Allozyme variation in the three extant populations of the narrowly endemic cycad Dioon angustifolium Miq. (Zamiaceae) from North-eastern Mexico

BACKGROUND AND AIMS: Dioon angustifolium was considered within D. edule. Recent morphometric and allozyme studies on D. edule have shown that D. angustifolium has originated from geographic isolation and is therefore considered to be a separate species. This cycad is endemic to north-eastern Mexico and is known only from three populations in the Sierra Madre Oriental mountain chain. Its populations are small when compared with its southern relative D. edule. In this study, genetic variation was determined within and between populations of D. angustifolium and the genetic consequences of habitat fragmentation and isolation of populations of this species were assessed. METHODS: Allozyme electrophoresis of 14 presumptive loci was used. The data were analysed with statistical approximations for estimating genetic diversity, structure, gene flow and recent genetic bottlenecks. KEY RESULTS: Means and standard deviations of genetic diversity estimators were: number of alleles per locus (A = 1.67 +/- 0.23), percentage of polymorphic loci (P = 52.4 +/- 23 %) and expected heterozygosity (H(E) = 0.218 +/- 0.093). The genetic variation attributable to differences among populations was 16.7 %. Mean gene flow between paired populations was Nm = 1.55 +/- 0.67, which is similar to that reported for endemic plant species of narrow geographical distribution and species with gravity-dispersed seed. A recent bottleneck is detected in the populations studied. CONCLUSIONS: Dioon angustifolium presents high levels of genetic diversity compared with other cycad species, in spite of small population sizes. The recent bottleneck effect did not effectively reduce the genetic variation to the extent of eliminating these populations. The distribution of D. angustifolium appears to be the result of historical biogeographical effects related to the Pleistocene glaciations. It is recommended that this species be catalogued in the IUCN Red List of Threatened Species and conservation efforts be made to preserve it.     

Morphological and geographic variation of the cycad Dioon edule Lindl. (Zamiaceae): ecological and evolutionary implications

The relationship in geographical distribution and morphological variation of leaflet width and length (diagnostic trait), between and within populations of Dioon edule Lindl., has been investigated throughout its known range in eastern Mexico (from the states of Nuevo León to Veracruz, north to south, respectively). A total of 1832 leaflets were measured for width and length from 154 plants distributed amongst five populations using four leaflet replicas from each of three leaves per plant. For leaflet width and length the variation among populations indicated significant stat-istical differences ( F _4,147 = 125.83; P  < 0.0001; R ² = 92.17% and F _4,147 = 9.04; P  < 0.001; R ² = 26.8%), respectively. With respect to leaflet width, the multiple range test showed three groups with a north to south distributional relationship along the range of the species. The correlation coefficient among paired populations, respect to geographical distance and the absolute value of the mean difference of leaflet width in each population, was positive, and different from zero ( r  = 0.82; P  = 0.013). A great variation of important ecological and evolutionary parameters was shown.  © 2003 The Linnean Society of London, Botanical Journal of the Linnean Society , 2003, 141 , 465–470.     

Population dynamics of the Mexican cycad Dioon edule Lindl. (Zamiaceae): life history stages and management impact

The demographic dynamics of three populations of Dioon edule Lindl. (Zamiaceae) were studied in a fragmented landscape using projection matrix modelling. Compared with other plant species, D. edule behaves like a tree life-form species. Density and spatial distribution patterns varied among populations according to models for animal-dispersed tree species. In all scenarios, λ was most sensitive to changes in abundance of adult plants. The elasticity reproductive component (F) for the three populations was zero and stasis values (L) were higher, this being a function of the permanence of non-reproductive individuals. It was detected that disturbance influences the population dynamics of D. edule as a function of adult plant persistence. This observation suggests that the conservation of adult plants is critical for D. edule and perhaps for all cycads species. Adult plant decapitation should be halted at the 'Monte Oscuro' population, subjected to sustainable management since 1990, if higher seed production is needed in rural nurseries.  © 2008 The Linnean Society of London, Botanical Journal of the Linnean Society , 2008, 157 , 381–391.     

