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USGS2021

Gulf Coast vicariance shapes phylogeographic history of a North American freshwater mussel species complex

Aim Freshwater mussels share habitat and are parasites of freshwater fishes during the larval life stage. Therefore, models of fish biogeography may also explain the historical biogeography of freshwater mussels. We tested this assumption using predictions of three biogeographic models constructed for northern Gulf of Mexico drainages on a freshwater mussel species complex. Specifically, we tested (1) if speciation was due to vicariant events of fluctuating sea levels that separated lineages east‐west of the Mobile Basin (Central Gulf Coast speciation hypothesis), (2) if the timing of divergences occurred 8.5–3.5 MYA (Gulf Coast allopatric speciation model) and (3) if diversification in Mississippi River populations was recent and for evidence of population increase consistent with range expansion into northern deglaciated regions (Pleistocene glaciation model). Location Eastern North America. Taxon Freshwater mussels (Bivalvia: Unionidae), Lampsilis teres and L. floridensis . Methods We collected 249 specimens from 73 localities across the group's distribution. We used three molecular markers (COI, NDI & ITSI) to conduct time calibrated Bayesian phylogenetic analyses, phylogeographic analyses (AMOVA & SAMOVA) and demographic analyses including Bayesian skyline plots. Results Lampsilis teres and L. floridensis are allopatric species whose distributions meet at the eastern edge of the Mobile Basin. Speciation was estimated to occur in the late Miocene. Populations from isolated river systems surrounding the Gulf of Mexico are almost all monophyletic. Mississippi drainage samples formed a shallow clade with recent diversification and showed evidence of recent population expansion. Main conclusions The historical biogeography of the L. teres species complex is broadly consistent with tested ichthyofaunal models. The timing of speciation and intraspecific divergences correspond to low sea‐level events suggesting that Gulf Coast sea‐level fluctuations are responsible for dispersal (sea‐level recession) and subsequent cladogenesis (sea‐level inundation). Consistent with numerous other freshwater studies, we found the Mobile Basin to be a suture zone, which may be due to the narrow, offshore continental shelf.

Journal of Biogeography↗

USGS1989

Biogeography of marine podocopid Ostracodes in Micronesia

Shallow-water podocopic marine Ostracoda from Micronesian lagoons in the Marianas, Caroline, Marshall and Gilbert Islands in the western Pacific were analysed to establish their diversity and zoogeography within Micronesia and the taxonomic affinities with ostracodes from other tropical regions. Sixty-four bottom lagoonal sediment samples from twelve islands and atolls yielded more than seventy species representing over thirty-two ostracode genera. Q-mode cluster analysis using Jaccard coefficients showed that, with few exceptions, all or most samples from a particular lagoon form distinct subgroups (Jaccard=0.45-0.50). At lower levels, five groups delineate faunal regions within Micronesia: the Gilbert Islands (Onotoa) in the southeast part of the region, the northern Marshall Islands (Enewetak, Rongelap, Bikini), the southestern Marshall Islands (Kwajalein, Jaluit, Majuro, Arno), the Marianas and Caroline Islands (Guam, Truk, Pohnpei) and Pingelap. Patterns of species diversity show Guam, Truk, Pohnpei, Pingelap, Kwajalein and Onotoa have the highest species richness (S=32-42) and Shannon-Wiener diversity values (H(S)=2.62-3.02) in the study area. Enewetak, Jaluit, Majuro and Arno show lower values (S=23-27, H(S)=2.29-2.70). Of the ostracode species living in Micronesia, 64.3% have Indo-West Pacific affinities, 7.1% are circumtropical, 5.7% have East Pacific-Caribbean affinities, 11.4% are endemic to Micronesia, and 11.4% have unknown affinities. If the southeast Asian region is a primary species-source, the results show that each Micronesian lagoon is equally likely to be colonized by dispersal from the source region, despite differences in distance from a hypothetical source. However, each lagoon has a distinct ostracode assemblage, probably the result of unique history of random colonization events, local extinctions and environmental disturbances.

