Showing papers in "Journal of Biogeography in 2000"

Maps of Pleistocene sea levels in Southeast Asia: shorelines, river systems and time durations

Abstract: Aim Glaciation and deglaciation and the accompanying lowering and rising of sea levels during the late Pleistocene are known to have greatly affected land mass configurations in Southeast Asia. The objective of this report is to provide a series of maps that estimate the areas of exposed land in the Indo-Australian region during periods of the Pleistocene when sea levels were below present day levels. Location The maps presented here cover tropical Southeast Asia and Austral-Asia. The east–west coverage extends 8000 km from Australia to Sri Lanka. The north–south coverage extends 5000 km from Taiwan to Australia. Methods Present-day bathymetric depth contours were used to estimate past shore lines and the locations of the major drowned river systems of the Sunda and Sahul shelves. The timing of sea level changes associated with glaciation over the past 250,000 years was taken from multiple sources that, in some cases, account for tectonic uplift and subsidence during the period in question. Results This report provides a series of maps that estimate the areas of exposed land in the Indo-Australian region during periods of 17,000, 150,000 and 250,000 years before present. The ancient shorelines are based on present day depth contours of 10, 20, 30, 40, 50, 75, 100 and 120 m. On the maps depicting shorelines at 75, 100 and 120 m below present levels the major Pleistocene river systems of the Sunda and Sahul shelves are depicted. Estimates of the number of major sea level fluctuation events and the duration of time that sea levels were at or below the illustrated level are provided. Main conclusions Previous reconstructions of sea-level change during the Pleistocene have emphasized the maximum lows. The perspective provided here emphasizes that sea levels were at their maximum lows for relatively short periods of time but were at or below intermediate levels (e.g. at or below 40 m below present-day levels) for more than half of each of the time periods considered.

Neotropical seasonally dry forests and Quaternary vegetation changes

Abstract: Seasonally dry tropical forests have been largely ignored in discussions of vegetation changes during the Quaternary. We distinguish dry forests, which are essentially tree-dominated ecosystems, from open savannas that have a xeromorphic fire-tolerant, grass layer and grow on dystrophic, acid soils. Seasonally dry tropical forests grow on fertile soils, usually have a closed canopy, have woody floras dominated by the Leguminosae and Bignoniaceae and a sparse ground flora with few grasses. They occur in disjunct areas throughout the Neotropics. The Chaco forests of central South America experience regular annual frosts, and are considered a subtropical extension of temperate vegetation formations. At least 104 plant species from a wide range of families are each found in two or more of the isolated areas of seasonally dry tropical forest scattered across the Neotropics, and these repeated patterns of distribution suggest a more widespread expanse of this vegetation, presumably in drier and cooler periods of the Pleistocene. We propose a new vegetation model for some areas of the Ice-Age Amazon: a type of seasonally dry tropical forest, with rain forest and montane taxa largely confined to gallery forest. This model is consistent with the distributions of contemporary seasonally dry tropical forest species in Amazonia and existing palynological data. The hypothesis of vicariance of a wider historical area of seasonally dry tropical forests could be tested using a cladistic biogeographic approach focusing on plant genera that have species showing high levels of endemicity in the different areas of these forests.

Mid-Holocene and glacial-maximum vegetation geography of the northern continents and Africa

Abstract: BIOME 6000 is an international project to map vegetation globally at mid-Holocene (6000 14C yr BP) and last glacial maximum (LGM, 18,000 14C yr BP), with a view to evaluating coupled climate-biosphere model results. Primary palaeoecological data are assigned to biomes using an explicit algorithm based on plant functional types. This paper introduces the second Special Feature on BIOME 6000. Site-based global biome maps are shown with data from North America, Eurasia (except South and Southeast Asia) and Africa at both time periods. A map based on surface samples shows the method's skill in reconstructing present-day biomes. Cold and dry conditions at LGM favoured extensive tundra and steppe. These biomes intergraded in northern Eurasia. Northern hemisphere forest biomes were displaced southward. Boreal evergreen forests (taiga) and temperate deciduous forests were fragmented, while European and East Asian steppes were greatly extended. Tropical moist forests (i.e. tropical rain forest and tropical seasonal forest) in Africa were reduced. In south-western North America, desert and steppe were replaced by open conifer woodland, opposite to the general arid trend but consistent with modelled southward displacement of the jet stream. The Arctic forest limit was shifted slighly north at 6000 14C yr BP in some sectors, but not in all. Northern temperate forest zones were generally shifted greater distances north. Warmer winters as well as summers in several regions are required to explain these shifts. Temperate deciduous forests in Europe were greatly extended, into the Mediterranean region as well as to the north. Steppe encroached on forest biomes in interior North America, but not in central Asia. Enhanced monsoons extended forest biomes in China inland and Sahelian vegetation into the Sahara while the African tropical rain forest was also reduced, consistent with a modelled northward shift of the ITCZ and a more seasonal climate in the equatorial zone. Palaeobiome maps show the outcome of separate, independent migrations of plant taxa in response to climate change. The average composition of biomes at LGM was often markedly different from today. Refugia for the temperate deciduous and tropical rain forest biomes may have existed offshore at LGM, but their characteristic taxa also persisted as components of other biomes. Examples include temperate deciduous trees that survived in cool mixed forest in eastern Europe, and tropical evergreen trees that survived in tropical seasonal forest in Africa. The sequence of biome shifts during a glacial-interglacial cycle may help account for some disjunct distributions of plant taxa. For example, the now-arid Saharan mountains may have linked Mediterranean and African tropical montane floras during enhanced monsoon regimes.