Species diversity and structure of phytophagous beetle assemblages along a latitudinal gradient: predicting the potential impacts of climate change

Abstract.  1. Assemblages of phytophagous beetles on Acacia were examined along a 1150 km latitudinal gradient in eastern Australia to investigate the potential effects of climate change on insect communities. The latitudinal gradient was used as a surrogate for differences in temperature. Several possible confounding variables were held constant by selecting comparable sites and emphasising a single host-plant species.
2. Total species richness increased towards the tropics, but there were no significant differences among latitudes for average species density, species richness, Fisher's α , or average Chao-1 index.
3. Beetles sampled along the gradient were classified into four climate change response groups, depending on their latitudinal range and apparent host specificity: cosmopolitan species, generalist feeders , climate generalists , and specialists . These four groups might respond differently to shifting climate zones. Cosmopolitan species (22% of community, found at more than one latitude and on more than one host plant) may be resilient to climate change. Generalist feeders (16%, found only at one latitude but found on more than one Acacia species) may well feed on several species but will have to move with their climatic envelope. Climate generalists (6%, found only on Acacia falcata but found at more than one latitude) may be constrained by the host species' ability to either cope with the changing climate or move with it. Finally, specialists (55%, found only on A. falcata and at only one latitude) may be forced to move poleward concurrently with their host species, or go extinct.
4. The analyses indicate that community structure may be fairly resilient to temperature change. The displacement or local extinction of species, especially the species that are found at only one latitude and on only one host plant, however, may lead to significant changes in community composition.     

The phylogeny and biogeography of Hakea (Proteaceae) reveals the role of biome shifts in a continental plant radiation

The frequency of evolutionary biome shifts during diversification has important implications for our ability to explain geographic patterns of plant diversity. Recent studies present several examples of biome shifts, but whether frequencies of biome shifts closely reflect geographic proximity or environmental similarity of biomes remains poorly known. We explore this question by using phylogenomic methods to estimate the phylogeny of Hakea, a diverse Australian genus occupying a wide range of biomes. Model‐based estimation of ancestral regions indicates that Hakea began diversifying in the Mediterranean biome of southern Australia in the Middle Eocene–Early Oligocene, and dispersed repeatedly into other biomes across the continent. We infer around 47 shifts between biomes. Frequencies of shifts between pairs of biomes are usually similar to those expected from their geographic connectedness or climatic similarity, but in some cases are substantially higher or lower than expected, perhaps reflecting how readily key physiological traits can be modified to adapt lineages to new environments. The history of frequent biome‐shifting is reflected in the structure of present‐day assemblages, which tend to be more phylogenetically diverse than null‐model expectations. The case of Hakea demonstrates that the radiation of large plant clades across wide geographic areas need not be constrained by dispersal limitation or conserved adaptations to particular environments.     

Diversity and assemblage structure of phytophagous Hemiptera along a latitudinal gradient: predicting the potential impacts of climate change

Aims The aims were (1) to assess the species richness and structure of phytophagous Hemiptera communities along a latitudinal gradient, (2) to identify the importance of rare species in structuring these patterns, and (3) to hypothesize about how phytophagous Hemiptera communities may respond to future climate change. Location East coast of Australia. Methods Four latitudes within the 1150 km coastal distribution of Acacia falcata were selected. The insect assemblage on the host plant Acacia falcata was sampled seasonally over two years. Congeneric plant species were also sampled at the sites. Results Ninety‐eight species of phytophagous Hemiptera were collected from A. falcata. Total species richness was significantly lower at the most temperate latitude compared to the three more tropical latitudes. We classified species into four climate change response groups depending on their latitudinal range and apparent host specificity. Pairwise comparisons between groups showed that the cosmopolitan, generalist feeders and specialists had a similar community structure to each other, but the climate generalists had a significantly different structure. Fifty‐seven species were identified as rare. Most of these rare species were phloem hoppers and their removal from the dataset led to changes in the proportional representation of all guilds in two groups: the specialist and generalist feeders. Main conclusions We found no directional increase in phytophagous Hemiptera species richness. This indicates that, at least in the short term, species richness patterns of these communities may be similar to that found today. As the climate continues to change, however, we might expect some increases in species richness at the more temperate latitudes as species migrate in response to shifting climate zones. In the longer term, more substantial changes in community composition will be expected because the rare species, which comprise a large fraction of these communities, will be vulnerable to both direct climatic changes, and indirect effects via changes to their host's distribution.     

