Royal British Columbia Museum

Abstract Systematic assessments of species extinction risk at regular intervals are necessary for informing conservation action 1,2 . Ongoing developments in taxonomy, threatening processes and research further underscore the need for reassessment 3,4 . Here we report the findings of the second Global Amphibian Assessment, evaluating 8,011 species for the International Union for Conservation of Nature Red List of Threatened Species. We find that amphibians are the most threatened vertebrate class (40.7% of species are globally threatened). The updated Red List Index shows that the status of amphibians is deteriorating globally, particularly for salamanders and in the Neotropics. Disease and habitat loss drove 91% of status deteriorations between 1980 and 2004. Ongoing and projected climate change effects are now of increasing concern, driving 39% of status deteriorations since 2004, followed by habitat loss (37%). Although signs of species recoveries incentivize immediate conservation action, scaled-up investment is urgently needed to reverse the current trends.

The widespread extinctions of large mammals at the end of the Pleistocene epoch have often been attributed to the depredations of humans; here we present genetic evidence that questions this assumption. We used ancient DNA and Bayesian techniques to reconstruct a detailed genetic history of bison throughout the late Pleistocene and Holocene epochs. Our analyses depict a large diverse population living throughout Beringia until around 37,000 years before the present, when the population's genetic diversity began to decline dramatically. The timing of this decline correlates with environmental changes associated with the onset of the last glacial cycle, whereas archaeological evidence does not support the presence of large populations of humans in Eastern Beringia until more than 15,000 years later.

It is widely accepted, based on data from the last few decades and on model simulations, that anthropogenic climate change will cause increased fire activity. However, less attention has been paid to the relationship between abrupt climate changes and heightened fire activity in the paleorecord. We use 35 charcoal and pollen records to assess how fire regimes in North America changed during the last glacial-interglacial transition (15 to 10 ka), a time of large and rapid climate changes. We also test the hypothesis that a comet impact initiated continental-scale wildfires at 12.9 ka; the data do not support this idea, nor are continent-wide fires indicated at any time during deglaciation. There are, however, clear links between large climate changes and fire activity. Biomass burning gradually increased from the glacial period to the beginning of the Younger Dryas. Although there are changes in biomass burning during the Younger Dryas, there is no systematic trend. There is a further increase in biomass burning after the Younger Dryas. Intervals of rapid climate change at 13.9, 13.2, and 11.7 ka are marked by large increases in fire activity. The timing of changes in fire is not coincident with changes in human population density or the timing of the extinction of the megafauna. Although these factors could have contributed to fire-regime changes at individual sites or at specific times, the charcoal data indicate an important role for climate, and particularly rapid climate change, in determining broad-scale levels of fire activity.

This study reports DNA barcodes for more than 1300 Lepidoptera species from the eastern half of North America, establishing that 99.3 per cent of these species possess diagnostic barcode sequences. Intraspecific divergences averaged just 0.43 per cent among this assemblage, but most values were lower. The mean was elevated by deep barcode divergences (greater than 2%) in 5.1 per cent of the species, often involving the sympatric occurrence of two barcode clusters. A few of these cases have been analysed in detail, revealing species overlooked by the current taxonomic system. This study also provided a large-scale test of the extent of regional divergence in barcode sequences, indicating that geographical differentiation in the Lepidoptera of eastern North America is small, even when comparisons involve populations as much as 2800 km apart. The present results affirm that a highly effective system for the identification of Lepidoptera in this region can be built with few records per species because of the limited intra-specific variation. As most terrestrial and marine taxa are likely to possess a similar pattern of population structure, an effective DNA-based identification system can be developed with modest effort.