Palaeovegetation of China: a pollen data‐based synthesis for the mid‐Holocene and last glacial maximum

Abstract: Pollen data from China for 6000 and 18,000 C-14 yr BP Were compiled and used to reconstruct palaeovegetation patterns, using complete taxon lists where possible and a biomization procedure that entailed the assignment of 645 pollen taxa to plant functional types. A set of 658 modern pollen samples spanning all biomes and regions provided a comprehensive test for this procedure and showed convincing agreement between reconstructed biomes and present natural vegetation types, both geographically and in terms of the elevation gradients in mountain regions of north-eastern and south-western China. The 6000 C-14 yr BP map confirms earlier studies in showing that the forest biomes in eastern China were systematically shifted northwards and extended westwards during the mid-Holocene. Tropical rain forest occurred on mainland China at sites characterized today by either tropical seasonal or broadleaved evergreen/warm mixed forest. Broadleaved evergreen/warm mixed forest occurred further north than today, and at higher elevation sites within the modern latitudinal range of this biome. The northern limit of temperate deciduous forest was shifted c. 800 km north relative to today. The 18,000 C-14 yr BP map shows that steppe and even desert vegetation extended to the modem coast of eastern China at the last glacial maximum, replacing today's temperate deciduous forest. Tropical forests were excluded from China and broadleaved evergreen/warm mixed forest had retreated to tropical latitudes, while taiga extended southwards to c. 43 degreesN.

Seven forms of rarity in mammals

Abstract: Conservation biologists have identified threats to the survival of about a quarter of the mammalian species; to identify patterns of rarity and commonness of mammals, we studied a global sample of 1212 species (about 28% of the mammals) using the ‘7 forms of rarity’ model (in which species are roughly divided into above and below the median for local population density, species’ range area, and number of habitat types). From a niche-based hypothesis of abundance and distribution, we predicted that mammals would exhibit a bimodal pattern of rarity and commonness, with an overabundance of species in the relatively rarest and most common categories; and just such a significant bimodal pattern emerged, with over a quarter of the species classified as exceedingly rare and a further quarter very common, supporting the niche-based hypothesis. Orders that include large mammals, including perissodactyls, primates, diprotodonts, and carnivores, exhibited significantly high proportions of relatively rare species; and tropical zoogeographic regions, especially Indomalaya, had relatively high proportions of species in the rarest category. Significant biases in the available data on mammals included under-sampling of small species like rodents and bats, and a relative paucity of data on zoogeographic regions outside of North America and Australia. Mammalian species listed as of conservation concern by the IUCN occurred in all cells of the model, indicating that even relatively common species can be listed as threatened under some conditions; but we also found that sixty-three species were relatively rare in all three criteria of the 7-forms model but were not listed as threatened, indicating potential candidates for further study. Mammals may be a group of animals where rarity or commonness is a natural aspect of species biology, both confirming and perhaps partly explaining the large proportion of mammals assigned threatened status.

Effects of fire protection on savanna structure in Central Brazil

Abstract: Aim Fire protection gradually changes the density of woody plants in numerous savannas around the world. In this study changes of structure in two tropical savanna areas with contrasting history of fire protection in the central Brazilian Cerrado is documented. Location Vegetation was sampled with line intercept transects in two adjacent sites in Brasilia, Federal District. These transects were located within a nature reserve protected from fire since 1972 and within an adjacent reserve area that burns every 2 years. Methods Five savanna physiognomies, from a low forest (‘cerradao’) to an open savanna (‘campo sujo’), were sampled in both sites. Results Fire protection increased the abundance of woody plants and favoured fire-sensitive species. With some exceptions, shrubs tended to be less affected by fire than trees. Species distribution was affected by a complex interaction of fire and physiognomy. Fire had the strongest effect on ‘campo sujo’ savanna, and a less significant effect on the intermediate physiognomies. Main conclusions Protection permits the establishment of fire sensitive species. A long enough protection against fire could lead to the appearance of more wooded physiognomies in the Cerrado.

Last glacial maximum biomes reconstructed from pollen and plant macrofossil data from northern Eurasia

Abstract: Pollen and plant macrofossil data from northern Eurasia were used to reconstruct the vegetation of the last glacial maximum (LGM: 18,000 ± 2000 14C yr bp) using an objective quantitative method for interpreting pollen data in terms of the biomes they represent ( Prentice et al., 1996 ). The results confirm previous qualitative vegetation reconstructions at the LGM but provide a more comprehensive analysis of the data. Tundra dominated a large area of northern Eurasia (north of 57°N) to the west, south and east of the Scandinavian ice sheet at the LGM. Steppe-like vegetation was reconstructed in the latitudinal band from western Ukraine, where temperate deciduous forests grow today, to western Siberia, where taiga and cold deciduous forests grow today. The reconstruction shows that steppe graded into tundra in Siberia, which is not the case today. Taiga grew on the northern coast of the Sea of Azov, about 1500 km south of its present limit in European Russia. In contrast, taiga was reconstructed only slightly south of its southern limit today in south-western Siberia. Broadleaved trees were confined to small refuges, e.g. on the eastern coast of the Black Sea, where cool mixed forest was reconstructed from the LGM data. Cool conifer forests in western Georgia were reconstructed as growing more than 1000 m lower than they grow today. The few scattered sites with LGM data from the Tien-Shan Mountains and from northern Mongolia yielded biome reconstructions of steppe and taiga, which are the biomes growing there today.