Genetic diversity of Saccharina latissima (Phaeophyceae) along a salinity gradient in the North Sea–Baltic Sea transition zone

The North Sea–Baltic Sea transition zone constitutes a boundary area for the kelp species Saccharina latissima due to a strong salinity gradient operating in the area. Furthermore, the existence of S. latissima there, along Danish waters, is fairly patchy as hard bottom is scarce. In this study, patterns of genetic diversity of S. latissima populations were evaluated along the salinity gradient area of Danish waters (here designated brackish) and were compared to reference sites (here designated marine) outside the gradient area, using microsatellite markers. The results showed that the S. latissima populations were structured into two clusters corresponding to brackish versus marine sites, and that gene flow was reduced both between clusters and between populations within clusters. In addition, results provided empirical evidence that marginal populations of S. latissima in the salinity gradient area exhibited a distinct genetic structure when compared to marine ones. Brackish populations were less diverse, more related, and showed increased differentiation over distance compared to marine populations. The isolation of the brackish S. latissima populations within the salinity gradient area of Danish waters in conjunction with their general low genetic diversity makes these populations vulnerable to ongoing environmental and climate change, predicted to result in declining salinity in the Baltic Sea area that may alter the future distribution and performance of S. latissima in the area.     

The importance (or lack thereof) of niche divergence to the maintenance of a northern species complex: the case of the long‐toed salamander (Ambystoma macrodactylum Baird)

The relative importance of ecological vs. nonecological factors for the origin and maintenance of species is an open question in evolutionary biology. Young lineages – such as the distinct genetic groups that make up the ranges of many northern species – represent an opportunity to study the importance of ecological divergence during the early stages of diversification. Yet, few studies have examined the extent of niche divergence between lineages in previously glaciated regions and the role of ecology in maintaining the contact zones between them. In this study, we used tests of niche overlap in combination with ecological niche models to explore the extent of niche divergence between lineages of the long‐toed salamander (Ambystoma macrodactylum Baird) species complex and to determine whether contact zones correspond to (divergent) niche limits. We found limited evidence for niche divergence between the different long‐toed salamander lineages, substantial overlap in the predicted distribution of suitable climatic space for all lineages and range limits that are independent of niche limits. These results raise questions as to the importance of ecological divergence to the development of this widespread species complex and highlight the potential for non‐ecological factors to play a more important role in the maintenance of northern taxa.     

Osmoregulatory capacity and the ability to use marine food sources in two coastal songbirds (Cinclodes: Furnariidae) along a latitudinal gradient

Cinclodes nigrofumosus and C. oustaleti are two closely related songbirds that inhabit the northern Chilean coast during the austral fall and winter.This stretch spans a dramatic north to south latitudinal gradient in rainfall and temperature. Whereas C. nigrofumosus lives exclusively on coastal environments, C. oustaleti shifts seasonally from coastal environments to inland freshwater ones. We used the δ¹³C of these two species’ tissues to investigate whether the reliance on marine versus terrestrial sources varied from the hyper-arid north to the wet south. We also investigated latitudinal variation in the renal traits that mediate how these birds cope with dehydration and a salty marine diet. Both species increased the incorporation of terrestrial carbon, as measured by δ¹³C, as terrestrial productivity increased southwards. However, C. nigrofumosus had consistently more positive (i.e. more marine) and less variable δ¹³C values than C. oustaleti. The osmoregulatory traits of both species varied with latitude as well. Urine osmolality decreased from extremely high values in the north to moderate values in the south, while C. nigrofumosus produced more concentrated urine than C. oustaleti. In both species, the proportion of kidney devoted to medullary tissue decreased from north to south, and kidney size increased significantly with latitude. Cinclodes nigrofumosus had larger kidneys with larger proportions of medullary tissue than C. oustaleti. C. nigrofumosus and C. oustaleti are terrestrial organisms subsidized by a rich marine environment where it is adjacent to an unproductive terrestrial. Variation in the reliance on marine food sources seems to be accompanied by adjustments in the osmoregulatory mechanisms used by these birds to cope with salt and dehydration.     

