How exceptional are the classic adaptive radiations of passerine birds?

We investigated whether celebrated cases of evolutionary radiations of passerine birds on islands have produced exceptional morphological diversity relative to comparable-aged radiations globally. Based on eight external measurements, we calculated the disparity in size and shape within clades, each of which was classified as being tropical or temperate and as having diversified in a continental or an island/archipelagic setting. We found that the distribution of disparity among all clades does not differ substantively from a normal distribution, which would be consistent with a common underlying process of morphological diversification that is largely independent of latitude and occurrence on islands. Disparity is slightly greater in island clades than in those from continents or clades consisting of island and noninsular taxa, revealing a small, but significant, effect of island occurrence on evolutionary divergence. Nonetheless, the number of highly disparate clades overall is no greater than expected from a normal distribution, calling into question the need to invoke key innovations, ecological opportunity, or other factors as stimuli for adaptive radiations in passerine birds.

Evolutionary and Ecological Processes Underlying Geographic Variation in Innate Bird Songs.

AbstractEcological and evolutionary processes underlying spatial variation in signals involved in mate recognition and reproductive isolation are crucial to understanding the causes of population divergence and speciation. Here, to test hypotheses concerning the causes of song divergence, we examine how songs of two sister species of Atlantic Forest suboscine birds with innate songs, the Pyriglena fire-eye antbirds, vary across their ranges. Specifically, we evaluated the influence of isolation by distance and introgressive hybridization, as well as morphological and environmental variation, on geographic variation in male songs. Analyses based on 496 male vocalizations from 63 locations across a 2,200-km latitudinal transect revealed clinal changes in the structure of songs and showed that introgressive hybridization increases both the variability and the homogenization of songs in the contact zone between the two species. We also found that isolation by distance, morphological constraints, the environment, and genetic introgression independently predicted song variation across geographic space. Our study shows the importance of an integrative approach that investigates the roles of distinct ecological and evolutionary processes that influence acoustic signal evolution.

Improving species delimitation for effective conservation: a case study in the endemic maple-leaf oak (Quercus acerifolia).

Species delimitation is challenging in lineages that exhibit both high plasticity and introgression. This challenge can be compounded by collection biases, which may downweight specimens morphologically intermediate between traditional species. Additionally, mismatch between named species and observable phenotypes can compromise species conservation. We studied the species boundaries of Quercus acerifolia, a tree endemic to Arkansas, U.S. We performed morphometric analyses of leaves and acorns from 527 field and 138 herbarium samples of Q. acerifolia and its close relatives, Q. shumardii and Q. rubra. We employed two novel approaches: sampling ex situ collections to detect phenotypic plasticity caused by environmental variation and comparing random field samples with historical herbarium samples to identify collection biases that might undermine species delimitation. To provide genetic evidence, we also performed molecular analyses on genome-wide SNPs. Quercus acerifolia shows distinctive morphological, ecological, and genomic characteristics, rejecting the hypothesis that Q. acerifolia is a phenotypic variant of Q. shumardii. We found mismatches between traditional taxonomy and phenotypic clusters. We detected underrepresentation of morphological intermediates in herbarium collections, which may bias species discovery and recognition. Rare species conservation requires considering and addressing taxonomic problems related to phenotypic plasticity, mismatch between taxonomy and morphological clusters, and collection biases.

Host-specific soil microbes contribute to habitat restriction of closely related oaks (Quercus spp.).

Habitat divergence among close relatives is a common phenomenon. Studying the mechanisms behind habitat divergence is fundamental to understanding niche partitioning, species diversification, and other evolutionary processes. Recent studies found that soil microbes regulate the abundance and diversity of plant species. However, it remains unclear whether soil microbes can affect the habitat distributions of plants and drive habitat divergence. To fill in this knowledge gap, we investigated whether soil microbes might restrict habitat distributions of closely related oaks (Quercus spp.) in eastern North America. We performed a soil inoculum experiment using two pairs of sister species (i.e., the most closely related species) that show habitat divergence: Quercus alba (local species) vs. Q. michauxii (foreign), and Q. shumardii (local) vs. Q. acerifolia (foreign). To test whether host-specific soil microbes are responsible for habitat restriction, we investigated the impact of local sister live soil (containing soil microbes associated with local sister species) on the survival and growth of local and foreign species. Second, to test whether habitat-specific soil microbes are responsible for habitat restriction, we examined the effect of local habitat live soil (containing soil microbes within local sister's habitats, but not directly associated with local sister species) on the seedlings of local and foreign species. We found that local sister live soil decreased the survival and biomass of foreign species' seedlings while increasing those of local species, suggesting that host-specific soil microbes could potentially mediate habitat exclusion. In contrast, local habitat live soil did not differentially affect the survival or biomass of the local vs. foreign species. Our study indicates that soil microbes associated with one sister species can suppress the recruitment of the other host species, contributing to the habitat partitioning of close relatives. Considering the complex interactions with soil microbes is essential for understanding the habitat distributions of closely related plants.

