Across island and continents, mammals are more successful invaders than birds (Reply)

Aim  The establishment success of exotic species is calculated as the fraction of introduced species that have become established, and invasion success is estimated as the fraction of established species that have spread significantly from their points of introduction. Records on species introductions are highly incomplete, so strong conclusions about the tens rule and invasibility of island and continents cannot be drawn.
Location  Global.
Methods  Using Jeschke (2008) as an example, we explain the issue of the inadequacy of data to draw conclusions about the tens rule and invasibility of island and continents.
Results  Lack of adequate data.
Main Conclusion  Jeschke (2008) probably overestimates the establishment and invasion success rates, so his conclusions about violation of the tens rule and that islands are not more susceptible to invasion than continents are misleading.     

Densely sampled phylogenetic analyses of the Lesser Short-toed Lark (Alaudala rufescens) — Sand Lark (A. raytal) species complex (Aves,Passeriformes) reveal cryptic diversity

The taxonomy of the Lesser/Asian Short-toed Lark Alaudala rufescens–cheleensis complex has been debated for decades, mainly because of minor morphological differentiation among the taxa within the complex, and different interpretations of the geographical pattern of morphological characters among different authors. In addition, there have been few studies based on non-morphological traits. It has recently been suggested based on a molecular study of the lark family Alaudidae that the Sand Lark A. raytal is nested within this complex. We here analysed mitochondrial cytochrome b (cyt b) from 130 individuals across the range of this complex (hereafter called Alaudala rufescens–raytal complex), representing all except two of the 18 currently recognized subspecies. We also analysed 11 nuclear markers from a subsample of these individuals, representing all of the clades found in the cyt b tree. Five primary clades were recovered, which confirmed that A. raytal is nested within this complex. Divergence time estimates among these five clades ranged from 1.76 to 3.16 million years (my; 95% highest posterior density [HPD] 1.0–4.51 my) or 1.99–2.53 my (95% HPD 0.96–4.3 my) in different analyses. Only four of the currently recognized subspecies were recovered as monophyletic in the cyt b tree. Our results call for a taxonomic revision, and we tentatively suggest that at least four species should be recognized, although we stress the need for an approach integrating molecular, morphological and other data that are not yet available.     

Alien species as a driver of recent extinctions

We assessed the prevalence of alien species as a driver of recent extinctions in five major taxa (plants, amphibians, reptiles, birds and mammals), using data from the IUCN Red List. Our results show that alien species are the second most common threat associated with species that have gone completely extinct from these taxa since AD 1500. Aliens are the most common threat associated with extinctions in three of the five taxa analysed, and for vertebrate extinctions overall.     

A mitochondrial-DNA-based phylogeny for some evolutionary-genetic model species of Colias butterflies (Lepidoptera, Pieridae)

We study the phylogenetic relationships among some North American Colias ("sulfur") butterflies, using mitochondrial gene sequences (ribosomal RNA, cytochrome oxidase I+II) totaling about 20% of the mitochondrial genome. We find that (1) the lowland species complex shows a branching order different from earlier views; (2) several montane and northern taxa may be more distinct than in earlier views; (3) one morphologically conservative Holarctic assemblage, C. hecla, is differentiated at the molecular-genetic level into at least three taxa which occupy distinct positions in the phylogeny and are sisters to diverse other taxa. These conclusions, constituting phylogenetic hypotheses, are supported by parsimony, maximum-likelihood, and Bayesian reconstruction algorithms. They are tested formally, by interior branch tests and paired-site tests, against alternative hypotheses derived from conventional species and subspecies naming combinations. In all cases our hypotheses are supported by these tests and the conventional alternatives are rejected. The "barcoding" subset of cytochrome oxidase I sequence identifies only some of the taxa supported by our full data set. Comparison of genetic divergence values among Colias taxa with those among related Pierid butterflies suggests that species radiations within Colias are comparatively younger. This emerging Colias phylogeny facilitates comparisons of genetic polymorphism and other adaptive mechanisms among taxa, thereby connecting micro- and macro-evolutionary processes.     

Exotic species introductions into South America: an underestimated threat?

