings on this sandplain grassland in southern Maine, USA was positively correlated (P<0.01) with foraging for invertebrates by striped skunks (Mephitis mephitis) in plots of primary skunk usage. The relationship between rates of predation and vegetation physiognomy was analyzed on three spatial scales, but no strong correlation was found at any level. The hypothesis that skunk predation at this site was incidental, rather than targeted towards birds' nests, was supported by the significant correlation between skunk invertebrate-foraging activity and nest predation, the absence of skunk foraging behavior consistent with active nest searching, the absence of a relationship between nest success and vegetative cover or proximity to a forest/grassland edge, the low density of breeding birds (<2.0 territories ha-') on these grasslands, and the small proportion of avian material, less than 1.5%, reported in the summer skunk diet. Although nest predation was high (58.0% overall), its unpredictability probably limits the utility of certain antipredator behaviors by birds nesting at this site. Such incidental nest losses can severely limit recruitment, and thus populations, of rare or endangered species.

The coenological organization of ectomycorrhizal macrofungi in the Laurentide Mountains of Quebec

Ectomycorrhizal macrofungus communities were classified across a sequence of four forest associations in the Laurentide Mountains. Cluster analysis was used to delimit two types of coenological units: grouping and sociological group. Three macrofungal groupings and two subgroupings, which vary with physiognomic and floristic–edaphic parameters of the forest sites, were delimited. Each of these grouping units is characterized by one of the six delimited sociological groups of fungi that are consecutively replaced following changes in latitude and altitude. When considering the similarity of species composition, Laurentide groupings appear to be isolated from other macrofungal assemblages in North American temperate forests and mainly affected by geographic partitioning of sites and by the physiognomy and type of forest vegetation within a given phytoclimatic region. As seen by the respective richness at the family level, Eurasian and Laurentide communities show similar patterns of zonal distribution, mainly owing to the role of climatic factors. A numerical analysis supported the hypothesis that the edaphic environment, as well as the vascular vegetation, might exert a significant influence on the delimitation and distribution of ectomycorrhizal communities. The interdependence of ectomycorrhizal communities and phytocoenoses at the ecosystem level is discussed. Key words: cluster analysis, community, ectomycorrhiza, macrofungus, macromycete, mycocoenology.

Detection and Classification of Muskox Habitat on Banks Island, Northwest Territories, Canada, Using Landsat Thematic Mapper Data

The feasibility of using Landsat Thematic Mapper data for mapping muskox summer habitat was tested on northern Banks Island, Northwest Territories. Digital image enhancement and classification techniques were examined to determine if summer foraging habitats could be detected and mapped using Thematic Mapper imagery. Interpretations of the satellite data were verified in the field during the summers of 1988 and 1989. The most important summer foraging habitats for muskoxen included the wet sedge meadow, graminoid tundra and graminoid/dwarf shrub tundra cover types. These lowland habitats were generally distinguishable on enhanced colour images and were easily differentiated from upland areas. The most suitable colour composite for differentiating muskox summer habitats was the near-infrared (band 4), shortwave infrared (band 5) and red (band 3) spectral bands displayed in red, green and blue respectively. Upland cover types, including dwarf shrub tundra, hummocky tundra and dwarf shrub/lichen barrens, were more difficult to differentiate because of spectral variability resulting from differences in plant cover and site characteristics. The classified image had an overall accuracy of 88%. The summer habitats of particular importance to muskoxen had classification accuracies of 84-89%. Detection of important foraging habitats on Thematic Mapper imagery is attributable to the spectral distinctiveness of wet graminoid communities and the high spectral sensitivity and spatial resolution of the infrared sensors, which allow detection of differences in surface moisture and vegetation physiognomy.Key words: bison habitat, boreal forest, digital image analysis, supervised classification, Landsat Thematic Mapper data, habitat mapping, Bison bison athubuscue, remote sensing

Nest-site characteristics of prairie shorebirds

Nest-site characteristics of eight shorebird species were studied during the period 1982–1984 at Last Mountain Lake National Wildlife Area in south central Saskatchewan. Plant species composition at nests varied significantly among species and was correlated with distance to bare ground and aquatic habitat. Flora at Wilson's phalarope (Phalaropus tricolor) nests was correlated with clutch initiation date, reflecting a seasonal shift in nesting habitat. Species differed significantly in cover types at nests. For American avocet (Recurvirostra americana), killdeer (Charadrius vociferus), and Wilson's phalarope, cover types differed between nests and random sites; cover types also differed between nest cups and the immediate surrounding habitat for killdeer and Wilson's phalarope. Interspecific differences in nest-site vegetation physiognomy were portrayed by the first two factors of a principal components analysis, which accounted for 73 and 22% of variance in the data, respectively. Principal components analysis assembled species on a habitat gradient that varied according to cover type, and vegetation density and heterogeneity.