The evaluation of the cultural significance of plants in ethnobotanical studies is an essential step in various types of investigations, including research on lexical retention of plant names in diverging languages, on trade and material exchange between groups, on subsistence strategies, and on folk classification. An index of cultural significance was developed, based on ethnobotanical data on Thompson and Lillooet, two Interior Salish groups of British Columbia. For a given plant taxon, the index is a composite of a wide variety of potential applications of a plant, ranked according to the contribution of each separate application to survival in traditional cultures, together with estimates of intensity and exclusivity of use for each. Example calculations of the derivation of the index are provided, as well as a summary of calculations for more than 500 plant species from Thompson and Lillooet territories. The index has some limitations, but overall it seems to provide a meaningful and valid assessment of the relative importance of a plant taxon in Thompson and Lillooet.

A list of all valid 6,206 extant and 296 fossil species of Ceratopogonidae described worldwide is provided, along with all their synonyms. A full citation and the country of origin of the type is given, with some larger countries also providing a more specific state or province. For the first time, worldwide, nomina dubia are identified. Numbers of species of each genus and subgenus are listed. Within subfamilies and tribes, genera are listed alphabetically. Five species have newly recognized authors, four have new names and 28 new combinations are recognized, with these listed in a table. A commentary on the state of the systematics in the family and particularly of Culicoides Latreille is given. The museums of the world are listed with the types of various authors of Ceratopogonidae species indicated. Authors providing regional catalogs, as well as summation of various collections are tabulated. The rate of description since 1758 indicates a steady progression of description, with, for example, 1,231 valid species described since the compilation of the world species by Borkent Wirth (1997), till the end of 2018. The diversity in each Region is compared and the numbers of species shared between adjacent Regions presented.

British Columbia Holocene vegetation and climate is reconstructed from pollen records. A coastal Pinus contorta paleobiome developed after glacier retreat under cool and probably dry climate. Cool moist forests involving Picea, Abies, Tsuga spp., and Pinus followed until the early Holocene. Pseudotsuga menziesii arrived and spread in the south 10 000-9000 BP, and Picea sitchensis - Tsuga heterophylla forests developed in the north. T. heterophylla increased 7500-7000 BP, and Cupressaceae expanded 5000-4000 BP. Bogs began to develop and expland. Modern vegetation arose 4000-2000 BP. There were early Holocene grass and Artemisia communities at mid-elevations and pine stands at high elevations in southern interior B.C. Forests expanded downslope and lakes formed 8500-7000 BP. Modern forests arose 4500-4000 BP while lower and upper tree lines declined. In northern B.C. non-arboreal communities preceded middle Holocene Picea forests. Abies, Pinus and Picea mariana predominated at various sites after 4000 BP. At 6000 BP Tsuga heterophylla (south) and Picea sitchensis (north) dominated the coast and islands and Quercus garryana and Pseudotsuga on southeast Vancouver Island, but Thuja plicata was infrequent. Southern Interior Plateau vegetation at 6000 BP was more open than today at middle to lower elevations, whereas forests covered the Northern Interior Plateau. Picea forests occurred in northern B.C. Holocene climate phases were: 1) warm dry "xerothermic" ca. 9500-7000 BP, 2) warm moist "mesothermic" ca. 7000-4500 BP, 3) moderate and moist 4500-0 BP, with increasing moisture 8500-6000 BP and cooling (?increased moisture) 4500-3000 BP. B.Cs Hypsithermal had dry and wet stages; 6000 BP occurred in the warm and wet mesothermic stage.