Journal of Biogeography↗

USGS2021

Palaeoceanographic changes in the late Pliocene promoted rapid diversification in pelagic seabirds

Aim Palaeoceanographic changes can act as drivers of diversification and speciation, even in highly mobile marine organisms. Shearwaters are a group of globally distributed and highly mobile pelagic seabirds. Despite a recent well-resolved phylogeny, shearwaters have controversial species limits, and show periods of both slow and rapid diversification. Here, we explore the role of palaeoceanographic changes on shearwaters' diversification and speciation. We investigate shearwater biogeography and the evolution of a key phenotypic trait, body size, and we assess the validity of their current taxonomy. Location Worldwide. Taxa Shearwaters (Order Procellariiformes, Family Procellariidae, Genera Ardenna , Calonectris and Puffinus ). Methods We generated genomic (ddRAD) data to infer a time-calibrated species tree for the shearwaters. We estimated ancestral ranges and evaluated the roles of founder events, vicariance and surface ocean currents in driving diversification. We performed phylogenetic generalised least squares to identify potential predictors of variability in body size along the phylogeny. To assess the validity of the current taxonomy, we analysed genomic patterns of recent shared ancestry and differentiation among shearwater taxa. Results We identified a period of high dispersal and rapid speciation during the Late Pliocene–early Pleistocene. Species dispersal appears to be favoured by surface ocean currents, and founder events are supported as the main mode of speciation in these highly mobile pelagic seabirds. Body mass shows significant associations with life strategies and local conditions. The current taxonomy shows some incongruences with the patterns of genomic divergence. Main Conclusions A reduction of neritic areas during the Pliocene seems to have driven global extinctions of shearwater species, followed by a subsequent burst of speciation and dispersal probably promoted by Plio-Pleistocene climatic shifts. Our findings extend our understanding on the drivers of speciation and dispersal of highly mobile pelagic seabirds and shed new light on the important role of palaeoceanographic events.

Journal of Biogeography↗

USGS2021

Abiotic stress and biotic factors mediate range dynamics on opposing edges

Aim In the face of global change, understanding causes of range limits are one of the most pressing needs in biogeography and ecology. A prevailing hypothesis is that abiotic stress forms cold (upper latitude/altitude) limits, whereas biotic interactions create warm (lower) limits. A new framework – Interactive Range-Limit Theory (iRLT) – asserts that positive biotic factors such as food availability can ameliorate abiotic stress along cold edges, whereas abiotic stress can have a positive effect and mediate biotic interactions (e.g., competition) along warm limits. Location Northeastern United States Taxon Carnivora Methods We evaluated two hypotheses of iRLT using occupancy and structural equation modeling (SEM) frameworks based on data collected over a 6-year period (2014–2019) of six carnivore species across a broad latitudinal (42.8–45.3°N) and altitudinal (3–1451 m) gradient. Results We found that snow directly limits populations, but prey or habitat availability can influence range dynamics along cold edges. For example, bobcats ( Lynx rufus ) and coyotes ( Canis latrans ) were limited by deep snow and long winters, but the availability of prey had a strong positive effect. Conversely, snow had a strong positive effect on the warm limits of Canada lynx ( Lynx canadensis ), countering the negative effect of competition with the phylogenetically similar bobcat and with coyotes, highlighting how climate mediates competition between species. Main conclusions We used an integrated dataset that included competitors and prey species collected at the same spatial and temporal scale. As such, this design, along with a causal modeling framework (SEM), allowed us to evaluate community-wide hypotheses at macroecological scales and identify coarse-scale drivers of species' range limits. Our study supports iRLT and underscores the need to consider direct and indirect mechanisms for studying range dynamics and species' responses to global change.

New Hampshire, Vermont↗

USGS2009

Biogeography of Iberian freshwater fishes revisited: The roles of historical versus contemporary constraints

Aim The question of how much of the shared geographical distribution of biota is due to environmental vs. historical constraints remains unanswered. The aim of this paper is to disentangle the contribution of historical vs. contemporary factors to the distribution of freshwater fish species. In addition, it illustrates how quantifying the contribution of each type of factor improves the classification of biogeographical provinces. Location Iberian Peninsula, south-western Europe ( c . 581,000 km 2 ). Methods We used the most comprehensive data on native fish distributions for the Iberian Peninsula, compiled from Portuguese and Spanish sources on a 20-km grid-cell resolution. Overall, 58 species were analysed after being categorized into three groups according to their ability to disperse through saltwater: (1) species strictly intolerant of saltwater (primary species); (2) species partially tolerant of saltwater, making limited incursions into saltwaters (secondary species); and (3) saltwater-tolerant species that migrate back and forth from sea to freshwaters or have invaded freshwaters recently (peripheral species). Distance-based multivariate analyses were used to test the role of historical (basin formation) vs. contemporary environmental (climate) conditions in explaining current patterns of native fish assemblage composition. Cluster analyses were performed to explore species co-occurrence patterns and redefine biogeographical provinces based on the distributions of fishes. Results River basin boundaries were better at segregating species composition for all species groups than contemporary climate variables. This historical signal was especially evident for primary and secondary freshwater fishes. Eleven biogeographical provinces were delineated. Basins flowing to the Atlantic Ocean north of the Tagus Basin and those flowing to the Mediterranean Sea north of the Mijares Basin were the most dissimilar group. Primary and secondary freshwater species had higher province fidelity than peripheral species. Main conclusions The results support the hypothesis that historical factors exert greater constraints on native freshwater fish assemblages in the Iberian Peninsula than do current environmental factors. After examining patterns of assemblage variation across space, as evidenced by the biogeographical provinces, we discuss the likely dispersal and speciation events that underlie these patterns.