Prior studies on the latitudinal extent and ecological impact of exotic plant species suggest that areas of high diversity, such as the Neotropics, may be relatively 'resistant' to invasions. To explore the generality of this assertion and assess the impact of alien species on continental tropical faunas, I compiled data for threatened Neotropical animals from the red data books of Bolivia, Brazil, Minas Gerais (a Brazilian state), Peru, and Venezuela. A total of 378 species (including both vertebrates and invertebrates) were considered. For each taxon, I recorded whether it is threatened by habitat conversion, overexploitation, and/or exotic species. As suggested by other researchers, exotic species introductions appear to be relatively unimportant in South America, threatening only 6% of animal taxa. However, many South American animals are themselves either recent invaders or survivors of the Great American Biotic Interchange (GABI) which began during the Pliocene. Here, I hypothesize that the GABI may have acted as an 'extinction filter', leaving faunal groups of mostly South American origin relatively more threatened by the current wave of exotic invaders than those with prominent North American representation. The data support this prediction. For taxa whose current diversity patterns were not strongly influenced by the GABI, exotic species are indeed an important threat. For example, alien invaders threaten 29% of continental fishes and 30% of amphibians, figures comparable to those recorded in temperate areas. As more information on these less-studied taxa becomes available, the magnitude of the threat posed by exotic species introductions will probably reveal itself to be large. Of critical importance is to assess the impact of invasions on biological realms that have only been recently exposed to alien taxa, such as the aquatic faunas of the numerous drainages that occur along the eastern and western slopes of the South American Andes. The results of these investigations provide predictions for similar research focussed on other continental tropical regions of the world.     

Mammals and meaningful taxonomic units: the debate about species concepts and conservation

Mammalian taxonomy based on the diagnosability version of the Phylogenetic Species Concept has recently been declared the only sound approach to mammalian classification. In this article, based on the underlying evolutionary ontology of species taxa, I explain the fallacy of these arguments and hold that in a ‘grey area’ after lineage sundering, completely objective species delimitation is impossible, making both lumping and splitting equally correct or incorrect. As a consequence, we may have to get used to the idea that described species are often a poor guide to the delineation of meaningful taxonomic units for conservation.     

Avian foraging studies: an overlooked source of distribution data for macroecological and conservation studies

Macroecological and biogeographical studies and their applicability for biological conservation vitally depend on distribution data. These are usually taken from distribution atlases and databases or maps found in identification guides. A previous study pointed to another little explored source of data — local faunistic studies. Here, I would like to draw attention to another potential source. Papers that analysed the food composition of some taxa (e.g. insectivorous birds) are an overlooked source of rich information on taxa distributions. These studies frequently include an 'Appendix' with a list of food items determined to the species level. These studies also contain data on abundances, number of samples, sampling time, and geographical location as a rule. Foraging birds naturally provide data on invertebrate distributions with good spatio-temporal coverage and reasonably large samples. Importantly, birds frequently collect rare and by entomological methods hardly detectable species (i.e. those living high, in tree canopies, in very dense vegetation, or with secretive lifestyle). Data from bird dietary studies may help to ameliorate one of the most serious problems in distribution studies — zero inflation. I briefly discuss pros and cons of this so far neglected source of biogeographical data.     

Correlates of species richness in North American bat families

Aim A near universal truth in North America is that species richness increases from the Arctic Circle to the Central American tropics. Latitude is regarded as a major explanatory variable in species density, although it is only a surrogate for underlying ecological variables. I aimed to elucidate those underlying ecological variables that are associated with variation in bat species richness across the entire North American continent, providing a portrait of the macroecology of the order Chiroptera and its familial components. Methods I determined the number of bat species recorded for every state in Mexico and the United States, every province or territory in Canada, and every country in Central America. For each of these entities (n = 99), I also gathered basic data on mean annual precipitation, variation across the year (July vs. January) in mean temperature, mean January temperature, range in elevation (topographic relief), per cent vegetative cover and median latitude. Using a variety of linear regression and model‐fitting techniques, I analysed the strength and direction of the relationship between species richness and environmental variables for the order Chiroptera as a whole and separately for each of four familial groups: Molossidae (free‐tailed bats), Phyllostomidae (New World leaf‐nosed bats), Vespertilionidae (evening bats), and a set of six families (the Desmodontidae, Emballonuridae, Furipteridae, Natalidae, Noctilionidae, and Thyropteridae) represented in North America relatively poorly. Results and main conclusions Save for the Vespertilionidae, species richness of bats increased towards the Panamanian Isthmus. The Phyllostomidae and the set of miscellaneous families are particularly speciose in tropical Central America, with many fewer species occurring through subtropical Mexico into (in some cases) the southernmost United States. The Molossidae extends farther north, sparingly into the middle of the United States. Species density of the Vespertilionidae peaks in central and western Mexico and the southernmost United States, declining south through tropical southern Mexico and Central America and north through the central United States into Canada. Annual precipitation, January temperature, and topography are good predictors of species richness in the Chiroptera and the Molossidae, precipitation, topography, and temperature range in the Phyllostomidae, January temperature and topography in the Vespertilionidae, and precipitation alone in the collection of families. Vegetative cover explained little variation in the Chiroptera as a whole or in any family. After accounting for the effects of the environmental variables, latitude explained an insignificant amount of the residual variation in species richness. Bat families differ in their ecology, so studies of bat biogeography in North America may be misleading if they are examined only at the ordinal level.     

