Putting biogeography’s cart back behind taxonomy’s horse: a response to Triantis et al.

In a recent paper, two of us discussed diversity patterns and diversification processes in the Azores flora. Triantis et al. (2012, Journal of Biogeography, 39, 1179–1184) challenged our hypothesis that palaeoclimatic differences had an effect on diversification rates and suggested that area, island age and isolation explain diversity patterns. They did not, however, fully address the results from our subsequent paper, in which we showed that diversity patterns evident from phylogeographic studies differ markedly from those suggested by checklists. Checklists are working hypotheses and we suggest that the discrepancies evident between molecular data and checklists may be indicative of deficiencies in our taxonomic understanding of the Azores flora. Patterns of molecular and morphological diversity need to be better understood, and the discrepancies between checklists and molecular data accounted for, before we can establish the relative importance of factors such as palaeoclimate, area, island age or isolation in generating endemic diversity patterns in the Azores flora.     

Potential natural vegetation and pre‐anthropic pollen records on the Azores Islands in a Macaronesian context

This paper discusses the concept of potential natural vegetation (PNV) in the light of the pollen records available to date for the Macaronesian biogeographical region, with emphasis on the Azores Islands. The classical debate on the convenience or not of the PNV concept has been recently revived in the Canary Islands, where pollen records of pre‐anthropic vegetation seemed to strongly disagree with the existing PNV reconstructions. Contrastingly, more recent PNV model outputs from the Azores Islands show outstanding parallelisms with pre‐anthropic pollen records, at least in qualitative terms. We suggest the development of more detailed quantitative studies to compare these methodologies as an opportunity for improving the performance of both. PNV modelling may benefit by incorporating empirical data on past vegetation useful for calibration and validation purposes, whereas palynology may improve past reconstructions by minimizing interpretative biases linked to differential pollen production, dispersal and preservation.     

Parasitoids of whiteflies (Hymenoptera: Aphelinidae,Eulophidae, Platygastridae; Hemiptera: Aleyrodidae) from the Macaronesian archipelagos of the Canary Islands,Madeira and the Azores

Data on the whitefly parasitoid species known from the Macaronesian archipelagos of the Canary Islands, Madeira and the Azores are presented, based largely on recently collected material. A total of 26 species are treated, including six new species, six new records for the Canary Islands, two new records for Madeira, and two new records for the Azores. All species are fully described and illustrated. New species described are: Encarsia atlantica Polaszek & Hernández; Encarsia levadicola Polaszek & Hernández; Encarsia melanostoma Polaszek & Hernández; Encarsia noahi Polaszek & Hernández; Euderomphale gomer LaSalle & Hernández; Euderomphale insularis LaSalle & Hernández. A fully illustrated identification key based on females is provided for recognition of whitefly parasitoids in these archipelagos. Data on the known distribution and hosts are provided, as well as references to biology and use in biological control.     

Of mice and the ‘Age of Discovery’: the complex history of colonization of the Azorean archipelago by the house mouse (Mus musculus) as revealed by mitochondrial DNA variation

Humans have introduced many species onto remote oceanic islands. The house mouse (Mus musculus) is a human commensal and has consequently been transported to oceanic islands around the globe as an accidental stowaway. The history of these introductions can tell us not only about the mice themselves but also about the people that transported them. Following a phylogeographic approach, we used mitochondrial D‐loop sequence variation (within an 849‐ to 864‐bp fragment) to study house mouse colonization of the Azores. A total of 239 sequences were obtained from all nine islands, and interpretation was helped by previously published Iberian sequences and 66 newly generated Spanish sequences. A Bayesian analysis revealed presence in the Azores of most of the D‐loop clades previously described in the domesticus subspecies of the house mouse, suggesting a complex colonization history of the archipelago as a whole from multiple geographical origins, but much less heterogeneity (often single colonization?) within islands. The expected historical link with mainland Portugal was reflected in the pattern of D‐loop variation of some of the islands but not all. A more unexpected association with a distant North European source area was also detected in three islands, possibly reflecting human contact with the Azores prior to the 15th century discovery by Portuguese mariners. Widening the scope to colonization of the Macaronesian islands as a whole, human linkages between the Azores, Madeira, the Canaries, Portugal and Spain were revealed through the sharing of mouse sequences between these areas. From these and other data, we suggest mouse studies may help resolve historical uncertainties relating to the ‘Age of Discovery’.     