Iberian Peninsula↗

USGS2006

Classification tree and minimum-volume ellipsoid analyses of the distribution of ponderosa pine in the western USA

Aim? Ponderosa pine (Pinus ponderosa Douglas ex Lawson & C. Lawson) is an economically and ecologically important conifer that has a wide geographic range in the western USA, but is mostly absent from the geographic centre of its distribution - the Great Basin and adjoining mountain ranges. Much of its modern range was achieved by migration of geographically distinct Sierra Nevada (P. ponderosa var. ponderosa) and Rocky Mountain (P. ponderosa var. scopulorum) varieties in the last 10,000 years. Previous research has confirmed genetic differences between the two varieties, and measurable genetic exchange occurs where their ranges now overlap in western Montana. A variety of approaches in bioclimatic modelling is required to explore the ecological differences between these varieties and their implications for historical biogeography and impending changes in western landscapes. Location? Western USA. Methods? We used a classification tree analysis and a minimum-volume ellipsoid as models to explain the broad patterns of distribution of ponderosa pine in modern environments using climatic and edaphic variables. Most biogeographical modelling assumes that the target group represents a single, ecologically uniform taxonomic population. Classification tree analysis does not require this assumption because it allows the creation of pathways that predict multiple positive and negative outcomes. Thus, classification tree analysis can be used to test the ecological uniformity of the species. In addition, a multidimensional ellipsoid was constructed to describe the niche of each variety of ponderosa pine, and distances from the niche were calculated and mapped on a 4-km grid for each ecological variable. Results? The resulting classification tree identified three dominant pathways predicting ponderosa pine presence. Two of these three pathways correspond roughly to the distribution of var. ponderosa, and the third pathway generally corresponds to the distribution of var. scopulorum. The classification tree and minimum-volume ellipsoid model show that both varieties have very similar temperature limitations, although var. ponderosa is more limited by the temperature extremes of the continental interior. The precipitation limitations of the two varieties are seasonally different, with var. ponderosa requiring significant winter moisture and var. scopulorum requiring significant summer moisture. Great Basin mountain ranges are too cold at higher elevations to support either variety of ponderosa pine, and at lower elevations are too dry in summer for var. scopulorum and too dry in winter for var. ponderosa. Main conclusions? The classification tree analysis indicates that var. ponderosa is ecologically as well as genetically distinct from var. scopulorum. Ecological differences may maintain genetic separation in spite of a limited zone of introgression between the two varieties in western Montana. Two hypotheses about past and future movements of ponderosa pine emerge from our analyses. The first hypothesis is that, during the last glacial period, colder and/or drier summers truncated most of the range of var. scopulorum in the central Rockies, but had less dramatic effects on the more maritime and winter-wet distribution of var. ponderosa. The second hypothesis is that, all other factors held constant, increasing summer temperatures in the future should produce changes in the distribution of var. scopulorum that are likely to involve range expansions in the central Rockies with the warming of mountain ranges currently too cold but sufficiently wet in summer for var. scopulorum. Finally, our results underscore the growing need to focus on genotypes in biogeographical modelling and ecological forecasting.

Journal of Biogeography↗

USGS2003

Actual evapotranspiration and deficit: Biologically meaningful correlates of vegetation distribution across spatial scales