Abstract Ice types, albedos and impurity content are characterized for the ablation zone of the Greenland ice sheet in Kronprinz Christians Land (80° N, 24° W). Along this ice margin the width of the ablation zone is only about 8 km. The emergence and melting of old ice in the ablation zone creates a surface layer of dust that was originally deposited with snowfall high on the ice sheet. This debris cover is augmented by locally derived wind-blown sediment. Subsequently, the surface dust particles often aggregate together to form centimetre-scale clumps that melt into the ice, creating cryoconite holes. The debris in the cryoconite holes becomes hidden from sunlight, raising the area-averaged albedo relative to surfaces with uniform debris cover. Spectral and broadband albedos were obtained for snow, ice hummocks, debris-covered ice, cryoconite-studded ice and barren tundra surfaces. Broadband ice albedos varied from 0.2 (for ice with heavy loading of uniform debris) to 0.6 (for ice hummocks with cryoconite holes). The cryoconite material itself has albedo 0.1 when wet. Areal distribution of the major surface types was estimated visually from a transect video as a function of distance from the ice edge (330 m a.s.l.). Ablation rates were measured along a transect from the ice margin to the slush zone 8 km from the margin (550 m a.s.l.), traversing both Pleistocene and Holocene ice. Ablation rates in early August averaged 2 cm d −1 . Impurity concentrations were typically 4.3 mg L −1 in the subsurface ice. Surface concentrations were about 16 g m −2 on surfaces with low impurity loading, and heavily loaded surfaces had concentrations as high as 1.4 kg m −2 . The mineralogical composition of the cryoconite material is comparable with that of the surrounding soils and with dust on a snowdrift in front of the ice margin, implying that much of the material is derived from local sources. A fine mode (clay) is present in the oldest ice but not in the nearby soil, suggesting that its origin is from wind deposition during Pleistocene glaciation.

Millennial-scale records of forest fire provide important baseline information for ecosystem management, especially in regions with too few recent fires to describe the historical range of variability. Charcoal records from lake sediments and soil profiles are well suited for reconstructing the incidence of past fire and its relationship to changing climate and vegetation. We highlight several records from western North America and their relevance in reconstructing historical forest dynamics, fire–climate relationships, and feedbacks between vegetation and fire under climate change. Climatic effects on fire regimes are evident in many regions, but comparisons of paleofire records sometimes show a lack of synchrony, indicating that local factors substantially affect fire occurrence, even over long periods. Furthermore, the specific impacts of vegetation change on fire regimes vary among regions with different vegetation histories. By documenting the effects on fire patterns of major changes in climate and vegetation, paleo-fire records can be used to test the mechanistic models required for the prediction of future variations in fire.

Sponges are suspension feeders that use flagellated collar-cells (choanocytes) to actively filter a volume of water equivalent to many times their body volume each hour. Flow through sponges is thought to be enhanced by ambient current, which induces a pressure gradient across the sponge wall, but the underlying mechanism is still unknown. Studies of sponge filtration have estimated the energetic cost of pumping to be <1% of its total metabolism implying there is little adaptive value to reducing the cost of pumping by using "passive" flow induced by the ambient current. We quantified the pumping activity and respiration of the glass sponge Aphrocallistes vastus at a 150 m deep reef in situ and in a flow flume; we also modeled the glass sponge filtration system from measurements of the aquiferous system. Excurrent flow from the sponge osculum measured in situ and in the flume were positively correlated (r>0.75) with the ambient current velocity. During short bursts of high ambient current the sponges filtered two-thirds of the total volume of water they processed daily. Our model indicates that the head loss across the sponge collar filter is 10 times higher than previously estimated. The difference is due to the resistance created by a fine protein mesh that lines the collar, which demosponges also have, but was so far overlooked. Applying our model to the in situ measurements indicates that even modest pumping rates require an energetic expenditure of at least 28% of the total in situ respiration. We suggest that due to the high cost of pumping, current-induced flow is highly beneficial but may occur only in thin walled sponges living in high flow environments. Our results call for a new look at the mechanisms underlying current-induced flow and for reevaluation of the cost of biological pumping and its evolutionary role, especially in sponges.

Blood residues from several animal species have been discovered on the surfaces of chert, basalt, and obsidian prehistoric tools (1000 to 6000 years old) from open-air sites along the western coast and in the northern boreal forest of Canada. A screening test has been developed to detect residual blood. Hemoglobin has been crystallized from the residues, and the species of origin determined.