Testing the role of interspecific competition in the evolutionary origin of elevational zonation: an example with Buarremon brush-finches (Aves, Emberizidae) in the neotropical mountains

Interspecific competition might drive the evolution of ecological niches and result in pairs of formerly competing species segregating along ecological gradients following a process of character displacement. This mechanism has been proposed to account for replacement of related species along gradients of elevation in many areas of the world, but the fundamental issue of whether competition is responsible for the origin of elevational replacements has not been tested. To test hypotheses about the role of interspecific competition in the origin of complementary elevational ranges, I combined molecular phylogenetics, phylogeography, and population genetic analyses on Buarremon torquatus and B. brunneinucha (Aves, Emberizidae), whose patterns of elevational distribution suggest character displacement or ecological release. The hypothesis that elevational distributions in these species changed in opposite directions as a result of competition is untenable because: (1) a historical expansion of the range of B. brunneinucha into areas occupied by B. torquatus was not accompanied by a shift in the elevational range of the former species; (2) when B. brunneinucha colonized the range of B. torquatus, lineages of the latter distributions had already diverged; and (3) historical trends in effective population size do not suggest populations with elevational ranges abutting those of putative competitors have declined as would be expected if competition caused range contractions. However, owing to uncertainty in coalescent estimates of historical population sizes, the hypothesis that some populations of B. torquatus have declined cannot be confidently rejected, which suggests asymmetric character displacement might have occurred. I suggest that the main role of competition in elevational zonation may be to act as a sorting mechanism that allows the coexistence along mountain slopes only of ecologically similar species that differ in elevational distributions prior to attaining sympatry. The contrasting biogeographic histories of B. brunneinucha and B. torquatus illustrate how present-day ecological interactions can have recent origins, and highlights important challenges for testing the hypothesis of character displacement in the absence of data on population history and robust reconstructions of the evolution of traits and geographic ranges.     

Phylogenetic,Spatial, and Species‐Trait Patterns across Environmental Gradients: the Case of Hydropsyche (Trichoptera) along the Loire River

When Illies published his concept on the longitudinal zonation of lotic invertebrates five decades ago (Illies, J., 1961: Internat. Rev. ges. Hydrobiol. 46 : 205–213), he defined a research topic that currently interests many ecologists because he linked speciation and phylogeny with spatial distribution and trait adaptation to environmental conditions prevailing along rivers. We tested these ideas analyzing nine species of the caddisfly genus Hydropsyche from the Loire River (France). A morphology‐based phylogeny illustrated that the oldest of our species occurred in the headwater and that specific phylogenetic distances from the root location in the tree were significantly related to the specific longitudinal occurrences in the Loire. Furthermore, traits such as oxygen consumption, optimal velocity for filter‐net‐building, and larval size were significantly related to the specific phylogenetic distances, indicating meaningful, gradual adaptations to environmental gradients prevailing along European rivers (particularly in water temperature, near‐bottom flow, sediment porosity). Thus, joining phylogeny, spatial distribution, and species traits provided insights into a central topic of contemporary ecology, the spatial patterns of speciation, taxonomic community structure (species distributions), and niche adaptation (traits). (© 2011 WILEY‐VCH Verlag GmbH & Co. KGaA, Weinheim)     

Euglenoid morphospecies replacement along a hydraulic gradient of the Lower Paraná Basin (Argentina)