Haemosporidian parasites and incubation period influence plumage coloration in tanagers (Passeriformes: Thraupidae).

Birds are highly visually oriented and use plumage coloration as an important signalling trait in social communication. Hence, males and females may have different patterns of plumage coloration, a phenomenon known as sexual dichromatism. Because males tend to have more complex plumages, sexual dichromatism is usually attributed to female choice. However, plumage coloration is partly condition-dependent; therefore, other selective pressures affecting individuals' success may also drive the evolution of this trait. Here, we used tanagers as model organisms to study the relationships between dichromatism and plumage coloration complexity in tanagers with parasitism by haemosporidians, investment in reproduction and life-history traits. We screened blood samples from 2849 individual birds belonging to 52 tanager species to detect haemosporidian parasites. We used publicly available data for plumage coloration, bird phylogeny and life-history traits to run phylogenetic generalized least-square models of plumage dichromatism and complexity in male and female tanagers. We found that plumage dichromatism was more pronounced in bird species with a higher prevalence of haemosporidian parasites. Lastly, high plumage coloration complexity in female tanagers was associated with a longer incubation period. Our results indicate an association between haemosporidian parasites and plumage coloration suggesting that parasites impact mechanisms of sexual selection, increasing differences between the sexes, and social (non-sexual) selection, driving females to develop more complex coloration.

Host life-history traits predict haemosporidian parasite prevalence in tanagers (Aves: Thraupidae).

Vector-borne parasites are important ecological drivers influencing life-history evolution in birds by increasing host mortality or susceptibility to new diseases. Therefore, understanding why vulnerability to infection varies within a host clade is a crucial task for conservation biology and for understanding macroecological life-history patterns. Here, we studied the relationship of avian life-history traits and climate on the prevalence of Plasmodium and Parahaemoproteus parasites. We sampled 3569 individual birds belonging to 53 species of the family Thraupidae. Individuals were captured from 2007 to 2018 at 92 locations. We created 2 phylogenetic generalized least-squares models with Plasmodium and Parahaemoproteus prevalence as our response variables, and with the following predictor variables: climate PC1, climate PC2, body size, mixed-species flock participation, incubation period, migration, nest height, foraging height, forest cover, and diet. We found that Parahaemoproteus and Plasmodium prevalence was higher in species inhabiting open habitats. Tanager species with longer incubation periods had higher Parahaemoproteus prevalence as well, and we hypothesize that these longer incubation periods overlap with maximum vector abundances, resulting in a higher probability of infection among adult hosts during their incubation period and among chicks. Lastly, we found that Plasmodium prevalence was higher in species without migratory behaviour, with mixed-species flock participation, and with an omnivorous or animal-derived diet. We discuss the consequences of higher infection prevalence in relation to life-history traits in tanagers.

Beta diversity, prevalence, and specificity of avian haemosporidian parasites throughout the annual cycle of Chilean Elaenia (Elaenia chilensis), a Neotropical austral migrant.