What are the greatest sizes that the largest marine megafauna obtain? This is a simple question with a difficult and complex answer. Many of the largest-sized species occur in the world's oceans. For many of these, rarity, remoteness, and quite simply the logistics of measuring these giants has made obtaining accurate size measurements difficult. Inaccurate reports of maximum sizes run rampant through the scientific literature and popular media. Moreover, how intraspecific variation in the body sizes of these animals relates to sex, population structure, the environment, and interactions with humans remains underappreciated. Here, we review and analyze body size for 25 ocean giants ranging across the animal kingdom. For each taxon we document body size for the largest known marine species of several clades. We also analyze intraspecific variation and identify the largest known individuals for each species. Where data allows, we analyze spatial and temporal intraspecific size variation. We also provide allometric scaling equations between different size measurements as resources to other researchers. In some cases, the lack of data prevents us from fully examining these topics and instead we specifically highlight these deficiencies and the barriers that exist for data collection. Overall, we found considerable variability in intraspecific size distributions from strongly left- to strongly right-skewed. We provide several allometric equations that allow for estimation of total lengths and weights from more easily obtained measurements. In several cases, we also quantify considerable geographic variation and decreases in size likely attributed to humans.

Glass sponges are conspicuous inhabitants of benthic communities in the cool waters of the Antarctic and north Pacific continental shelf. We used an ROV outfitted with a new device for simultaneous sampling of water inhaled and exhaled by the sponges to provide the first data on the nutritional ecology and metabolism of two glass sponge species in their natural deep‐water habitat (120–160 m). Aphrocallistes vastus and Rhabdocalyptus dawsoni were found to be mostly bacteriovores, removing up to 95% of the bacteria (median removal was 79% for both species) and heterotrophic protists (&lt;10 µm) from the water they filter. The relatively scarce microbial cells were efficiently selected from a ‘soup’ of suspended clay and detritus particles (microorganisms accounted for ~1% of the total ambient suspended solids). Removal of planktonic microorganisms (2.2 ± 1.3 µmol carbon [C] C L −1 and 0.37 ± 0.17 µmol nitrogen [N] L −1 ) accounted for the entire total organic C uptake and ammonium excretion by both species, with no evidence for dissolved organic uptake. Similar results were obtained in laboratory experiments in which dissolved organic C was directly measured. Despite the massive siliceous sponge skeleton, silica uptake was below detection levels (0.28 µmol L −1 ), supporting previous suggestions of low growth rates in Hexactinellida. Reported mean sponge abundances of &gt;1 individual m −2 indicate that the sponge filtering activity may significantly affect the deep microbial community and benthic‐pelagic mass exchange in some north‐east Pacific fjords.

A comparison of paleobotanical records with archeological and ethnographic evidence from the Pacific Northwest shows a strong correlation between the expansion of Western red cedar (Thuja plicata) in coastal forests between 5000 and 2500 years ago and the evolution of a massive woodworking technology by native cultures. This suggests that an important component of cultural development was environmentally constrained until large cedar trees, the basic resource for canoe-building and plank-house construction, had become available in late Holocene time.

We studied the evolutionary history of the Parmeliaceae (Lecanoromycetes, Ascomycota), one of the largest families of lichen-forming fungi with complex and variable morphologies, also including several lichenicolous fungi. We assembled a six-locus data set including nuclear, mitochondrial and low-copy protein-coding genes from 293 operational taxonomic units (OTUs). The lichenicolous lifestyle originated independently three times in lichenized ancestors within Parmeliaceae, and a new generic name is introduced for one of these fungi. In all cases, the independent origins occurred c. 24 million yr ago. Further, we show that the Paleocene, Eocene and Oligocene were key periods when diversification of major lineages within Parmeliaceae occurred, with subsequent radiations occurring primarily during the Oligocene and Miocene. Our phylogenetic hypothesis supports the independent origin of lichenicolous fungi associated with climatic shifts at the Oligocene-Miocene boundary. Moreover, diversification bursts at different times may be crucial factors driving the diversification of Parmeliaceae. Additionally, our study provides novel insight into evolutionary relationships in this large and diverse family of lichen-forming ascomycetes.