Molecular phylogeny in endemic weevils: revision of the genera of Macaronesian Cryptorhynchinae (Coleoptera: Curculionidae)

A molecular phylogeny and lineage age estimates are presented for the Macaronesian representatives of the weevil subfamily Cryptorhynchinae, using two mitochondrial genes (cytochrome c oxidase subunit 1 and 16S). The Bayesian reconstruction is supplemented by observations on morphology, ecology, and reproductive biology. The present study often corroborates the groups previously outlined in higher‐level informal taxonomies. These and further groups are now assigned new taxonomic status. The following genera and subgenera are described (formerly Acalles): Aeoniacalles gen. nov. , Canariacalles gen. nov. , Ficusacalles gen. nov. , Madeiracalles gen. nov. , Silvacalles gen. nov. (with Tolpiacalles subgen. nov. , Tagasastacalles subgen. nov. ), Sonchiacalles gen. nov. , Echiumacalles gen. nov. (monotypic), Lauriacalles gen. nov. (monotypic), and Pseudodichromacalles gen. nov. (monotypic; formerly Dichromacalles). For the western Palaearctic genus Acalles Schoenherr, 1825 the first subgenus Origoacalles subgen. nov. is described and for the genus Onyxacalles Stüben, 1999 the first subgenus Araneacalles subgen. nov. ; Paratorneuma Roudier 1956 resyn. Except for one species of Acalles (Origoacalles), all of these new higher taxa are endemic to the Macaronesian Islands. All new taxa are presented, together with their host plants and further data, in a synoptic tabular overview. Based on the results of our phylogenetic analysis, we advocate the hypothesis that the evolution of the species in the new genera (of which most group into a ‘Macaronesian clade’) began in the comparatively arid succulent bush zone and that the shady and humid laurel forest of the thermo‐Canarian and thermo‐Madeiran zone was entered much later. Our reconstruction implies that the Canarian and Madeiran archipelagos were colonized by Cryptorhynchinae at least seven times from the continent but saw only one considerable adaptive radiation. It also becomes apparent that it is the ancestor species of the genus Canariacalles– and not Pseudodichromacalles– that features a close connection to the south‐western European and north‐western African species of Dichromacalles s.s. Finally, a key is presented for all genera and subgenera of the Macaronesian Cryptorhynchinae. © 2010 The Linnean Society of London, Zoological Journal of the Linnean Society, 2010, 160 , 40–87.     

Faunal response to sea‐level and climate change in a short‐lived seaway: Jurassic of the Western Interior,USA

Understanding how regional ecosystems respond to sea‐level and environmental perturbations is a main challenge in palaeoecology. Here we use quantitative abundance estimates, integrated within a sequence stratigraphic and environmental framework, to reconstruct benthic community changes through the 13 myr history of the Jurassic Sundance Seaway in the western United States. Sundance Seaway communities are notable for their low richness and high dominance relative to most areas globally in the Jurassic, and this probably reflects steep temperature and salinity gradients along the 2000 km length of the Seaway that hindered colonization of species from the open ocean. Ordination of samples shows a main turnover event at the Middle–Upper Jurassic transition, which coincided with a shift from carbonate to siliciclastic depositional systems in the Seaway, probably initiated by northward drift from subtropical latitudes to more humid temperate latitudes, and possibly global cooling. Turnover was not uniform across the onshore–offshore gradient, but was higher in offshore environments. The higher resilience of onshore communities to third‐order sea‐level fluctuations and to the change from a carbonate to a siliciclastic system was driven by a few abundant eurytopic species that persisted from the opening to the closing of the Seaway. Lower stability in offshore facies was instead controlled by the presence of more volatile stenotopic species. Such increased onshore stability in community composition contrasts with the well‐documented onshore increase in taxonomic turnover rates, and this study underscores how ecological analyses of relative abundance may contrast with taxonomically based analyses. We also demonstrate the importance of a stratigraphic palaeobiological approach to reconstructing the links between environmental and faunal gradients, and how their evolution through time produces local stratigraphic changes in community composition.     