A morphometric study of three closely related taxa in the European Isoetes velata complex

Isoetes velata A. Braun is an amphibious plant with Mediterranean and Ibero-Atlantic distribution, showing marked morphological variability. Within the same geographical area, closely related restricted-range taxa are also present, including I. velata ssp. asturicense (restricted to mountainous areas of the north-west Iberian Peninsula) and I. boryana (restricted to the southern French Atlantic coast). These related taxa have unclear taxonomy and are both threatened. We performed a morphometric study of specimens of the three taxa. Our results suggest that I. velata ssp. asturicense is better considered as a distinct species, I. asturicense Laínz (Laínz), distinguishable from the other two taxa by mean megaspore length ( N  ≥ 30) < 360 µm. In contrast, our results do not support consideration of I. boryana as a separate species, and we here propose that this taxon be considered as a variety, I. velata A. Braun var. boryana M.I. Romero & C. Real stat. nov.   © 2005 The Linnean Society of London, Botanical Journal of the Linnean Society , 2005, 148 , 459–464.     

Associations of leaf miners and leaf gallers with island plants of different residency histories

Aim Introduced plant species are less likely to be attacked by herbivores than are native plant species. Isolated oceanic islands provide an excellent model system for comparing the associations between herbivore species and plant species of different residency histories, namely endemic, indigenous (non‐endemic) or introduced (naturalized or cultivated) species. My aim was to test the prediction that, on isolated oceanic islands, introduced plant species have a lower tendency to have an association with insect herbivores than do endemic and indigenous plant species. Location Ogasawara (Bonin) Islands in the western Pacific Ocean. Methods I examined the presence/absence of leaf‐mining and leaf‐galling insect species on 71 endemic, 31 indigenous, 18 naturalized and 31 cultivated (introduced but not naturalized) species of woody plants from 2004 to 2008. Results Leaf‐mining insect species were found on 53.5%, 35.5%, 11.1% and 16.1% and leaf‐galling species were found on 14.1%, 9.7%, 5.6% and 0% of endemic, indigenous, naturalized and cultivated plant species, respectively. Species of Lepidoptera (moths) and Hemiptera (primarily psyllids) comprised the dominant types of leaf miners and leaf gallers, respectively. Main conclusions The incidence of leaf miners and leaf gallers differed as a function of residency history of the plant species. Introduced (naturalized and cultivated) species were less frequently associated with leaf miners and leaf gallers than were native (endemic and indigenous) species, indicating that the leaf‐mining and leaf‐galling insect species, most of which feed on leaves of a particular native plant genus (i.e. they show oligophagy), have not yet begun to utilize most introduced plant species.     

Considerations regarding species distribution models for forest insects

1. Species distribution models, or SDMs, have become important decision support tools by answering fundamental questions about where species, including invasive species, are likely to survive and thrive based on environmental conditions.
2. For an inexperienced modeller or model reviewer, the terminology and technical aspects of SDMs can be overwhelming, and even well-trained modellers can struggle to understand the implications of various modelling choices.
3. Here, I outline some key considerations with respect to SDMs, focusing on their application to forest insects. Foremost, I assert that a model should be developed and evaluated with attention to relationships between an insect and its hosts, as those relationships determine much about the places the insect may occupy.
4. In my view, the most successful models are constructed carefully and incorporate honest assessments of their limitations, sources of error and uncertainty, and the degree of linkage between the model and the real-world circumstances it is meant to portray.