Trajectories to extinction: spatial dynamics of the contraction of geographical ranges

Abstract: Aim We examined the range contraction of 309 declining species of animals and plants to determine if the contraction dynamics better matched predictions based on the demographic characteristics of historical populations (demographic hypothesis) or based on the contagion-like spread of extinction forces (contagion hypothesis). Location Species included in the analysis came from all biogeographic regions. Methods We obtained range maps for 309 species from literature or through personal correspondence with authorities. Hypotheses were contrasted by examining the sequence of changes in the proportion (C) of the remnant range that fell within the central region of the historical range. Monte Carlo simulations and polynomial regressions were employed to examine changes in C during the process of range contraction. Results The results of the Monte Carlo simulations indicated that more species had observed range contractions consistent with the contagion hypothesis than expected by chance (z-score = 2.922, P = 0.002). The Monte Carlo analysis also indicated that the number of species whose observed range contractions were consistent with the demographic hypothesis was no greater than expected by chance (z-score = 0.337, P = 0.367). The results of the polynomial regression analysis for the two most common taxonomic groups (mammals and birds) and for all geographical regions (Australia, Africa, Eurasia, and North America) we examined also supported the contagion hypothesis. Main conclusions Most of the examined range contractions are consistent with the contagion hypothesis and that the most likely contagion is human related disturbance. These results have important implications for the conservation of endangered species.

Pollen‐based biome reconstruction for southern Europe and Africa 18,000 yr bp

Abstract: Pollen data from 18,000 C-14 yr sp were compiled in order to reconstruct biome distributions at the last glacial maximum in southern Europe and Africa. Biome reconstructions were made using the objective biomization method applied to pollen counts using a complete list of dryland taxa wherever possible. Consistent and major differences from present-day biomes are shown. F orest and xerophytic woods/scrub were replaced by steppe, both in the Mediterranean region and in southern Africa, except in south-western Cape Province where fynbos (xerophytic scrub) persisted. Sites in the tropical highlands, characterized today by evergreen forest, were dominated by steppe and/or xerophytic vegetation (cf. today's Ericaceous belt and Afroalpine grass land) at the last glacial maximum. Available data from the tropical lowlands are sparse but suggest that the modern tropical rain forest was largely replaced by tropical seasonal forest while the modern seasonal or dry forests were encroached on by savanna or steppe. Montane forest elements descended to lower elevations than today.

Cladistic biogeography of the Mexican transition zone

Abstract: Biogeographic relationships among nine montane areas of endemism across the transition zone between North and South America are analysed cladistically based on phylogenetic hypotheses of thirty-three resident monophyletic taxa of insects, fish, reptiles, and plants. Areas of endemism include the Arizona mountains (AZ), Sonoran Desert (SD), Sierra Madre Occidental (OCC), southern Sierra Madre Occidental (SOC), Sierra Madre Oriental (ORI), Sierra Transvolcanica (TRAN), Sierra Madre del Sur (SUR), Chiapan-Guatemalan Highlands (CGH), and Talamancan Cordillera (TC). Area relationships are summarized using Brooks Parsimony Analysis and Assumption 0, with the former resulting in more defensible biogeographic hypotheses. Areas of endemism are dividable into two monophyletic groups; a northern group including AZ, SD, OCC, and ORI, and a southern group consisting of TC, CGH, TRAN, SUR, and the isolated southern regions of the Sierra Madre Occidental (SOC). The northern set of areas are characterized by recent, probably Pleistocene, isolation and prevalent widespread species, whereas the southerly areas probably diverged after Pliocene closure of the Panamanian isthmus. The southern areas are redundantly represented on many of the taxon-area cladograms by endemic species, indicative of much higher levels of endemism in the Sierra Transvolcanica and further south. Use of a general area cladogram in such a transition zone permits explicit exploration of biogeographic patterns and establishes a predictive framework for taxonomy and conservation prioritization.

Fire regimes and forest changes in mid and upper montane forests of the southern Cascades, Lassen Volcanic National Park, California, U.S.A.