Migratory birds are implicated in dispersing haemosporidian parasites over great geographic distances. However, their role in sharing these vector-transmitted blood parasites with resident avian host species along their migration flyway is not well understood. We studied avian haemosporidian parasites in 10 localities where Chilean Elaenia, a long-distance Neotropical austral migrant species, spends part of its annual cycle to determine local parasite transmission among resident sympatric host species in the elaenia's distributional range across South America. We sampled 371 Chilean Elaenias and 1,818 birds representing 243 additional sympatric species from Brazilian wintering grounds to Argentinian breeding grounds. The 23 haemosporidian lineages found in Chilean Elaenias exhibited considerable variation in distribution, specialization, and turnover across the 10 avian communities in South America. Parasite lineage dissimilarity increased with geographic distance, and infection probability by Parahaemoproteus decreased in localities harbouring a more diverse haemosporidian fauna. Furthermore, blood smears from migrating Chilean Elaenias and local resident avian host species did not contain infective stages of Leucocytozoon, suggesting that transmission did not take place in the Brazilian stopover site. Our analyses confirm that this Neotropical austral migrant connects avian host communities and transports haemosporidian parasites along its distributional range in South America. However, the lack of transmissive stages at stopover site and the infrequent parasite lineage sharing between migratory host populations and residents at breeding and wintering grounds suggest that Chilean Elaenias do not play a significant role in dispersing haemosporidian parasites, nor do they influence local transmission across South America.

Evolution of coastal forests based on a full set of mangrove genomes.

Genomic studies are now poised to explore whole communities of species. The ~70 species of woody plants that anchor the coastal ecosystems of the tropics, collectively referred to as mangroves, are particularly suited to this exploration. In this study, we de novo sequenced the whole genomes of 32 mangroves, which we combined with other sequences of 30 additional species, comprising almost all mangroves globally. These community-wide genomic data will be valuable for ecology, evolution and biodiversity research. While the data revealed 27 independent origins of mangroves, the total phylogeny shows only modest increases in species number, even in coastal areas of active speciation, suggesting that mangrove extinction is common. A possible explanation for common extinction is the frequent sea-level rises and falls (SLRs and SLFs) documented in the geological record. Indeed, near-extinctions of species with extremely small population size (N) often happened during periods of rapid SLR, as revealed by the genome-wide heterozygosity of almost all mangroves. Reduction in N has possibly been further compounded by population fragmentation and the subsequent accumulation of deleterious mutations, thus pushing mangroves even closer to extinction. Crucially, the impact of the next SLR will be exacerbated by human encroachment into these mangrove habitats, potentially altering the ecosystems of tropical coasts irreversibly.

Evolutionary assembly of flowering plants into sky islands.

Alpine floras (plants in the vegetation belts above the climatic treelines) experience cold climates, particularly in temperate mountains during winter, but they are generally species-rich. Yet, biogeographers have not determined whether these floras represent evolutionarily independent but convergent assemblages drawn from their regional floras, or whether they originated from particular clades pre-adapted to harsh conditions. Here, we analyse the evolutionary relationships of angiosperm (flowering plant) species in 63 alpine floras worldwide (~7,000 species) in comparison with their regional floras (~94,000 species) and with the entire global flora. We find that each of the alpine floras represents an assemblage of more closely related species than their respective regional floras. The degree of phylogenetic clustering of species in alpine floras in tropical mountains exceeds that in temperate mountains. However, in relation to the global flora, temperate alpine floras are phylogenetically closely related subsets of floras that colonized cold temperate areas during interglacial periods. We conclude that alpine floras include a few dominant families that have evolved tolerance to low temperature, and that evolutionary niche conservatism explains their phylogenetic clustering, compared with species in their regional species pools.

Loss of forest cover and host functional diversity increases prevalence of avian malaria parasites in the Atlantic Forest.

Host phylogenetic relatedness and ecological similarity are thought to contribute to parasite community assembly and infection rates. However, recent landscape level anthropogenic changes may disrupt host-parasite systems by impacting functional and phylogenetic diversity of host communities. We examined whether changes in host functional and phylogenetic diversity, forest cover, and minimum temperature influence the prevalence, diversity, and distributions of avian haemosporidian parasites (genera Haemoproteus and Plasmodium) across 18 avian communities in the Atlantic Forest. To explore spatial patterns in avian haemosporidian prevalence and taxonomic and phylogenetic diversity, we surveyed 2241 individuals belonging to 233 avian species across a deforestation gradient. Mean prevalence and parasite diversity varied considerably across avian communities and parasites responded differently to host attributes and anthropogenic changes. Avian malaria prevalence (termed herein as an infection caused by Plasmodium parasites) was higher in deforested sites, and both Plasmodium prevalence and taxonomic diversity were negatively related to host functional diversity. Increased diversity of avian hosts increased local taxonomic diversity of Plasmodium lineages but decreased phylogenetic diversity of this parasite genus. Temperature and host phylogenetic diversity did not influence prevalence and diversity of haemosporidian parasites. Variation in the diversity of avian host traits that promote parasite encounter and vector exposure (host functional diversity) partially explained the variation in avian malaria prevalence and diversity. Recent anthropogenic landscape transformation (reduced proportion of native forest cover) had a major influence on avian malaria occurrence across the Atlantic Forest. This suggests that, for Plasmodium, host phylogenetic diversity was not a biotic filter to parasite transmission as prevalence was largely explained by host ecological attributes and recent anthropogenic factors. Our results demonstrate that, similar to human malaria and other vector-transmitted pathogens, prevalence of avian malaria parasites will likely increase with deforestation.