Avian Resource Use in a Caribbean Pine Plantation

I studied resource use by birds in a Caribbean pine (Pinus caribaea) plantation in Luquillo Experimental Forest, Puerto Rico, from 1981 to 1986. I used multivariate analyses to quantify and provide a classification of foraging relationships among bird species. The analysis of food-searching behavior by birds indicated that resource use was closely associated with vegetation physiognomy. The presence of specific kinds of vegetation was important in determining the presence and abundance of some bird species. Frugivores and nectarivores foraged in understory flowering and fruiting shrubs. Among insectivores, foliage-gleaners were found mainly in the canopy and hoverers were concentrated in the understory. Dead hanging leaves and vine thickets provided important foraging substrates for some species. Forest managers can improve habitat for birds in plantations by providing habitat heterogeneity through maintenance and encouragement of understory vegetation diversity, vine thickets, and native upperstory trees. J. WILDL. MANAGE. 52(2):274-279 Information on the impacts of land management practices on neotropical birds, and particularly the changes that occur when heterogenous tropical forest ecosystems are converted to pine plantations, is limited. In Australia, exotic pine (Pinus radiata) plantations supported 50% of breeding bird species found in the original eucalyptus forests (Gepp 1976, Suckling et al. 1976, Driscoll 1977). Only Falkenberg et al. (1983) has investigated bird communities in neotropical pine plantations; they found fewer birds in Caribbean pine plantations than in native forest sites in Jamaica. Resource use by birds in neotropical pine plantations has not been quantitatively examined. Information on bird use of neotropical pine plantations is needed because 33% of all tropical plantations are in the neotropics and the most commonly planted species are pine (Pinus spp.) (Evans 1982). The objectives of my research were to describe avian resource use in a Caribbean pine plantation and analyze niche structure, food resource relationships, and other factors associated with the bird assemblage patterns. This study was conducted under the auspices of the Institute of Tropical Forestry, U.S. Forest Service, Puerto Rico, and the U.S. Man and the Biosphere Program on Tropical Forests. I am indebted to A. E. Lugo for guidance, cooperation, and helpful suggestions. I also profited from discussions with W. J. Arendt, G. P. Bauer, C. E. Bock, B. B. Cintron, C. A. Delannoy, J. C. Figueroa, G. E. Freeman, J. A. Moreno, T. K. Nakamura, T. R. Strong, F. H. Wadsworth, P. L. Weaver, and J. W. Wiley. I gratefully acknowledge the valuable field assistance of A. Arendt, W. J. Arendt, and E. Hernandez-Prieto and the forest structural information provided by A. E. Lugo. Financial support came from a National Research Council (Ford Found.) Postdoctoral Fellowship, Grant-in-Aid from the University of Colorado, and a National Geographic Society Grant.

Food resources of the wild Barbary macaque (Macaca sylvanus) in high‐altitude fir forest, Ghomaran Rif, Morocco

A small geographically isolated population of the Barbary macaque inhabits a high‐altitude fir forest habitat (Abies pinsapo) in the Ghomaran region of the Rif mountains of northern Morocco. The climate of this region is Mediterranean, but the altitude (1600–2100 m) causes winters to be cold (as low as ‐8.0 d̀C) with snow occurring from November to May (snowfall as deep as 1.5 m). The primary winter feeding adaptation is the ability to ingest high quantities of fir foliage; in spring, the macaques took a high diversity of leafy food items from all vegetation layers; in summer, the macaques foraged terrestrially for a high diversity of food items including seeds, small fruits, bulbous geophytes, and animal foods (including tadpoles from small streams); in autumn, the macaques returned to arboreal foraging, primarily feeding on oak acorns (Quercus ilex), fir seeds and yew fruit (Taxus baccata). The macaques were capable of ingesting 100 of 195 (51%) of all identified plant species in the region, although during the four‐month winter, the macaques only averaged 12.5 common food items. A comparison of the study area with the prime habitat of the Barbary macaque‐the high‐altitude cedar forests of the Moroccan Moyen Atlas‐indicates that climate and vegetation physiognomy are highly similar in both regions. Correspondingly, there is a high degree of similarity in macaque diet in both regions in terms of feeding behaviour by season, food diversity and specific feeding techniques. In the Ghomara, the winter feeding adaptation of fir foliage eating parallels that of the Barbary macaque in cedar forest (winter foraging for cedar foliage). This enables the Barbary macaque to exploit the Ghomaran fir forest habitat during the cold, snowy winters much the same as it does cedar forest habitat throughout a major portion of its geographical range.

Small Mammal Populations of an Old-Field Chronosequence: Successional Patterns and Associations with Vegetation

We sampled the small mammal fauna of 18 successional old-fields, aged 2–57 years since abandonment from agriculture, and examined patterns of diversity and abundance along successional and productivity gradients. Through time, vegetational standing crop, cover, litter, and plant species composition changed, but vegetational physiognomy did not; thus, we examined small mammal populations within a grassland chronosequence. Small mammal densities were low and were not strongly associated with successional time. Species richness and density, however, were strongly associated with vegetational standing crop and nitrogen content. Peromyscus leucopus densities were best correlated with variability in vegetation ( SD vegetational biomass), whereas density of Microtus pennsylvanicus and Sorex cinereus were best correlated with vegetational nitrogen content. Overall, the data suggest that the diversity and abundance of small mammals were limited by nitrogen.