When is a "cryptic" species not a cryptic species: A consideration from the Holarctic micro-landsnail genus Euconulus (Gastropoda: Stylommatophora).

Naive use of molecular data may lead to ambiguous conclusions, especially within the context of "cryptic" species. Here, we integrated molecular and morphometric data to evaluate phylogenetic relationships in the widespread terrestrial micro-snail genus, Euconulus. We analyzed mitochondrial (16S + COII) and nuclear (ITS1 + ITS2) sequence across 94 populations from Europe, Asia and North America within the nominate species E. alderi, E. fulvus and E. polygyratus, and used the southeastern USA E. chersinus, E. dentatus, and E. trochulus as comparative outgroups. Phylogeny was reconstructed using four different reconstruction methods to identify robust, well-supported topological features. We then performed discriminant analysis on shell measurements between these genetically-identified species-level clades. These analyses provided evidence for a biologically valid North American "cryptic" species within E. alderi. However, while highly supported polyphyletic structure was also observed within E. fulvus, disagreement in placement of individuals between mtDNA and nDNA clades, lack of morphological differences, and presence of potential hybrids imply that these lineages do not rise to the threshold as biologically valid cryptic species, and rather appear to simply represent a complex of geographically structured populations within a single species. These results caution that entering into a cryptic species hypothesis should not be undertaken lightly, and should be optimally supported along multiple lines of evidence. Generally, post-hoc analyses of macro-scale features should be conducted to attempt identification of previously ignored diagnostic traits. If such traits cannot be found, i.e. in the case of potentially "fully cryptic" species, additional criteria should be met to propound a cryptic species hypothesis, including the agreement in tree topology among both mtDNA and nDNA, and little (or no) evidence of hybridization based on a critical analysis of sequence chromatograms. Even when the above conditions are satisfied, it only implies that the cryptic species hypothesis is plausible, but should optimally be subjected to further careful examination.

The macroecology of sustainability.

The discipline of sustainability science has emerged in response to concerns of natural and social scientists, policymakers, and lay people about whether the Earth can continue to support human population growth and economic prosperity. Yet, sustainability science has developed largely independently from and with little reference to key ecological principles that govern life on Earth. A macroecological perspective highlights three principles that should be integral to sustainability science: 1) physical conservation laws govern the flows of energy and materials between human systems and the environment, 2) smaller systems are connected by these flows to larger systems in which they are embedded, and 3) global constraints ultimately limit flows at smaller scales. Over the past few decades, decreasing per capita rates of consumption of petroleum, phosphate, agricultural land, fresh water, fish, and wood indicate that the growing human population has surpassed the capacity of the Earth to supply enough of these essential resources to sustain even the current population and level of socioeconomic development.

Macroecology Meets Macroeconomics: Resource Scarcity and Global Sustainability.

The current economic paradigm, which is based on increasing human population, economic development, and standard of living, is no longer compatible with the biophysical limits of the finite Earth. Failure to recover from the economic crash of 2008 is not due just to inadequate fiscal and monetary policies. The continuing global crisis is also due to scarcity of critical resources. Our macroecological studies highlight the role in the economy of energy and natural resources: oil, gas, water, arable land, metals, rare earths, fertilizers, fisheries, and wood. As the modern industrial technological-informational economy expanded in recent decades, it grew by consuming the Earth's natural resources at unsustainable rates. Correlations between per capita GDP and per capita consumption of energy and other resources across nations and over time demonstrate how economic growth and development depend on "nature's capital". Decades-long trends of decreasing per capita consumption of multiple important commodities indicate that overexploitation has created an unsustainable bubble of population and economy.

The age of island-like habitats impacts habitat specialist species richness.

While the effects of contemporaneous local environment on species richness have been repeatedly documented, much less is known about historical effects, especially over large temporal scales. Using fen sites in the Western Carpathian Mountains with known radiocarbon-dated ages spanning Late Glacial to modern times (16 975-270 cal years before 2008), we have compiled richness data from the same plots for three groups of taxa with contrasting dispersal modes: (1) vascular plants, which have macroscopic propagules possessing variable, but rather low, dispersal abilities; (2) bryophytes, which have microscopic propagules that are readily transported long distances by air; and (3) terrestrial and freshwater mollusks, which have macroscopic individuals with slow active migration rates, but which also often possess high passive dispersal abilities. Using path analysis we tested the relationships between species richness and habitat age, area, isolation, and altitude for these groups. When only matrix-derived taxa were considered, no significant positive relation was noted between species richness and habitat size or age. When only calcareous-fen specialists were considered, however, habitat age was found to significantly affect vascular plant richness and, marginally, also bryophyte richness, whereas mollusk richness was significantly affected by habitat area. These results suggest that in inland insular systems only habitat specialist (i.e., interpatch disperser and/or relict species) richness is influenced by habitat age and/or area, with habitat age becoming more important as species dispersal ability decreases.