Correlative approaches to understanding the climatic controls of vegetation distribution have exhibited at least two important weaknesses: they have been conceptually divorced across spatial scales, and their climatic parameters have not necessarily represented aspects of climate of broad physiological importance to plants. Using examples from the literature and from the Sierra Nevada of California, I argue that two water balance parameters—actual evapotranspiration (AET) and deficit (D)—are biologically meaningful, are well correlated with the distribution of vegetation types, and exhibit these qualities over several orders of magnitude of spatial scale (continental to local). I reach four additional conclusions. (1) Some pairs of climatic parameters presently in use are functionally similar to AET and D; however, AET and D may be easier to interpret biologically. (2) Several well-known climatic parameters are biologically less meaningful or less important than AET and D, and consequently are poorer correlates of the distribution of vegetation types. Of particular interest, AET is a much better correlate of the distributions of coniferous and deciduous forests than minimum temperature. (3) The effects of evaporative demand and water availability on a site's water balance are intrinsically different. For example, the ‘dry’ experienced by plants on sunward slopes (high evaporative demand) is not comparable to the ‘dry’ experienced by plants on soils with low water-holding capacities (low water availability), and these differences are reflected in vegetation patterns. (4) Many traditional topographic moisture scalars—those that additively combine measures related to evaporative demand and water availability—are not necessarily meaningful for describing site conditions as sensed by plants; the same holds for measured soil moisture. However, using AET and D in place of moisture scalars and measured soil moisture can solve these problems.

Journal of Biogeography↗

USGS2001

Forest area and distribution in the Mississippi alluvial valley: Implications for breeding bird conservation

Knowing the current forest distribution and patch size characteristics is integral to the development of geographically defined, habitat-based conservation objectives for breeding birds. Towards this end, we classified 2.6 million ha of forest cover within the Mississippi Alluvial Valley using 1992 thematic mapper satellite imagery. Although historically this area, from southern Illinois to southern Louisiana, was dominated by forested wetlands, forest cover remains on less than 25% of the floodplain. Remaining forest cover is comprised of > 38,000 discrete forest patches > 2 ha. Mean patch area (64.1±5.2 ha; x̄ ±SE) was highly skewed towards small fragment size. Larger patches had a higher proportion of more hydric forest cover classes than did smaller patches which had a higher proportion of less hydric forest cover classes. Public lands accounted for 16% of remaining forested wetlands. Fewer than 100 forest patches exceeded our hypothesized habitat objective (4000 ha minimum contiguous forest area) intended to support self-sustaining populations of forest breeding birds. To increase the number of forest patches exceeding 4000 ha contiguous area, and thereby increase the likelihood of successful forest bird conservation, we recommend afforestation adjoining existing forest fragments ≥1012 ha and focused within designated Forest Bird Conservation Regions.

Journal of Biogeography↗

USGS2001

Partitioning sources of variation in vertebrate species richness

Aim: To explore biogeographic patterns of terrestrial vertebrates in Maine, USA using techniques that would describe local and spatial correlations with the environment. Location: Maine, USA. Methods: We delineated the ranges within Maine (86,156 km2) of 275 species using literature and expert review. Ranges were combined into species richness maps, and compared to geomorphology, climate, and woody plant distributions. Methods were adapted that compared richness of all vertebrate classes to each environmental correlate, rather than assessing a single explanatory theory. We partitioned variation in species richness into components using tree and multiple linear regression. Methods were used that allowed for useful comparisons between tree and linear regression results. For both methods we partitioned variation into broad-scale (spatially autocorrelated) and fine-scale (spatially uncorrelated) explained and unexplained components. By partitioning variance, and using both tree and linear regression in analyses, we explored the degree of variation in species richness for each vertebrate group that Could be explained by the relative contribution of each environmental variable. Results: In tree regression, climate variation explained richness better (92% of mean deviance explained for all species) than woody plant variation (87%) and geomorphology (86%). Reptiles were highly correlated with environmental variation (93%), followed by mammals, amphibians, and birds (each with 84-82% deviance explained). In multiple linear regression, climate was most closely associated with total vertebrate richness (78%), followed by woody plants (67%) and geomorphology (56%). Again, reptiles were closely correlated with the environment (95%), followed by mammals (73%), amphibians (63%) and birds (57%). Main conclusions: Comparing variation explained using tree and multiple linear regression quantified the importance of nonlinear relationships and local interactions between species richness and environmental variation, identifying the importance of linear relationships between reptiles and the environment, and nonlinear relationships between birds and woody plants, for example. Conservation planners should capture climatic variation in broad-scale designs; temperatures may shift during climate change, but the underlying correlations between the environment and species richness will presumably remain.

Journal of Biogeography↗

USGS2001

Pollen-based biomes for Beringia 18,000, 6000 and 0 14C yr BP

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.