The Pillars of Hercules as a bathymetric barrier to gene flow promoting isolation in a global deep‐sea shark (Centroscymnus coelolepis)

Knowledge of the mechanisms limiting connectivity and gene flow in deep‐sea ecosystems is scarce, especially for deep‐sea sharks. The Portuguese dogfish (Centroscymnus coelolepis) is a globally distributed and near threatened deep‐sea shark. C. coelolepis population structure was studied using 11 nuclear microsatellite markers and a 497‐bp fragment from the mtDNA control region. High levels of genetic homogeneity across the Atlantic (Φ_ST = ?0.0091, F_ST = 0.0024, P > 0.05) were found suggesting one large population unit at this basin. The low levels of genetic divergence between Atlantic and Australia (Φ_ST = 0.0744, P < 0.01; F_ST = 0.0015, P > 0.05) further suggested that this species may be able to maintain some degree of genetic connectivity even across ocean basins. In contrast, sharks from the Mediterranean Sea exhibited marked genetic differentiation from all other localities studied (Φ_ST = 0.3808, F_ST = 0.1149, P < 0.001). This finding suggests that the shallow depth of the Strait of Gibraltar acts as a barrier to dispersal and that isolation and genetic drift may have had an important role shaping the Mediterranean shark population over time. Analyses of life history traits allowed the direct comparison among regions providing a complete characterization of this shark's populations. Sharks from the Mediterranean had markedly smaller adult body size and size at maturity compared to Atlantic and Pacific individuals. Together, these results suggest the existence of an isolated and unique population of C. coelolepis inhabiting the Mediterranean that most likely became separated from the Atlantic in the late Pleistocene.     

Testing,as opposed to supporting,the Mid‐domain Hypothesis: a response to Lees and Colwell (2007)

Both our analysis ( Kerr et al. 2006 ), and Lees and Colwell's (2007) reanalysis, of patterns of bird and mammal diversity on Madagascar show that the central peak of richness predicted by the Mid‐Domain Hypothesis (MDH) is not observed. Lees and Colwell emphasize an observation consistent with MDH predictions: a latitudinal mid‐domain richness peak in the rainforest biome. They find (but do not mention) that no analogous peak is observed in the other two main Madagascan biomes. MDH fails nearly all its tests in Madagascar.     

Mitochondrial phylogeography and subspecies of the wide‐ranging sub‐Saharan leopard tortoise Stigmochelys pardalis (Testudines: Testudinidae) – a case study for the pitfalls of pseudogenes and GenBank sequences

The leopard tortoise (Stigmochelys pardalis) is the most widely distributed sub‐Saharan tortoise species, with a range extending from the Horn of Africa all over eastern Africa to the Republic of South Africa, Namibia and southernmost Angola. Using 1938 bp of mitochondrial DNA (cyt b gene, partial ND4 gene plus adjacent tRNA genes) from a nearly range‐wide sampling, we examined its phylogeographic structure and compared our findings with previously published GenBank sequences. We identified seven major clades that are largely parapatrically distributed. A few records of distinct haplotypes at the same locality or in close proximity could be the result of translocation of tortoises by man. The greatest diversity occurs in the south of the species’ range, with five out of the seven clades. Testing for isolation‐by‐distance suggests that the observed phylogeographic structure is the result of restricted geographical gene flow and not of historical vicariance. This is in sharp contrast to wide‐ranging thermophilic reptiles from the western Palaearctic, whose phylogeographic structure was significantly shaped by Pleistocene range interruptions, but also by earlier dispersal and vicariant events. Most cyt b sequences of S. pardalis from GenBank turned out to be nuclear pseudogenes, or to be of chimerical origin from such pseudogenes and authentic mitochondrial sequences, which argues for caution regarding uncritical usage of GenBank sequences. The recent revalidation of the two subspecies of S. pardalis was based on such a chimerical sequence that was erroneously identified with the subspecies S. p. babcocki. Furthermore, according to our data, the distribution of mitochondrial clades does match neither the traditional subspecies ranges nor the pronounced geographical size variation of leopard tortoises. We conclude that there is no rationale for recognizing subspecies within S. pardalis.     