Ecological niche divergence between extant and glacial land snail populations explained.

The presence of Last Glacial Maximum (LGM) biotic communities without modern counterparts is well known. It is particularly evident in central European fossil LGM land snails whose assemblages represent an odd mix of species that are currently limited to either xeric or wetland habitats. Here we document a genetically verified discovery of the modern calcareous wetland species Pupilla alpicola on Iceland, where it is limited to dry grasslands. This species also represents a common European LGM fossil, and its new records from Iceland help explain puzzling shifts of some glacial land snails of xeric grassland habitats to open wetlands today. Similarities between the climates of modern Iceland and LGM Eurasia suggest that this species did not become limited to wetlands in continental Europe until after the Late Pleistocene-Holocene climate transition. These results are a strong reminder that assumptions of ecological uniformity must be questioned and that the quality and robustness of palaeoecological reconstructions is dependent upon adequate knowledge of the full autecological range of species over time.

Evolutionary pattern and process within the Vertigo gouldii (Mollusca: Pulmonata, Pupillidae) group of minute North American land snails.

A phylogenetic analysis of 19 sibling taxa in the Vertigo gouldii group was conducted on 73 individuals sampled across North America using DNA sequence data of the mitochondrial genes cytochrome oxidase subunit 1 (CO1) and 16S ribosomal RNA (16S), and the internal transcribed spacer-2 of the nuclear ribosomal RNA (ITS-2) gene. The results of these analyses were found incongruent with previous taxonomic concepts used to define the V. gouldii group and its composite taxa that were based entirely on conchological features. The mtDNA sequence data suggest that some previous members of the traditional V. gouldii group may be more closely related to V. modesta. They also suggest that V. gouldii may itself consist of seven species-level branches spread across two deeply rooted clades. Revision of geographical distributions on the basis of these analyses suggests that these Vertigo species may commonly possess continental-sized ranges in spite of their minute size and limited active dispersal ability. High levels of sympatry within the group are also confirmed, with up to four species being known to co-occur within single microsites. These data also suggest that rates of diversification have been non-constant. Assuming a 1%/my rate of base pair substitution, a 10-fold diversification pulse is indicated from 6.7-7.0 myBP, which would be co-incident with known mid-late Miocene global climate changes.

The wealth of species: ecological communities, complex systems and the legacy of Frank Preston.

General statistical patterns in community ecology have attracted considerable recent debate. Difficulties in discriminating among mathematical models and the ecological mechanisms underlying them are likely related to a phenomenon first described by Frank Preston. He noted that the frequency distribution of abundances among species was uncannily similar to the Boltzmann distribution of kinetic energies among gas molecules and the Pareto distribution of incomes among wage earners. We provide additional examples to show that four different 'distributions of wealth' (species abundance distributions, species-area and species-time relations, and distance decay of compositional similarity) are not unique to ecology, but have analogues in other physical, geological, economic and cultural systems. Because these appear to be general statistical patterns characteristic of many complex dynamical systems they are likely not generated by uniquely ecological mechanistic processes.

Spatial constraint of peatland butterfly occurrences within a heterogeneous landscape.

The occurrence of ten butterfly taxa (Clossiana eunomia dawsonii, Clossiana freija, Clossiana frigga, Clossiana titania, Coenonympha inornata, Erebia discoidalis, Incisalia augustinus, Lycaena dorcas, Lycaena epixanthe, Oeneis jutta) was analyzed within three acid peatland habitat types from the Lake Superior drainage basin of northwestern Wisconsin. Both first- (nearest-neighbor spatial analysis) and second-order (Ripley's K) spatial point process statistics were used to identify the extents over which each distribution pattern significantly deviated from random expectations. Versions of these tests were used that identified significant spatial pattern uncorrelated to habitat location and habitat preference. These analyses documented non-random occurrence patterns in seven species. Deviations from random were largely confined to two extents: <50 km and 70-100+ km. The majority of non-random patterns at <50 km extents were examples of aggregation, while the majority of non-random patterns noted at the 70-100+ km scale were examples of segregation. These results demonstrate that even for winged animals inside a limited landscape, spatially constrained processes can be important determinants of distribution. It is likely that metapopulation dynamics and dispersal limitation help explain why aggregation is dominant at small scales. The mechanisms underlying the predominance of segregation at large scales are less clear, but may be related to migration history and/or weak environmental gradients.

The Malthusian-Darwinian dynamic and the trajectory of civilization.

Two interacting forces influence all populations: the Malthusian dynamic of exponential growth until resource limits are reached, and the Darwinian dynamic of innovation and adaptation to circumvent these limits through biological and/or cultural evolution. The specific manifestations of these forces in modern human society provide an important context for determining how humans can establish a sustainable relationship with the finite Earth.