1. This study deals with the distribution of euglenoid morphospecies along the length of the Lower Paraná River floodplain. Six sites located in the Paraná River, Estevez stream and Montiel shallow lake were surveyed monthly from May 1995 to April 1996. 2. Eighty‐eight infrageneric taxa were registered though the total density percentage of euglenoids seldom exceeded 15% of total phytoplankton. The increasing species richness towards the lake is mostly the result of a high contribution of species of Trachelomonas and naked infrageneric taxa of Euglena, Lepocinclis and Phacus, while the distribution of Strombomonas species along the transect is fairly constant. 3. The richness of euglenoid morphospecies follows a pattern opposite to that found for the entire phytoplankton community. The distribution of euglenoids differs among the three habitats defined in this study: river, stream, shallow lake. These differences depend on the hydraulic condition of each habitat, as revealed by canonical correspondence analysis. 4. The degree of association with each type of aquatic habitat varied among the species and differed between water phases. Strombomonas fluviatilis was equally constant along the length of the ecotone, while S. girardiana and S. scabra were particularly associated with the Paraná River and P. longicauda, L. truncata, T. hispida and T. similis with the lake. 5. The flood pulse establishes a clear pattern both in species richness and abundance of euglenoids. The number of species recorded during high water was more than twice that recorded during low water.     

Phylogeography of the genus Dasiphora (Rosaceae) in the Qinghai‐Tibetan Plateau: divergence blurred by expansion

Plateau uprisings and climatic oscillations are considered to have caused extensive allopatric divergences that account for the rich species diversity of the Qinghai‐Tibetan Plateau (QTP). However, secondary contact during range shifts in the Quaternary glacial cycles or inter‐uplift stages may have restored the gene flow between species and so counteracted these divergences, particularly in rapidly‐adapting dominant elements. We tested this hypothesis by determining the phylogeographical history of Dasiphora (Rosaceae), a genus of two species that are widely distributed on the QTP and co‐exist in numerous localities. We sequenced two chloroplast DNA fragments (rbcL, trnT‐L) for 559 individuals from 87 populations. Bayesian methods were used to identify phylogenetic relationships and to estimate divergence times. Demographic histories were inferred using neutrality tests, mismatch distribution analysis, and coalescent simulation. A total of 112 haplotypes that clustered into three major groups were identified. The formation of these groups and their subgroups was dated to between the Pliocene and the late Pleistocene. In addition, we found that some groups underwent multiple extensive expansions. Species‐specific haplotypes were identified for each species, although these haplotypes phylogenetically intermixed. These results suggest that recent plateau uplifts and climatic oscillations might have caused the deep divergences observed within this genus. However, later range expansions probably blurred these divergences and possible species boundaries. Our results shed new light on the complex evolutionary history of the QTP alpine plants. © 2014 The Linnean Society of London, Biological Journal of the Linnean Society, 2014, 111 , 777–788.     

Reproductive characteristics of the flower breeding Drosophila hibisci Bock (Drosophilidae) along a latitudinal gradient in eastern Australia: relation to flower and habitat features

For many insect species, egg and larval substrate characteristics are significantly correlated with interspecific differences in female reproductive allocation and egg size-number tradeoffs. We tested the hypothesis that a similar pattern occurred within the Australian drosophilid, Drosophila hibisci, that is restricted throughout its life cycle to flowers of species in the genus Hibiscus. These plants occur as small, isolated, normally monospecific stands that should facilitate differentiation of the fly populations in relation to specific oviposition and larval substrates. Data from 38 sites ranging from 20.8̀ to 34.4̀ S latitude in eastern Australia indicated no relationship between female body size, egg size, or ovariole numbers and floral size or mass among four species of Hibiscus. However, the flies did show a latitudinal cline in ovariole number that was independent of floral variation. Females averaged 15–20 ovarioles per female in the south (32–34̀ S latitude) and 10–12 ovarioles in the north (21–22̀ S latitude). The increase in ovariole number with latitude was due to divergence in the ovariole number of the largest females. In contrast, small females in the north and south had the same number of ovarioles. Reproductive allocation of female flies in the northern region was less than females in the southern region. The latitudinal divergence in ovariole number was not associated with habitat differences (density of trees, density of flies and beetles), nor with differences in floral characteristics (flower weight, petal length, yeast species present). Short term weather patterns in daily temperature and rainfall preceding collections pardy explain the variation in ovariole number. These observations in conjunction with preliminary genetic results suggest the cline is associated with genetic differences that interact with environmental determinants such as the temperature during larval development.     