Journal of Biogeography↗

USGS2001

Biomes of western North America at 18,000, 6000 and 0 14C yr BP reconstructed from pollen and packrat midden data

A new compilation of pollen and packrat midden data from western North America provides a refined reconstruction of the composition and distribution of biomes in western North America for today and for 6000 and 18,000 radiocarbon years before present (14C yr BP). Modern biomes in western North America are adequately portrayed by pollen assemblages from lakes and bogs. Forest biomes in western North America share many taxa in their pollen spectra and it can be difficult to discriminate among these biomes. Plant macrofossils from packrat middens provide reliable identification of modern biomes from arid and semiarid regions, and this may also be true in similar environments in other parts of the world. However, a weighting factor for trees and shrubs must be used to reliably reconstruct modern biomes from plant macrofossils. A new biome, open conifer woodland, which includes eurythermic conifers and steppe plants, was defined to categorize much of the current and past vegetation of the semiarid interior of western North America. At 6000 14C yr BP, the forest biomes of the coastal Pacific North-west and the desert biomes of the South-west were in near-modern positions. Biomes in the interior Pacific North-west differed from those of today in that taiga prevailed in modern cool/cold mixed forests. Steppe was present in areas occupied today by open conifer woodland in the northern Great Basin, while in the central and southern Rocky Mountains forests grew where steppe grows today. During the mid-Holocene, cool conifer forests were expanded in the Rocky Mountains (relative to today) but contracted in the Sierra Nevada. These differences from the forests of today imply different climatic histories in these two regions between 6000 14C yr BP and today. At 18,000 14C yr BP, deserts were absent from the South-west and the coverage of open conifer woodland was greatly expanded relative to today. Steppe and tundra were present in much of the region now covered by forests in the Pacific North-west.

Journal of Biogeography↗

USGS2008

GIS interpolations of witness tree records (1839-1866) for northern Wisconsin at multiple scales

To construct forest landscape of pre-European settlement periods, we developed a GIS interpolation approach to convert witness tree records of the U.S. General Land Office (GLO) survey from point to polygon data, which better described continuously distributed vegetation. The witness tree records (1839-1866) were processed for a 3-million ha landscape in northern Wisconsin, U.S.A. at different scales. We provided implications of processing results at each scale. Compared with traditional GLO mapping that has fixed mapping scales and generalized classifications, our approach allows presettlement forest landscapes to be analysed at the individual species level and reconstructed under various classifications. We calculated vegetation indices including relative density, dominance, and importance value for each species, and quantitatively described the possible outcomes when GLO records are analysed at three different scales (resolution). The 1 x 1-section resolution preserved spatial information but derived the most conservative estimates of species distributions measured in percentage area, which increased at coarser resolutions. Such increases under the 2 x 2-section resolution were in the order of three to four times for the least common species, two to three times for the medium to most common species, and one to two times for the most common or highly contagious species. We marred the distributions of hemlock and sugar maple from the pre-European settlement period based on their witness tree locations and reconstructed presettlement forest landscapes based on species importance values derived for all species. The results provide a unique basis to further study land cover changes occurring after European settlement.

Wisconsin↗

USGS2008

Richness and diversity of helminth communities in tropical freshwater fishes: Empirical evidence

Aim: Published information on the richness and diversity of helminth parasite communities in tropical freshwater fishes is reviewed in response to expectations of species-rich parasite communities in tropical regions. Location: Areas covered include the tropics and some subtropical areas. In addition, the north temperate area of the nearctic zone is included for comparison. Methods: Data from 159 communities in 118 species of tropical freshwater fish, summarized from 46 published studies, were used for this review. Parasite community descriptors used in the analyses included component community richness and calculated mean species richness. Data from 130 communities in 47 species of nearctic north temperate freshwater fish were summarized from 31 studies and used for comparison. Results: The component helminth communities of many tropical freshwater fish are species-poor, and considerable proportions of fish from certain parts of the tropics, e.g. West African drainages, are uninfected or lightly infected. Mean helminth species richness was low and equaled or exceeded 2.0 in only 22 of 114 communities. No single group of helminths was identified as a dominant component of the fauna and species composition was variable among and within broader geographical areas. The richest enteric helminth assemblages were found in mochokid and clariid catfish with a mixed carnivorous diet, whereas algal feeders, herbivores and detritivores generally had species-poor gut helminth communities. Comparisons indicated that certain areas in the north temperate region had higher helminth species richness in fishes than areas in the tropics. Main conclusions: Expectations of high species richness in helminth communities of tropical freshwater fishes are not fulfilled by the data. Direct comparisons of infracommunities and component communities in host species across widely separated phylogenetic and geographical lines are inappropriate. Examination of latitudinal differences in richness of monophyletic parasite groups or of compound communities may uncover patterns different from those found in this study. Richness of helminth communities may be ultimately determined not by the number of host species present but by the degree of divergence of host lineages and by their diversification modes. A phylogenetic framework for hosts and parasites will reveal if increased host species richness within host clades, when host speciation is accompanied by habitat or diet specialization, or both, leads to lower helminth diversity in host species by fragmentation of a core helminth fauna characteristic or specific of the larger host clade. This pattern may be analysed in the context of cospeciation and acquisition from other unrelated hosts (host-sharing or host-switching).