     

The frog filter: amphibian introduction bias driven by taxonomy,body size and biogeography

Aim Invasive species often exhibit a highly non‐random suite of traits relative to non‐invasive taxa, and these biases reflect strong selection at a series of steps along the invasion pathway. Here we investigate traits that are favoured in the first of these steps: the introduction of species outside their native geographic range. We use the global introduction of amphibians as our case study. Location Global. Methods We examined whether taxonomy, body size and attributes of the native geographic range predict the likelihood of an introduction event in amphibians. We then determined whether these characteristics differed between amphibian species that have been introduced intentionally and those that have been introduced unintentionally (i.e. stowaways). Results Comparisons of introduced and non‐introduced amphibians reveal significant biases with respect to taxonomic position (c. 50% of all introduced species originate from only 5% of all families), and characteristics of the native geographic range (most introduced species originate from the Northern Hemisphere, occupy a wide geographic range and are sympatric with high densities of humans). Many of these biases presumably reflect the ease and likelihood of capturing and transporting such animals. Additionally, intentionally introduced species are of larger than average body size, occupy higher elevations in their native ranges, and are more likely to originate from the Northern Hemisphere than unintentionally introduced species. Main conclusions Introduced amphibian species are not a random subset of the global amphibian fauna with respect to taxonomic affiliation and biogeography, and this restricts our ability to make robust generalizations regarding the ecological determinants of introduction success. Nevertheless, many of our findings are similar to those revealed by previous analyses of vertebrate and invertebrate introductions. Thus, our study suggests that biases in the anthropogenic mechanisms involved in transporting species around the globe are surprisingly consistent across broadly divergent taxa.     

Weak links: 'Rapoport's rule' and large-scale species richness patterns

Many hypotheses have been proposed to explain regional species richness patterns. Among these, ‘Rapoport's rule’ has sparked considerable controversy by stating that the latitudinal gradient in species richness can be explained indirectly as a function of narrower geographic ranges for species at low latitudes. Annual climatic variability, or deviation from mean climatic conditions, has been hypothesized to moderate this phenomenon. Furthermore, taxa that avoid much of this seasonality, such as temperate zone insects that enter diapause or species that migrate, were predicted to show reduced latitudinal gradients in richness. I test the suggested link between ‘Rapoport's rule’ and species richness for two higher level insect taxa as well as for the class Mammalia. Although these taxa exhibit the well-known latitudinal gradient in species richness, simple annual climatic variability and deviation from mean annual climatic conditions provide very poor predictions of species richness in each of them. Potential evapotranspiration, a measurement of ambient climatic energy, explains most of the observed variance in regional species richness patterns for all three taxa, consistent with the species richness-energy hypothesis. I find no support for an indirect link between ‘Rapoport's rule’ and terrestrial species richness patterns in North America.     

Biological traits predict shifts in geographical ranges of freshwater invertebrates during climatic warming and drying

Aim To test the ability of biological traits to predict climate‐related changes in geographical ranges of running‐water invertebrates. Location The Australian state of New South Wales and the Australian Capital Territory. Methods I analysed data from 8928 biomonitoring samples collected during a 16‐year period of generally rising air temperatures and declining precipitation. I used quantile regression to test for expansions and contractions on the climatically cooler, warmer, drier and wetter edges of the ranges of 120 invertebrate taxa, and correlated these shifts with the traits of thermophily (degree of preference for high versus low temperature) and rheophily (preference for flowing versus still water). Results The most commonly inferred range shifts were cool‐edge expansion plus warm‐edge contraction (71 taxa) and wet‐edge expansion plus dry‐edge contraction (71), but contractions from both cool and warm extremes (36) and from both dry and wet extremes (28) were also frequent. High‐temperature preference was associated with cool‐edge expansion and low‐temperature preference with wet‐edge expansion and contraction from all other extremes. A preference for flow was associated with wet‐edge expansion and dry‐edge contraction. Main conclusions Trait analysis has potential for predicting which species will expand their ranges and which will contract, but needs to be coupled with assessment of how the landscape provides each species with opportunities to track or avoid climate change. Improved quantification of climatically relevant traits and integration of trait analysis with species distribution modelling are likely to be beneficial.     