Spatial synchrony in the response of a long range migratory species (Salmo salar) to climate change in the North Atlantic Ocean

A major challenge in understanding the response of populations to climate change is to separate the effects of local drivers acting independently on specific populations, from the effects of global drivers that impact multiple populations simultaneously and thereby synchronize their dynamics. We investigated the environmental drivers and the demographic mechanisms of the widespread decline in marine survival rates of Atlantic salmon (Salmo salar) over the last four decades. We developed a hierarchical Bayesian life cycle model to quantify the spatial synchrony in the marine survival of 13 large groups of populations (called stock units, SU) from two continental stock groups (CSG) in North America (NA) and Southern Europe (SE) over the period 1971–2014. We found strong coherence in the temporal variation in postsmolt marine survival among the 13 SU of NA and SE. A common North Atlantic trend explains 37% of the temporal variability of the survivals for the 13 SU and declines by a factor of 1.8 over the 1971–2014 time series. Synchrony in survival trends is stronger between SU within each CSG. The common trends at the scale of NA and SE capture 60% and 42% of the total variance of temporal variations, respectively. Temporal variations of the postsmolt survival are best explained by the temporal variations of sea surface temperature (SST, negative correlation) and net primary production indices (PP, positive correlation) encountered by salmon in common domains during their marine migration. Specifically, in the Labrador Sea/Grand Banks for populations from NA, 26% and 24% of variance is captured by SST and PP, respectively and in the Norwegian Sea for populations from SE, 21% and 12% of variance is captured by SST and PP, respectively. The findings support the hypothesis of a response of salmon populations to large climate‐induced changes in the North Atlantic simultaneously impacting populations from distant continental habitats.     

Testing the limits of resistance: a 19‐year study of Mediterranean grassland response to grazing regimes

A synthesis of a long‐term (19 years) study assessing the effects of cattle grazing on the structure and composition of a Mediterranean grassland in north‐eastern Israel is presented, with new insights into the response of the vegetation to grazing management and rainfall. We hypothesized that the plant community studied would be resistant to high grazing intensities and rainfall variability considering the combined long history of land‐use and unpredictable climatic conditions where this community evolved. Treatments included manipulations of stocking densities (moderate, heavy, and very heavy) and of grazing regimes (continuous vs. seasonal), in a factorial design. The effect of interannual rainfall variation on the expression of grazing impacts on the plant community was minor. The main effects of grazing on relative cover of plant functional groups were related to early vs. late seasonal grazing. Species diversity and equitability were remarkably stable across all grazing treatments. A reduction in tall grass cover at higher stocking densities was correlated with increased cover of less palatable groups such as annual and perennial thistles, as well as shorter and prostrate groups such as short annual grasses. This long‐term study shows that interannual fluctuations in plant functional group composition could be partly accounted for by grazing pressure and timing, but not by the measured rainfall variables. Grazing affected the dominance of tall annual grasses. However, the persistence of tall grasses and more palatable species over time, despite large differences in grazing pressure and timing, supports the idea that Mediterranean grasslands are highly resistant to prolonged grazing. Indeed, even under the most extreme grazing conditions applied, there were no signs of deterioration or collapse of the ecosystem. This high resistance to grazing intensity and interannual fluctuation in climatic conditions should favor the persistence of the plant community under forecasted increasing unpredictability due to climate change.