Journal of Biogeography↗

USGS2002

Native American impacts on fire regimes of the California coastal ranges

Aim Native American burning impacts on California shrubland dominated landscapes are evaluated relative to the natural lightning fire potential for affecting landscape patterns. Location Focus was on the coastal ranges of central and southern California. Methods Potential patterns of Indian burning were evaluated based upon historical documents, ethnographic accounts, archaeological records and consideration of contemporary land management tactics. Patterns of vegetation distribution in this region were evaluated relative to environmental factors and the resilience of the dominant shrub vegetation to different fire frequencies. Results Lightning fire frequency in this region is one of the lowest in North America and the density of pre-Columbian populations was one of the highest. Shrublands dominate the landscape throughout most of the region. These woody communities have weak resilience to high fire frequency and are readily displaced by annual grasses and forbs under high fire frequency. Intact shrublands provided limited resources for native Americans and thus there was ample motivation for using fire to degrade this vegetation to an open mosaic of shrubland/grassland, not unlike the agropastoral modification of ecologically related shrublands by Holocene peoples in the Mediterranean Basin. Alien-dominated grasslands currently cover approximately one-quarter of the landscape and less than 1% of these grasslands have a significant native grass presence. Ecological studies in the Californian coastal ranges have failed to uncover any clear soil or climate factors explaining grassland and shrubland distribution patterns. Main conclusions Coastal ranges of California were regions of high Indian density and low frequency of lightning fires. The natural vegetation dominants on this landscape are shrubland vegetation that often form dense impenetrable stands with limited resources for Native Americans. Natural fire frequencies are not high enough to maintain these landscapes in habitable mixtures of shrublands and grasslands but such landscape mosaics are readily produced with additional human subsidy of ignitions. It is hypothesized that a substantial fraction of the landscape was type converted from shrubland to grassland and much of the landscape that underwent such type conversion has either been maintained by Euro-American land management practices or resisted recolonization of native shrublands. It appears that these patterns are disturbance dependent and result from anthropogenic alteration of landscapes initiated by Native Americans and sustained and expanded upon by Euro-American settlers.

Journal of Biogeography↗

USGS2002

Availability of food resources, distribution of invasive species, and conservation of a Hawaiian bird along a gradient of elevation

Aim: We evaluated how an elevation gradient affects: (1) the availability of food required by a specialist seed-eater, Loxioides bailleui Oustalet (Drepanidinae), or pallia, and hence the distribution of this endangered Hawaiian bird, and (2) the distribution of alien threats to Loxioides populations, their primary foods, and their dry-forest habitat, and hence strategies for their conservation. Location: We worked throughout the subalpine forest that encircles Mauna Kea Volcano, Hawai'i Island, but we focused our studies mainly on the western slope between 2000 and 3000 m elevation, where the gradient of elevation was greatest and palila were most abundant. Methods: We determined phenology and productivity patterns of the endemic dry-forest tree species, Sophora chrysophylla (Salisb.) Seem. (Fabaceae), or ma??mane, which provides Loxioides with most of their food, and another common endemic tree, Myoporum sandwicense A. Gray (Myoporaceae), or naio, which provides some resources, along a 786-m elevation gradient at monthly intervals for 10 years (Sophora only). We also determined the availability each month of moth larvae (Lepidoptera) for that were important in the diet of nestling and adult palila. In addition, we documented the incidence of parasitism on moth larvae by several wasp (Hymenoptera) and fly (Diptera) species, and we determined the distribution of predatory wasps and ants (Hymenoptera), which potentially threaten insect prey of birds. Percentage cover of alien grass species that pose fire threats in palila habitat and other weeds were assessed during one survey. Small mammal abundance and distribution were determined by trapping during three (rodent) or five (carnivore) surveys. Results: Sophora flower and seed (pod) availability varied predictably along the elevation gradient, with about 4 months separating peaks in reproduction at high and low elevations. This, together with highly variable production of flowers and pods within elevation strata, resulted in Sophora resources being available to Loxioides throughout the year on the western slope of Mauna Kea. Sophora produced flowers and pods more seasonally where gradients of elevation were short; thus, resources were available less consistently. In contrast, Myoporum produced flowers and fruits with little variation with respect to season or elevation. The availability of important insect prey of Loxioides was also related to elevation, in part because threats to Lepidoptera larvae from parasitic wasps were generally less at higher elevations. Threats to insect prey from predatory ants was also less at higher elevations but the abundance of predatory wasps was not related to elevation. Several weeds that pose the most serious threats to Loxioides habitat were more abundant at mid and low elevations, and alien grass cover was somewhat greater at mid elevation, thereby increasing fire risks in the centre of Loxioides habitat. Predatory mammals, in particular Felis catus Linnaeus, were common throughout the subalpine forest of Mauna Kea. However, Rattus rattus Linnaeus was rare, especially at higher elevations, whereas Mus musculus Linnaeus was more abundant at lower elevations. Main conclusions: Loxioides are concentrated in habitat that is distributed along a substantial gradient of elevation at least in part because food is available throughout the year and threats to food resources are less concentrated. To recover Loxioides elsewhere in its former range, habitats must be restored and alien threats reduced along extensive elevation gradients. Conservation along environmental gradients will likely benefit other Hawaiian birds that track the availability of food across landscapes or that have been stranded in the higher portions of their original ranges because of the greater impacts of alien diseases, predators, food competitors, and habitat stressors at lower elevations.