Development syndromes in New World temperate and tropical songbirds.

We studied avian development in 49 to 153 species of temperate and tropical New World passerine birds to determine how growth rates, and incubation and nestling periods, varied in relation to other life-history traits. We collected growth data and generated unbiased mass and tarsus growth rate estimates (mass n = 92 species, tarsus n = 49 species), and measured incubation period (n = 151) and nestling period (n = 153), which we analyzed with respect to region, egg mass, adult mass, clutch size, parental care type, nest type, daily nest predation rate (DMR), and nest height. We investigated covariation of life-history and natural-history attributes with the four development traits after controlling for phylogeny. Species in our lowland tropical sample grew 20% (incubation period), 25% (mass growth rate), and 26% (tarsus growth rate) more slowly than in our temperate sample. Nestling period did not vary with respect to latitude, which suggests that tropical songbirds fledge in a less well-developed state than temperate species. Suboscine species typically exhibited slower embryonic and post-embryonic growth than oscine passerines regardless of their breeding region. This pattern of slow development in tropical species could reflect phylogenetic effects based on unknown physiological attributes. Time-dependent nest mortality was unrelated to nestling mass growth rate, tarsus growth rate, and incubation period, but was significantly associated with nestling period. This suggests that nest predation, the predominant cause of nest loss in songbirds, does not exert strong selection on physiologically constrained traits, such as embryonic and post-embryonic growth, among our samples of temperate and lowland tropical songbird species. Nestling period, which is evolutionarily more labile than growth rate, was significantly shorter in birds exposed to higher rates of nest loss and nesting at lower heights, among other traits. Differences in life-history variation across latitudes provide insight into how unique ecological characteristics of each region influence physiological processes of passerines, and thus, how they can shape the evolution of life histories. While development traits clearly vary with respect to latitude, trait distributions overlap broadly. Life-history and natural history associations differ for each development trait, which suggests that unique selective pressures or constraints influence the evolution of each trait.

Haemosporidian parasites of resident and wintering migratory birds in The Bahamas.

In temperate regions, some avian haemosporidian parasites have evolved seasonal transmission strategies, with chronic infections relapsing during spring and transmission peaking during the hosts' breeding season. Because lineages with seasonal transmission strategies are unlikely to produce gametocytes in winter, we predicted that (1) resident birds living within wintering areas of Neotropical migrants would unlikely be infected with North American parasite lineages; and (2) if infected, wintering migratory birds would be more likely to harbor Plasmodium spp. rather than Parahaemoproteus spp. or Haemoproteus spp. parasites in their bloodstreams, as only Plasmodium produces life stages, other than gametocytes, that infect red blood cells. To test these predictions, we used molecular detection and microscopy to compare the diversity and prevalence of haemosporidian parasites among year-round residents and wintering migratory birds during February 2016, on three islands of The Bahamas archipelago, i.e., Andros, Grand Bahama, and Great Abaco. Infection prevalence was low and comparable between migratory (15/111) and resident (15/129) individuals, and it did not differ significantly among islands. Out of the 12 lineages detected infecting migratory birds, five were transmitted in North America; four lineages could have been transmitted during breeding, wintering, or migration; and three lineages were likely transmitted in The Bahamas. Resident birds mostly carried lineages endemic to the Caribbean region. All North American-transmitted parasite lineages detected among migratory birds were Plasmodium spp. Our findings suggest that haemosporidian parasites of migrants shift resource allocation seasonally, minimizing the production of gametocytes during winter, with low risk of infection spillover to resident birds.

Nest attendance by tropical and temperate passerine birds: Same constancy, different strategy.