Parmelina quercina (Parmeliaceae, Lecanorales) includes four phylogenetically supported morphospecies

Morphological and phylogenetic relationships of the worldwide Mediterranean lichen forming fungus, Parmelina quercina , have been studied. Specimens from western Europe, western North America and southern Australia were analysed using molecular data (nuITS rDNA, nuLSU rDNA and mtSSU rDNA) and selected morphological features (upper cortex maculae, scanning electron microscopy examination of the epicortex, ascospores and conidia shape and size, and amphithecial retrorse rhizines). The results conclusively reveal that: (1) there is not one single species but four separate species in the Mediterranean or sub Mediterranean areas of the world. Parmelina quercina and Parmelina carporrhizans (Euroasiatic species), Parmelina coleae sp. nov. (North America) and Parmelina elixia sp. nov. (Australia); (2) largely debated P. carporrhizans is not a synonym of P. quercina but supported as a valid species circumscribed to Macaronesic relict sites; (3) the geographical isolation of the Australian population is correlated with a large genetic distance; (4) morphological characters (ascospores and conidial variability and thallus epicortex) correlate with the phylogenetic hypothesis; (5) the new or revalidated species within Parmelina quercina are not cryptic species but morphologically recognizable taxa.  © 2007 The Linnean Society of London, Biological Journal of the Linnean Society , 2007, 91 , 455–467.     

Electrophoretic delineation of species boundaries within the short-necked freshwater turtles of Australia (Testudines: Ghelidae)

A total of 222 specimens from 55 populations of short-necked chelid turtle was collected from drainages in Australia and Papua New Guinea. Two populations were initially considered to belong to different diagnosable taxa if all individuals in one population could be distinguished from all individuals in the other by fixed allozyme differences. When two populations or diagnosable taxa shared allozymes at all presumptive loci, their profiles were combined into a single diagnosable taxon. Comparisons between populations and emerging diagnosable taxa were repeated until no further changes were possible. The species Elseya dentata comprised five clearly diagnosable taxa, differing by between 4 and 19 fixed allozyme differences. The currently recognized El. latisternum and El. novaeguineae were each a single diagnosable taxon, and there were three diagnosable taxa, including a sibling pair, that could not be assigned to a currendy described Elseya species. In contrast, all forms of Emydura were very closely related, with no two taxa differing by more than three fixed allozyme differences. There were three diagnosable taxa in the north (Em. victoriae, Em. subglobosa and one new form), though support for them was marginal. In the south, Em. macquarii, Em. krefflii and Em. signata formed only a single diagnosable taxon, even sharing rare alleles. If the phylogenetic species concept is adopted, there is support for recognition of 16 species of short-necked turtle in Australia, including Pseudemydura umbrina. Currendy only 10 are described. Our data also provide evidence of reproductive isolation in some cases (sympatric or parapatric), and comparative evidence (sensu Mayr) in others, than the traditional biological species concept applies also to these diagnosable taxa.     

Species concepts and species reality: salvaging a Linnaean rank

The validity of the species category (rank) as a distinct level of biological organization has been questioned. Phenetic, cohesion and monophyletic species concepts do not delimit species-level taxa that are qualitatively distinct from lower or higher taxa: all organisms throughout the tree of life exhibit varying degrees of similarity, cohesion, and monophyly. In contrast, interbreeding concepts delimit species-level taxa characterized by a phenomenon (regular gene flow) not found in higher taxa, making the species category a distinct level of biological organization. Only interbreeding concepts delimit species-level taxa that are all comparable according to a biologically meaningful criterion and qualitatively distinct from entities assigned to other taxonomic categories. Consistent application of interbreeding concepts can result in counterintuitive taxonomies--e.g. many wide polytypic species in plants and narrow cryptic species in animals. However, far from being problematic, such differences are biologically illuminating--reflecting differing barriers to gene flow in different clades. Empirical problems with interbreeding concepts exist, but many of these also apply to other species concepts, whereas others are not as severe as some have argued. A monistic view of species using interbreeding concepts will encounter strong historical inertia, but can save the species category from redundancy with other categories, and thus justify continued recognition of the species category.