Journal of Biogeography↗

USGS2003

Large-scale phytogeographical patterns in East Asia in relation to latitudinal and climatic gradients

Aim: This paper aims at determining how different floristic elements (e.g. cosmopolitan, tropical, and temperate) change with latitude and major climate factors, and how latitude affects the floristic relationships between East Asia and the other parts of the world. Location: East Asia from the Arctic to tropical regions, an area crossing over 50?? of latitudes and covering the eastern part of China, Korea, Japan and the eastern part of Russia. Methods: East Asia is divided into forty-five geographical regions. Based on the similarity of their world-wide distributional patterns, a total of 2808 indigenous genera of seed plants found in East Asia were grouped into fourteen geographical elements, belonging to three major categories (cosmopolitan, tropical and temperate). The 50??-long latitudinal gradient of East Asia was divided into five latitudinal zones, each of c. 10??. Phytogeographical relationships of East Asia to latitude and climatic variables were examined based on the forty-five regional floras. Results: Among all geographical and climatic variables considered, latitude showed the strongest relationship to phytogeographical composition. Tropical genera (with pantropical, amphi-Pacific tropical, palaeotropical, tropical Asia-tropical Australia, tropical Asia-tropical Africa and tropical Asia geographical elements combined) accounted for c. 80% of the total genera at latitude 20??N and for c. 0% at latitude 55-60??N. In contrast, temperate genera (including holarctic, eastern Asia-North America, temperate Eurasia, temperate Asia, Mediterranean, western Asia to central Asia, central Asia and eastern Asia geographical elements) accounted for 15.5% in the southernmost latitude and for 80% at 55-60??N, from where northward the percentage tended to level off. The proportion of cosmopolitan genera increased gradually with latitude from 5% at the southernmost latitude to 21% at 55-60??N, where it levelled off northward. In general, the genera present in a more northerly flora are a subset of the genera present in a more southerly flora. Main conclusions: The large-scale patterns of phytogeography in East Asia are strongly related to latitude, which covaries with several climatic variables such as temperature. Evolutionary processes such as the adaptation of plants to cold climates and current and past land connections are likely responsible for the observed latitudinal patterns.