The latitudinal gradient of species diversity among North American grasshoppers (Acrididae) within a single habitat: a test of the spatial heterogeneity hypothesis

Abstract. The spatial heterogeneity hypothesis predicts a positive relationship between habitat complexity and species diversity: the greater the heterogeneity of a habitat, the greater the number of species in that habitat. On a regional scale, this hypothesis has been proposed to explain the increases in species diversity from the poles to the tropics: the tropics are more diverse because they contain more habitats. On the local scale, the spatial heterogeneity hypothesis suggests that the tropics are more diverse because they contain more microhabitats. The positive relationship between habitat heterogeneity and species diversity, on the local scale, is well documented. In this paper, we test whether habitat heterogeneity on the local scale can explain the latitudinal gradient of species diversity on the regional scale. We determined the latitudinal gradient of species diversity of 305 species of North American grasshoppers using published distribution maps. We compared the slope of this multihabitat (regional-scale) gradient with the slope of a within-habitat (local-scale) gradient in the prairie grasslands. Our results show no significant difference between the slopes at the two scales. We tested the generality of our results by comparing multi- and within-habitat latitudinal gradients of species diversity for ants, scorpions and mammals using data from the literature. These results are in accordance with those from grasshoppers. We can therefore reject the local-scale spatial heterogeneity hypothesis as a mechanism explaining the regional-scale latitudinal gradient of species diversity. We discuss alternative mechanisms that produce this gradient.     

Diatom assemblages along a former intertidal gradient: A palaeoecological study of a subboreal clay layer (Western coastal plain,Belgium)

Lateral and vertical variations in diatom taphocoenoses were studied in a clayey intercalation in the upper peat bed at Booitshoeke (western coastal plain, Belgium). The palaeoecological evolution, linked to sedimentary facies changes along the ecotone from fen to tidal flat, apparently involved two successive periods of increased local tidal activity. This result could be inferred from a weakly developed salt-marsh horizon within the clay, traceable by diatom analysis over a distance of a few hundred meters. Although influenced by taphonomic processes, the composition of the fossil assemblages appeared to reflect the relative position of the palaeoenvironment along the tidal gradient fairly accurately. Taphonomic groups of taxa could be distinguished, characterizing respectively the transition from fen to salt-marsh, the high salt-marsh, the low salt-marsh and high mud-flat, and the more polyhaline mud-flat. Sediments of drier salt-marsh conditions showed a very poor taphocoenosis, of mainly allochthonous origin, dominated by marine tychoplanktonic species with durable frustules. Some possible causes for the slight increase in marine activity during the mid-Subboreal, which resulted in the formation of the deposits considered here, are discussed.     

Evaluating higher taxa as surrogates of harvestmen biodiversity (Arachnida: Opiliones) along a latitudinal gradient in the Atlantic Forest

Habitat loss and fragmentation have highlighted the importance of monitoring remaining habitats. For megadiverse groups such as arthropods, of which many species are still being discovered, the use of higher taxonomic levels as substitutes for diversity may be a useful tool. The aim of this study was to evaluate the use of substitute taxonomic resolutions to assess the richness and composition of Laniatores harvestmen (Arachnida: Opiliones). The five resolutions selected were as follows: genus, family, subfamily, indicator taxa and intermediate resolution (combination of genus and species identification levels). In addition, we evaluated whether the diversity substitutes provide good estimates of latitudinal gradients. Nineteen Atlantic Forest sites located along a latitudinal gradient in northeastern Brazil were sampled. We recorded a total of 88 harvestmen species/morpho-species, distributed in 7 families, 15 subfamilies and 36 genera. Genus and intermediate resolution were excellent substitutes for harvestmen species richness. The efficiency differed according to the substitute resolution used. Four resolutions were adequate to replace the harvestmen composition: genus, intermediate resolution, indicator taxa, and subfamily. The number of harvestmen species recorded was significantly different between Seasonal Semideciduous Forest and Costal Atlantic Rainforest. The same relationship was also observed the same relationship was observed when we consider genus and intermediate resolution. Our results suggest the use of genus as a substitute for richness and composition of harvestmen for reducing monitoring costs and providing evaluation in a shorter time and a more practical way.