Abstract: Aim Spatial and temporal variation in fire regime parameters and forest structure were assessed. Location A 2630-ha area of mid- and upper montane forest in Lassen Volcanic National Park (LVNP). Methods Two hypotheses were tested concerned with fire-vegetation relationships in southern Cascades forests: (1) fire regime parameters (return interval, season of burn, fire size, rotation period) vary by forest dominant, elevation and slope aspect; and (2) fire exclusion since 1905 has caused forest structural and compositional changes in both mid- and upper montane forests. The implications of the study for national park management are also discussed. Results Fire regime parameters varied by forest compositional group and elevation in LVNP. Median composite and point fire return intervals were shorter in low elevation Jeffrey pine ( Pinus jeffreyi ) (JP) (4‐6 years, 16 years) and Jeffrey pine‐white fir ( Abies concolor ) (JP-WF) (5‐10 years, 22 years) and longer in high elevation red fir ( Abies magnifica )— western white pine ( Pinus monticola ) (RF-WWP) forests (9‐27 years, 70 years). Median fire return intervals were also shorter on east-facing (6‐9 years, 16.3 years) and longer on south- (11 years, 32.5 years) and west-facing slopes (22‐28 years, 54-years) in all forests and in each forest composition group. Spatial patterns in fire rotation length were the same as those for fire return intervals. More growing season fires also occurred in JP (33.1%) and JP-WF (17.5%) than in RF-WWP (1.1%) forests. A dramatic decline in fire frequency occurred in all forests after 1905. Conclusions Changes in forest structure and composition occurred in both mid- and upper montane forests due to twentieth-century fire exclusion. Forest density increased in JP and JP-WF forests and white fir increased in JP-WF forests and is now replacing Jeffrey pine. Forest density only increased in some RF-WWP stands, but not others. Resource managers restoring fire to these now denser forests need to burn larger areas if fire is going to play its pre-settlement role in montane forest dynamics.

Ecological biogeography of North American mammals: species density and ecological structure in relation to environmental gradients

Abstract: Aim To evaluate the relationship of climate and physiography to species density and ecological diversity of North American mammals. Location North America, including Mexico and Central America. Methods Species density, size structure and trophic structure of mammalian faunas and nine environmental variables were documented for quadrats covering the entire continent. Spatial autocorrelation of species density and the environmental variables illustrated differences in their spatial structure at the continental scale. We used principal component analysis to reduce the dimensionality of the climatic variables, linear multiple regression to determine which environmental variables best predict species density for the continent and several regions of the continent, and canonical ordination to evaluate how well the environmental variables predict ecological structure of mammalian faunas over North America. Results In the best regression model, five environmental variables, representing seasonal extremes of temperature, annual energy and moisture, and elevation, predicted 88% of the variation in species density for the whole continent. Among different regions of North America, the environmental variables that predicted species density vary. Changes in the size and trophic structure of mammalian faunas accompany changes in species density. Redundancy analysis demonstrated that environmental variables representing winter temperature, frostfree period, potential and actual evapotranspiration, and elevation account for 77% of the variation in ecological structure. Main conclusions The latitudinal gradient in mammalian species density is strong, but most of it is explained by variation in the environmental variables. Each ecological category peaks in species richness under particular environmental conditions. The changes of greatest magnitude involve the smallest size categories (< 10 g, 11–100 g), aerial insectivores and frugivores. Species in these categories, mostly bats, increase along a gradient of decreasing winter temperature and increasing annual moisture and frostfree period, trends correlated with latitude. At the opposite end of this gradient, species in the largest size category (101–1000 kg) increase in frequency. Species in size categories 3 (101–1000 g), 5 (11–100 kg) and 6 (101–1000 kg), herbivores, and granivores increase along a longitudinal gradient of increasing annual potential evapotranspiration and elevation. Much of the spatial pattern is consistent with ecological sorting of species ranges along environmental gradients, but differential rates of speciation and extinction also may have shaped the ecological diversity of extant North American mammals.

Plant dispersal in boreal rivers and its relation to the diversity of riparian flora.

Abstract: Aim The paper has four major objectives: (1) to determine whether diaspore mimics accurately represent dispersal dynamics of real diaspores in a free-flowing river; (2) to estimate distance travelled and reasons for stranding of floating diaspores along a free-flowing river; (3) to test if species composition and seedling recruitment vary with the ability of the riverbank to trap waterborne drift; and (4) to compare diaspore dispersal in a free-flowing river with that in a regulated river where current velocity has been reduced. Location The field work was conducted in two 7th-order boreal rivers in northern Sweden, the free-flowing Vindel River and the regulated Ume River. Methods We performed a series of dispersal experiments. We tested the usefulness wooden cubes of diaspore mimics for performing dispersal experiments by releasing cubes and achenes of Helianthus annuus and compare their dispersal patterns in the free-flowing Vindel River. We used the cubes to identify 50-m long sections along the river with different trapping capacity, i.e. the number of stranded diaspore mimics within a 50-m section. We then related the number of stranded diaspore mimics to the vascular plant flora, the proportions of species with long or short floating times (i.e. more than or less than 2 days, respectively), the number of seedlings, and to environmental variables within the sections. We also released wooden cubes in a run-of-river impoundment to determine the dispersal capacity of diaspores in a regulated river. Results The cubes were useful as diaspore mimics. They dispersed similarly to achenes of H. annuus. The stranding pattern of diaspore mimics was significantly associated with water current. Species richness of vascular plants per 50-m section increased with the number of stranded mimics. Seedling recruitment, and the proportions of species with short-floating and long-floating diaspores, did not vary with the number of stranded mimics. The ability of a river to transport diaspores downstream was strongly reduced by impoundment. Main conclusions We conclude that patterns of species richness of riparian vegetation is in part determined by the ability of the riverbank to trap waterborne diaspores, but differences in floating ability among species did not affect the species composition along free-flowing rivers.