Parental care in birds varies among species and geographic regions. Incubation behavior influences embryonic development rate and varies substantially among species.We studied attendance at the nest by videoing nests or collecting data from the literature for 112 species in north temperate and lowland tropical sites, then associated patterns of incubation on- and off-bouts with species and environmental traits.Songbirds nesting at low elevations incubate their eggs for an average of 74.1% (±12.9 SD, n = 60 species) of the time in temperate regions and 71.0% (±12.2 SD, n = 52 species) in tropical regions during daylight hours, and 84.3% (±8.2 SD) and 85.3% (±6.2 SD), respectively, of each 24-hr cycle.While these attendance percentages do not differ significantly between latitudes, our data also show that lowland tropical songbirds make fewer visits to the nest and, consequently, have longer on-bouts and off-bouts during incubation. This pattern in attendance reflects a latitudinal contrast in parental care strategy, where lowland tropical birds reduce visits to the nest by increasing on- and off-bout lengths while maintaining the same proportion of time spent incubating their eggs (constancy).Similar constancy across latitude suggests that tropical and temperate birds may be similarly constrained to maintain elevated egg temperatures for normal embryo growth.The different attendance strategies adopted in each region may reflect differences in ambient temperature, adult foraging time, and nest predation rate. Consistently warm ambient temperatures likely allow tropical birds to take longer off-bouts, and thereby to reduce activity around the nest, compared to temperate birds.

Phylogenetic dispersion and diversity in regional assemblages of seed plants in China.

Species assemble into communities through ecological and evolutionary processes. Phylogenetic niche conservatism-the tendency of species to retain ancestral ecological distributions-is thought to influence which species from a regional species pool can persist in a particular environment. We analyzed data for seed plants in China to test hypotheses about the distribution of species within regional floras. Of 16 environmental variables, actual evapotranspiration, minimum temperature of the coldest month, and annual precipitation most strongly influenced regional species richness, phylogenetic dispersion, and phylogenetic diversity for both gymnosperms (cone-bearing plants) and angiosperms (flowering plants). For most evolutionary clades at, and above, the family level, the relationships between metrics of phylogenetic dispersion (i.e., average phylogenetic distance among species), or phylogenetic diversity, and the 3 environmental variables were consistent with the tropical niche conservatism hypothesis, which predicts closer phylogenetic relatedness and reduced phylogenetic diversity with increasing environmental stress. The slopes of the relationships between phylogenetic relatedness and the 3 environmental drivers identified in this analysis were steeper for primarily tropical clades, implying greater niche conservatism, than for primarily temperate clades. These observations suggest that the distributions of seed plants across large-scale environmental gradients in China are constrained by conserved adaptations to the physical environment, i.e., phylogenetic niche conservatism.

Population structure of avian malaria parasites.

The geographic distribution of genetic diversity in malaria parasite populations (Apicomplexa: Haemosporida) presumably influences local patterns of virulence and the evolution of host-resistance, but little is known about population genetic structure in these parasites. We assess the distribution of genetic diversity in the partial Domain I of apical membrane antigen 1 (AMA1) in three mtDNA-defined lineages of avian Plasmodium to determine spatial population structure and host-parasite genetic relationships. We find that one parasite lineage is genetically differentiated in association with a single host genus and among some locations, but not with respect to other hosts. Two other parasite lineages are undifferentiated with respect to host species but exhibit geographic differentiation that is inconsistent with shared geographic barriers or with isolation-by-distance. Additional differentiation within two other lineages is unassociated with host species or location; in one case, we tentatively interpret this differentiation as the result of mitochondrial introgression from one of the lineages into a second lineage. More sampling of nuclear genetic diversity within populations of avian Plasmodium is needed to rule out coinfection as a possible confounding factor. If coinfections are not responsible for these findings, further assessment is needed to determine the frequency of mitonuclear discordance and its implications for defining parasite lineages based on mitochondrial genetic variation. OPEN RESEARCH BADGES: This article has earned an Open Data Badge for making publicly available the digitally-shareable data necessary to reproduce the reported results. The data is available at Genbank https://www.ncbi.nlm.nih.gov/genbank/, accession numbers MK965548-MK965653 and MK929797-MK930264.

Speciation Rate Is Independent of the Rate of Evolution of Morphological Size, Shape, and Absolute Morphological Specialization in a Large Clade of Birds.