Journal of Biogeography↗

USGS2003

Species area relationships in mediterranean-climate plant communities

Aim To determine the best-fit model of species–area relationships for Mediterranean-type plant communities and evaluate how community structure affects these species–area models. Location Data were collected from California shrublands and woodlands and compared with literature reports for other Mediterranean-climate regions. Methods The number of species was recorded from 1, 100 and 1000 m 2 nested plots. Best fit to the power model or exponential model was determined by comparing adjusted r 2 values from the least squares regression, pattern of residuals, homoscedasticity across scales, and semi-log slopes at 1–100 m 2 and 100–1000 m 2 . Dominance–diversity curves were tested for fit to the lognormal model, MacArthur's broken stick model, and the geometric and harmonic series. Results Early successional Western Australia and California shrublands represented the extremes and provide an interesting contrast as the exponential model was the best fit for the former, and the power model for the latter, despite similar total species richness. We hypothesize that structural differences in these communities account for the different species–area curves and are tied to patterns of dominance, equitability and life form distribution. Dominance–diversity relationships for Western Australian heathlands exhibited a close fit to MacArthur's broken stick model, indicating more equitable distribution of species. In contrast, Californian shrublands, both postfire and mature stands, were best fit by the geometric model indicating strong dominance and many minor subordinate species. These regions differ in life form distribution, with annuals being a major component of diversity in early successional Californian shrublands although they are largely lacking in mature stands. Both young and old Australian heathlands are dominated by perennials, and annuals are largely absent. Inherent in all of these ecosystems is cyclical disequilibrium caused by periodic fires. The potential for community reassembly is greater in Californian shrublands where only a quarter of the flora resprout, whereas three quarters resprout in Australian heathlands. Other Californian vegetation types sampled include coniferous forests, oak savannas and desert scrub, and demonstrate that different community structures may lead to a similar species–area relationship. Dominance–diversity relationships for coniferous forests closely follow a geometric model whereas associated oak savannas show a close fit to the lognormal model. However, for both communities, species–area curves fit a power model. The primary driver appears to be the presence of annuals. Desert scrub communities illustrate dramatic changes in both species diversity and dominance–diversity relationships in high and low rainfall years, because of the disappearance of annuals in drought years. Main conclusions Species–area curves for immature shrublands in California and the majority of Mediterranean plant communities fit a power function model. Exceptions that fit the exponential model are not because of sampling error or scaling effects, rather structural differences in these communities provide plausible explanations. The exponential species–area model may arise in more than one way. In the highly diverse Australian heathlands it results from a rapid increase in species richness at small scales. In mature California shrublands it results from very depauperate richness at the community scale. In both instances the exponential model is tied to a preponderance of perennials and paucity of annuals. For communities fit by a power model, coefficients z and log c exhibit a number of significant correlations with other diversity parameters, suggesting that they have some predictive value in ecological communities.

Journal of Biogeography↗

USGS2004

Twentieth century demographic changes in cirio and cardón in Baja California, Mexico

Aim&ensp; Our purpose was to discern long-term large-scale patterns of survivorship and recruitment of two dominant, charismatic and useful desert plants, cirio [Fouquieriaceae; Fouquieria columnaris (Kell.) Kell. ex Curran] and cardón [Cactaceae; Pachycereus pringlei (S.Watson) Britton & Rose], and to evaluate the effects of physical and human factors. Location&ensp; The study included 77 sites distributed over c. 13,200 km 2 in the northern Vizcaíno Region of the Sonoran Desert, state of Baja California, México. Method&ensp; Censuses used identified individuals ( n = 3780 cirio and 2246 cardón) and were based on repeat photography. Time intervals between censuses ranged from 29 to 96 years, ending in 1996–2002. Earlier repetition (1963) of old photographs at 14 sites allowed analysis of temporal variation. The effect of elapsed time was modelled with Weibull functions for survivorship and linear functions for recruitment. To explain the residual variations, we tested categorical and continuous variables related to substrate, geomorphology, climate and geography. Human impacts were weighed with historical and quantitative analysis. Results&ensp; Elapsed time accounted for 45% of the variation of survivorship in cirio but only 35% in cardón. The fitted Weibull curves were used to estimate longevity: all-size cohorts of 1000 individuals would die out in 388 (cirio) and 723 years (cardón). Recruitment in cirio was linearly related to time ( r 2 = 0.71) and averaged < 0.003 year −1 but was less than mortality at 90% of the sites. For cardón, recruitment averaged > 0.005 year −1 , was linearly related to time ( r 2 = 0.66) and was less than mortality at only 33% of the sites. Vital rates were not strongly correlated between mid- and late-century but were mostly similar to the long-term regional rates. Removing the factor of elapsed time, the residual variations showed that survivorship was greater on sites protected from winds for cirio and was less on very coarse soils for cardón. Recruitment increased with latitude in cirio and was greater on southern exposures for both species. The residual variations were not clearly affected, at a regional scale, by other factors thought to be important, including elevation, distance to the Pacific Ocean, geology, slope gradient, soil stability, older vs. young alluvial soils, and soil Ca : Mg and Na : K ratios. Human impacts have been sporadic and heterogeneous but locally strong; our quantitative indices of accessibility did not show regionally significant effects. Blowdown by hurricanes is a sporadic cause of major mortality for cirio but not necessarily for cardón. Main conclusions&ensp; At our scales, effects of time outweigh those of substrate, and human impact was scant. Large patterns were pronounced: cirio experienced a prolonged, widespread decline in the 1900s, while cardón fluctuated in different directions and degree among local populations. Cirio was found to be inherently much shorter-lived than cardón. We also suggest that recruitment in cirio was low due to long periods with relatively dry winters that affected the entire region, while spatial heterogeneity of cardón recruitment seemed more related to the variation in summer rains.

Journal of Biogeography↗