Fires and land-cover change in the tropics: a remote sensing analysis at the landscape scale

Abstract: Aim Fires are an important landscape disturbance which interact in a complex way with land-use and land-cover change. The objective of this study is to understand the role of fires in vegetation-cover change and, conversely, the role of land use as a controlling factor of fires. Location The study sites are located in Mate Grosso, Brazil, in Central Africa and on the border between Kenya and Tanzania. Methods The role of vegetation fire is addressed through a landscape-scale analysis of the spatial association between maps of active fires and maps of land-cover change derived from remote sensing data for the different sites in Africa and South America. Results The empirical results of this study clearly support the idea that fires have widely varying impacts on land cover in savanna and forest ecosystems. Fires play different roles within the different components of landscape mosaics and at different times of the land-cover change trajectory. The impact of fires on vegetation is mainly controlled by land use. Conclusions There is thus a need to consider the socioeconomic purpose of biomass burning and the context in which such activities are undertaken. In forest ecosystems, a statistically significant relationship exists between the occurrence of fires and forest-cover changes. One could not conclude however, that fires are always the cause of the change in land cover, nor that fires are a reliable indicator of 'hot spots' of deforestation. Current low spatial resolution information on fire activity derived from remote sensing systems can be prone to inaccuracies due to a poor co-location of fire with respect to land-cover data, and temporal sampling problems affecting fire data.

Mammalian responses to Middle Holocene climatic change in the Great Basin of the western United States

Abstract: In spite of decades of intense research directed toward understanding the climates and ecology of the Great Basin (western United States) during the past 10,000 years, the responses of mammals to the extreme aridity of the Middle Holocene ( c. 8000‐ 5000 years ago) in this region have been poorly understood. Using a well-dated small mammal sequence from Homestead Cave, north-central Utah, I show that the Middle Holocene small mammal faunas of this area underwent a decrease in species richness and evenness, driven largely by a series of local extinctions and near-extinctions coupled with a dramatic increase in the abundance of taxa well-adapted to xeric conditions. At the end of this period, some taxa that require relatively mesic habitats began to increase in abundance immediately, others did not rebound in abundance until several thousand years later, while still others have not returned at all. This suite of responses has been difficult to detect because climatic change at the beginning of the Middle Holocene was so much more substantial than that which occurred toward its end.

Late Quaternary biomes of Canada and the eastern United States

Abstract: Pollen data have been used to construct biome maps for today, 6000 14 C yr bp and 18,000 14 C yr bp for Canada and the eastern United States. The inferred modern biome distributions agree well with independent reconstructions of North American vegetation prior to European settlement. Some discrepancies between the pollen data and the modern potential vegetation are caused by post-settlement clearing of the landscape and the consequent increase of herbaceous types in the recent pollen record. Biome distributions at 6000 14 C yr bp reflected the warmer and drier conditions then prevalent in the continental interior, but the overall position of biomes was similar to that of today. The boreal treeline in North America was not significantly north of its present position, in contrast to the 100‐200 km shift reported for Siberia. At the last glacial maximum (18,000 14 C yr bp ), steppe and tundra were prevalent in the Midwest and northwestern Canada, and coniferous forests and woodlands grew in eastern North America. The open vegetation at 18,000 14 C yr bp was probably due to drier conditions and/or lower concentrations of atmospheric CO 2 . The composition and physical structure of biomes is not constant over time. MidHolocene biomes were similar in structure to those of today, but shifts in the relative import ance of individual plant functional types are large enough that the physical properties of biomes, such as albedo, canopy conductance and surface roughness, are likely to have varied even during the Holocene. Last glacial maximum biomes were structurally different from their modern counterparts. The biome maps therefore may obscure significant vegetational changes in space and time during the late Quaternary. The difference between the highest and next highest affinity scores for each sample measures how strongly affinity scores discriminate among biomes. For many biomes, the difference is not large, and affinity score ties are not uncommon, highlighting the import ance of tie-break procedures when using the biomization method.

Pollen‐based biomes for Beringia 18,000, 6000 and 0 14C yr bp†

Abstract: The objective biomization method developed by Prentice et al. (1996) for Europe was extended using modern pollen samples from Beringia and then applied to fossil pollen data to reconstruct palaeovegetation patterns at 6000 and 18,000 14C yr bp. The predicted modern distribution of tundra, taiga and cool conifer forests in Alaska and north-western Canada generally corresponds well to actual vegetation patterns, although sites in regions characterized today by a mosaic of forest and tundra vegetation tend to be preferentially assigned to tundra. Siberian larch forests are delimited less well, probably due to the extreme under-representation of Larix in pollen spectra. The biome distribution across Beringia at 6000 14C yr bp was broadly similar to today, with little change in the northern forest limit, except for a possible northward advance in the Mackenzie delta region. The western forest limit in Alaska was probably east of its modern position. At 18,000 14C yr bp the whole of Beringia was covered by tundra. However, the importance of the various plant functional types varied from site to site, supporting the idea that the vegetation cover was a mosaic of different tundra types.