Whether ecological differences between species evolve in parallel with lineage diversification is a fundamental issue in evolutionary biology. These processes might be connected if conditions that favor the proliferation of species, such as release from competitors, facilitate the evolution of novel ecological relationships. Despite this, phylogenetic studies do not consistently identify such a connection. Conversely, if higher diversity caused species to become increasingly specialized ecologically, then lineage diversification might become dissociated from ecological diversification. In this analysis, we ask whether the rate of lineage diversification in a large clade of birds is correlated with morphological specialization and with rates of morphological evolution. We find that morphological variation is related to species richness within clades but that rates of morphological evolution are decoupled from the rate of lineage diversification. Additionally, morphological specialization within lineages is independent of the rate at which lineages diversify, with the results apparently robust against false negative inference. This dissociation is likely a consequence of the major ecomorphological differences between avian clades arising early in their evolutionary history, with comparatively little variation added subsequently, while avian diversification has been driven predominantly by geographic isolation and sexual selection. Accordingly, biodiversity appears to be limited by the extent to which taxa can subdivide exploited regions of ecological space and not just overall ecological opportunity.

The global biogeography of avian haemosporidian parasites is characterized by local diversification and intercontinental dispersal.

The biogeographic histories of parasites and pathogens are infrequently compared with those of free-living species, including their hosts. Documenting the frequency with which parasites and pathogens disperse across geographic regions contributes to understanding not only their evolution, but also the likelihood that they may become emerging infectious diseases. Haemosporidian parasites of birds (parasite genera Plasmodium, Haemoproteus and Leucocytozoon) are globally distributed, dipteran-vectored parasites. To date, over 2000 avian haemosporidian lineages have been designated by molecular barcoding methods. To achieve their current distributions, some lineages must have dispersed long distances, often over water. Here we quantify such events using the global avian haemosporidian database MalAvi and additional records primarily from the Americas. We scored lineages as belonging to one or more global biogeographic regions based on infection records. Most lineages were restricted to a single region but some were globally distributed. We also used part of the cytochrome b gene to create genus-level parasite phylogenies and scored well-supported nodes as having descendant lineages in regional sympatry or allopatry. Descendant sister lineages of Plasmodium, Haemoproteus and Leucocytozoon were distributed in allopatry in 11, 16 and 15% of investigated nodes, respectively. Although a small but significant fraction of the molecular variance in cytochrome b of all three genera could be explained by biogeographic region, global parasite dispersal likely contributed to the majority of the unexplained variance. Our results suggest that avian haemosporidian parasites have faced few geographic barriers to dispersal over their evolutionary history.

Host species, and not environment, predicts variation in blood parasite prevalence, distribution, and diversity along a humidity gradient in northern South America.

Environmental factors strongly influence the ecology and evolution of vector-borne infectious diseases. However, our understanding of the influence of climatic variation on host-parasite interactions in tropical systems is rudimentary. We studied five species of birds and their haemosporidian parasites (Plasmodium and Haemoproteus) at 16 sampling sites to understand how environmental heterogeneity influences patterns of parasite prevalence, distribution, and diversity across a marked gradient in water availability in northern South America. We used molecular methods to screen for parasite infections and to identify parasite lineages. To characterize spatial heterogeneity in water availability, we used weather-station and remotely sensed climate data. We estimated parasite prevalence while accounting for spatial autocorrelation, and used a model selection approach to determine the effect of variables related to water availability and host species on prevalence. The prevalence, distribution, and lineage diversity of haemosporidian parasites varied among localities and host species, but we found no support for the hypothesis that the prevalence and diversity of parasites increase with increasing water availability. Host species and host × climate interactions had stronger effects on infection prevalence, and parasite lineages were strongly associated with particular host species. Because climatic variables had little effect on the overall prevalence and lineage diversity of haemosporidian parasites across study sites, our results suggest that independent host-parasite dynamics may influence patterns in parasitism in environmentally heterogeneous landscapes.

Emma Lucy Braun's forest plots in eastern North America.

Relative abundances of tree species are presented for the 348 forest plots described in E. Lucy Braun's (1950) book, Deciduous Forests of Eastern North America (Hafner, New York, facsimile reprint 1972). Information about the plots includes forest type, location with latitude and longitude, WorldClim climate variables, and sources of original studies where applicable. No copyright restrictions are associated with the use of this data set. Please cite this article when the data are used in other publications.