Species richness and endemism in the Western Australian flora.

Abstract: Aim Estimates of endemic and non-endemic native vascular plant species in each of the three Western Australian Botanical Provinces were made by East in 1912 and Beard in 1969. The present paper contains an updated assessment of species endemism in the State. Location Western Australia comprises one third of the continental Australian land mass. It extends from 13° to 35° S and 113° to 129° W. Methods Western Australia is recognized as having three Botanical Provinces (Northern, Eremaean and South-West) each divided into a number of Botanical Districts. Updated statistics for number of species and species endemism in each Province are based on the Census of Western Australian Plants data base at the Western Australian Herbarium (Western Australian Herbarium, 1998 onwards). Results The number of known species in Western Australia has risen steadily over the years but reputed endemism has declined in the Northern and Eremaean Provinces where cross-continental floras are common. Only the isolated South-West Province retains high rates of endemism (79%). Main conclusions With 5710 native species, the South-West Province contains about the same number as the California Floristic Province which has a similar area. The Italian mediterranean zone also contains about this number but in a smaller area, while the much smaller Cape Floristic Region has almost twice as many native species. The percentage of endemic species is highest at the Cape, somewhat less in south-western Australia and less again in California. Italy, at 12.5%, has the lowest value. Apart from Italy, it is usual for endemism to reach high values in the largest plant families. In Western Australia, these mainly include woody sclerophyll shrubs and herbaceous perennials with special adaptations to environmental conditions. While those life forms are prominent in the Cape, that region differs in the great importance of herbaceous families and succulents, both of which are virtually absent from Western Australia. In California and Italy, most endemics are in families of annual, herbaceous perennial and soft shrub plants. It is suggested that the dominant factor shaping the South-West Province flora is the extreme poverty of the area’s soils, a feature that emphasizes sclerophylly, favours habitat specialization and ensures relatively many local endemic species.

The diversity and abundance of North American butterflies vary with habitat disturbance and geography

Abstract: Aim We used data from the annual Fourth of July Butterfly Count for the years 1989–97 to examine patterns of species richness and total butterfly abundance across North America and within topographically diverse and disturbed landscapes. Location We analysed counts from 514 different locations in North America. The counts represent all areas of the USA and southern Canada, with a few Mexican sites as well, although most counts were in the eastern USA. Methods First, we standardized published count data according to the effort expended per count (total party-hours). Using regression analysis and analysis of variance, we then examined the impact of latitude, longitude, topographical relief, habitat disturbance and different climatic measures on the species richness and total abundance of butterflies per count. We also examined the abundance of exotic species in disturbed landscapes. Results Our analyses suggest that: (1) species richness is highest at low latitudes and near Rocky Mountain longitudes; (2) the total abundance of individuals is highest in northern US latitudes and Great Plains longitudes; (3) species richness but not total abundance increases with greater topographical relief; (4) species richness and diversity indices are lower in more disturbed habitats; and (5) the abundance of the introduced Pieris rapae (L.) is greater in more disturbed habitats. Main conclusions Different factors control the abundance and species richness of North American butterflies. Along with geographical location, habitat disturbance and topographical variability affect species richness. Our analysis also shows the value of broad-based monitoring regimes, such as the North American Fourth of July Butterfly Count.

Isozyme variation and recent biogeographical history of the long-lived conifer Fitzroya cupressoides

Abstract: Aim Palaeoenvironmental records of Pleistocene glaciation and associated vegetation changes in Patagonia have led to the hypothesis that during the last glacial maximum (LGM) tree species survived locally in favourable habitats. If present populations originated from spread from only one refugium, such as an ice-free area of coastal Chile (Single Refugium hypothesis), we would expect that eastern populations would be genetically depauperate and highly similar to western populations. In contrast, if the ice cap was not complete and tree species persisted in forest patches on both slopes of the Andes (Multiple Refugia hypothesis), we would expect a greater degree of genetic divergence between populations either on opposite sides of the Cordillera (Cordillera Effect scenario) or towards its present-day southern distributional limit where the ice sheet reached its maximum coverage (Extent-of-the-Ice scenario). Location We tested this refugia hypothesis using patterns of isozyme variation in populations sampled over the entire modern range of the endemic conifer Fitzroya cupressoides (Mol.) Johnst. (Cupressaceae) in temperate South America. Methods Fresh foliage was collected from twenty-four populations and analysed by horizontal electrophoresis on starch gels. Results Twenty-one putative loci were reliably scored and 52% were polymorphic in at least one population. Populations from the eastern slope of the Andes were genetically more variable than those from the western slope; the former had a greater mean number of alleles per locus, a larger total number of alleles and rare alleles, and higher polymorphism. Genetic identities within western populations were greater than within eastern populations. Discriminant analyses using allelic frequencies of different grouping schedules of populations were non significant when testing for the Single Refugium hypothesis whereas significant results were obtained for the Multiple Refugia hypothesis. Main conclusions Our results indicate that present Fitzroya populations are the result of spreading from at least two, but possibly more, glacial refugia located in Coastal Chile and on the southern flanks of the Andes in Argentina.

Patterns of distribution for southern Australian marine echinoderms and decapods

Abstract: Aim To relate patterns of distribution of marine echinoderms and decapods around southern Australia to major ecological and historical factors. Location Shallow-water (0–100 m) marine waters off southern Australia, south of 30° S. Methods (1) Record the presence/absence of known echinoderm and decapod species in cells of c. 1° latitude and longitude, along the coast of southern mainland Australia and Tasmania. (2) Describe patterns in species composition, species richness and endemism through gradient analysis, ordination and cluster analysis. (3) Relate these patterns to distance and temperature gradients, the area of continental shelf, the average size of species range, and known historical factors. Results Species composition varied with both latitude and longitude. Species richness was relatively constant from east to west but graded with latitude from high in the warm-temperate regions around Perth and Sydney to low in cool-temperate southern Tasmania. Species richness was not related to the area of continental shelf or average species range size. Species turnover was not correlated with rates of temperature change. It was problematic to separate distance from temperature gradients, but there was evidence that the southern distribution limits of some species are related to minimum sea surface temperature. Within the taxonomic groups surveyed, evolutionary radiation has been largely limited to a few cosmopolitan species-rich genera. Main conclusions There are historical as well as ecological hypotheses explaining the latitudinal gradient of marine species richness in southern Australia: (1) the continual invasion and speciation of species of tropical origin as Australia has split from Gondwana and drifted northward; (2) progressive extinction of some Gondwanan cool-temperate species at the limits of their range; (3) low level of immigration of additional cool-temperate species; and (4) some in situ endemic speciation.

Using between-model comparisons to fine-tune linear models of species ranges.

Abstract: Aim To offer an objective approach to some of the problems associated with the development of logistic regression models: how to compare different models, determination of sample size adequacy, the influence of the ratio of positive to negative cells on model accuracy, and the appropriate scale at which the hypothesis of a non-random distribution should be tested. Location Test data were taken from Southern Africa. Methods The approach relies mainly on the use of the AUC (Area under the Curve) statistic, based on ROC (threshold Receiver Operating Characteristic) plots, for between-model comparisons. Data for the distribution of the bont tick Amblyomma hebraeum Koch (Acari: Ixodidae) are used to illustrate the methods. Results Methods for the estimation of minimum sample sizes and more accurate hypothesis-testing are outlined. Logistic regression is robust to the assumption that uncollected cells can be scored as negative, provided that the sample size of cells scored as positive is adequate. The variation in temperature and rainfall at localities where A. hebraeum has been collected is significantly lower than expected from a random sample of points across the data set, suggesting that within-site variation may be an important determinant of its distribution. Main conclusions Between-model comparisons relying on AUCs can be used to enhance objectivity in the development and refinement of logistic regression models. Both between-site and within-site variability should be considered as potentially important factors determining species distributions.

Californian mixed-conifer forests under unmanaged fire regimes in the Sierra San Pedro Mártir, Baja California, Mexico.

Abstract: Aim This study appraises historical fire regimes for Californian mixed-conifer forests of the Sierra San Pedro Martir (SSPM). The SSPM represents the last remaining mixed-conifer forest along the Pacific coast still subject to uncontrolled, periodic ground fire. Location The SSPM is a north–south trending fault bound range, centred on 31°N latitude, 100 km SE of Ensenada, Baja California. Methods We surveyed forests for composition, population structure, and historical dynamics both spatially and temporally over the past 65 years using repeat aerial photographs and ground sampling. Fire perimeter history was reconstructed based on time-series aerial photographs dating from 1942 to 1991 and interpretable back to 1925. A total of 256 1-ha sites randomly selected from aerial photographs were examined along a chronosequence for density and cover of canopy trees, density of snags and downed logs, and cover of non-conifer trees and shrubs. Twenty-four stands were sampled on-the-ground by a point-centred quarter method which yielded data on tree density, basal area, frequency, importance value, and shrub and herb cover. Results Forests experience moderately intense understory fires that range in size to 6400 ha, as well as numerous smaller, low intensity burns with low cumulative spatial extent. SSPM forests average 25–45% cover and 65–145 trees per ha. Sapling densities were two to three times that of overstory trees. Size-age distributions of trees ≥ 4 cm dbh indicate multi-age stands with steady-state dynamics. Stands are similar to Californian mixed conifer forests prior to the imposition of fire suppression policy. Livestock grazing does not appear to be suppressing conifer regeneration. Main conclusions Our spatially-based reconstruction shows the open forest structure in SSPM to be a product of infrequent, intense surface fires with fire rotation periods of 52 years, rather than frequent, low intensity fires at intervals of 4–20 years proposed from California fire-scar dendrochronology (FSD) studies. Ground fires in SSPM were intense enough to kill pole-size trees and a significant number of overstory trees. We attribute long fire intervals to the gradual build-up of subcontinuous shrub cover, conifer recruitment and litter accumulation. Differences from photo interpretation and FSD estimates are due to assumptions made with respect to site-based (point) sampling of fire, and nonfractal fire intensities along fire size frequency distributions. Fire return intervals determined by FSD give undue importance to local burns which collectively use up little fuel, cover little area, and have little demographic impact on forests.

Molecular evidence for dispersal rather than vicariance as the origin of flightless insect species on the Chatham Islands, New Zealand

Abstract: Aim The aim was to use mitochondrial DNA sequence data to test between vicariance and oversea dispersal explanations for the origin of the Chatham Islands biota. Location New Zealand and the Chatham Islands, separated by c. 800 km in the south-west Pacific Ocean. Methods DNA sequences from the mitochondrial gene cytochrome oxidase I (COI) were obtained from four genera of relatively large and flightless insects (Coleoptera— Geodorcus, Mecodema; Orthoptera—Talitropsis; Blattoidea—Celatoblatta). These were used to test alternative hypotheses for the origin of the Chatham taxa. Results Phylogenetic analysis revealed the Chatham taxa in each genus to be monophyletic. Genetic distances exhibited by these genera, between taxa found on the Chatham Islands and mainland New Zealand were relatively low (11.2, 2.8, 3.0 and 4.9%, respectively). Main conclusions Even allowing for variation in molecular evolutionary rates, these genetic distances indicate phylogenetic separation of New Zealand and Chatham insect lineages in the Pliocene (2–6 Ma). Such dates are more than one order of magnitude too recent to be explained by vicariant (tectonic) processes. Oversea dispersal from New Zealand to the Chatham Islands is implicated and this conclusion is in keeping with the taxonomy of the endemic avifauna, flora and fossil molluscan fauna.

Centrifugal speciation and centres of origin

Abstract: Aim To discuss the development and usefulness of the theory of centrifugal speciation and its practical application to the centre of origin hypothesis. Location The Indo-West Pacific Ocean. Methods Utilization of patterns demonstrating species diversity, generic age, dispersal tracks, phylogenetics, extinction and genetic diversity. Results The centrifugal speciation hypothesis appears to provide a suitable explanation for the way in which centres of origin theoretically operate. Recent information about the centre of origin in the marine East Indies suggests that it is the source of evolutionary radiation for the Indo-West Pacific. Its mechanism appears to conform to the centrifugal prediction. Main conclusions The East Indies Triangle is the origin of a series of dynamic systems that extend across the entire Indo-West Pacific. These systems are apparently maintained by a continuous outflow from the centre of origin. A series of species replacements from the East Indies over the past 10 million years would create and perpetuate the systems that have been identified.

Characterization of the spatio‐temporal patterns of global fire activity using satellite imagery for the period April 1992 to March 1993

Abstract: Aim This paper describes the characteristics of the spatio-temporal distribution of vegetation fires as detected from satellite data for the 12 months April 1992 to March 1993. Location Fires are detected daily at a spatial resolution of 1 km for all land areas of the globe. Methods From the fire location information a daily gridded product at 0.5∞ by 0.5∞ has been constructed. Two methods of characterizing the spatio-temporal pattern of vegetation fires are discussed. The first applies empirical orthogonal function analysis to the monthly series of gridded data. The second approach defines and extracts a number of spatial and temporal parameters from the gridded product. The descriptive parameters extracted are used in a cluster analysis in order to group cells with similar characteristics into a small number of classes. Results Using daily global satellite observations, it is possible to characterize the spatial and temporal variability in fire activity. Most of this variability is within the tropical belt, where the majority of fire activity is concentrated, nonetheless fire was also detected in temperate and boreal regions. The period in which fire occurred varied from region to region. Parameterization provided a very synthetic view of this variability facilitating regional intercomparison. Clustering identifies five classes of fire activity, each of which can be associated with particular climatic conditions, vegetation types and land-use. Main conclusions Global monitoring of vegetation fire from satellite is possible. The analysis provides a coherent, consistent and synoptic view of global fire activty with one data set. The type of information extracted can be of use in global atmospheric chemistry modelling and for studying the role of fire in relation to global change issues.

A fire history of a subalpine forest in south‐eastern Wyoming, USA

Abstract: Fire history was determined for part of the Routt-Medicine Bow National Forest in south-eastern Wyoming using fire-scar and age-class analysis. A composite chronology of fire events was used to determine mean fire intervals (MFI) for pre-EuroAmerican settlement, EuroAmerican settlement (before 1868 AD), EuroAmerican settlement and modern (after 1912) periods, for all fires and stand-replacing fires. Point-scale MFI was also determined using grand means from individual trees. Stand-replacing fires were reconstructed to determine fire rotation. MFI for the entire time period is 5.5-8.4 years. MFI decreased from 9.3 to 15.7-1.9-2.9 years from the preto post-EuroAmerican settlement periods, and increased during the modern period. Point-scale MFIs are longer than MFI of the study area. Fire rotation is 182 years for the total period of record, but increased from 127 years during the pre-EuroAmerican settlement period to 170 years during the EuroAmerican settlement period. Fire rotation during the modern period dramatically increased to 27,035 years. Results suggest fire suppression may have influenced the fire regime. Comparison of regional fire events with fire events from this study indicate regional weather has an important influence on Rocky Mountain fire regimes.