Study of all flies (Diptera) collected for one year from a four-hectare (150 x 266 meter) patch of cloud forest at 1,600 meters above sea level at Zurquí de Moravia, San José Province, Costa Rica (hereafter referred to as Zurquí), revealed an astounding 4,332 species. This amounts to more than half the number of named species of flies for all of Central America. Specimens were collected with two Malaise traps running continuously and with a wide array of supplementary collecting methods for three days of each month. All morphospecies from all 73 families recorded were fully curated by technicians before submission to an international team of 59 taxonomic experts for identification. Overall, a Malaise trap on the forest edge captured 1,988 species or 51% of all collected dipteran taxa (other than of Phoridae, subsampled only from this and one other Malaise trap). A Malaise trap in the forest sampled 906 species. Of other sampling methods, the combination of four other Malaise traps and an intercept trap, aerial/hand collecting, 10 emergence traps, and four CDC light traps added the greatest number of species to our inventory. This complement of sampling methods was an effective combination for retrieving substantial numbers of species of Diptera. Comparison of select sampling methods (considering 3,487 species of non-phorid Diptera) provided further details regarding how many species were sampled by various methods. Comparison of species numbers from each of two permanent Malaise traps from Zurquí with those of single Malaise traps at each of Tapantí and Las Alturas, 40 and 180 km distant from Zurquí respectively, suggested significant species turnover. Comparison of the greater number of species collected in all traps from Zurquí did not markedly change the degree of similarity between the three sites, although the actual number of species shared did increase. Comparisons of the total number of named and unnamed species of Diptera from four hectares at Zurquí is equivalent to 51% of all flies named from Central America, greater than all the named fly fauna of Colombia, equivalent to 14% of named Neotropical species and equal to about 2.7% of all named Diptera worldwide. Clearly the number of species of Diptera in tropical regions has been severely underestimated and the actual number may surpass the number of species of Coleoptera. Various published extrapolations from limited data to estimate total numbers of species of larger taxonomic categories (e.g., Hexapoda, Arthropoda, Eukaryota, etc.) are highly questionable, and certainly will remain uncertain until we have more exhaustive surveys of all and diverse taxa (like Diptera) from multiple tropical sites. Morphological characterization of species in inventories provides identifications placed in the context of taxonomy, phylogeny, form, and ecology. DNA barcoding species is a valuable tool to estimate species numbers but used alone fails to provide a broader context for the species identified.

BACKGROUND: Detecting and controlling the movements of invasive species, such as insect pests, relies upon rapid and accurate species identification in order to initiate containment procedures by the appropriate authorities. Many species in the tussock moth genus Lymantria are significant forestry pests, including the gypsy moth Lymantria dispar L., and consequently have been a focus for the development of molecular diagnostic tools to assist in identifying species and source populations. In this study we expand the taxonomic and geographic coverage of the DNA barcode reference library, and further test the utility of this diagnostic method, both for species/subspecies assignment and for determination of geographic provenance of populations. METHODOLOGY/PRINCIPAL FINDINGS: Cytochrome oxidase I (COI) barcodes were obtained from 518 individuals and 36 species of Lymantria, including sequences assembled and generated from previous studies, vouchered material in public collections, and intercepted specimens obtained from surveillance programs in Canada. A maximum likelihood tree was constructed, revealing high bootstrap support for 90% of species clusters. Bayesian species assignment was also tested, and resulted in correct assignment to species and subspecies in all instances. The performance of barcoding was also compared against the commonly employed NB restriction digest system (also based on COI); while the latter is informative for discriminating gypsy moth subspecies, COI barcode sequences provide greater resolution and generality by encompassing a greater number of haplotypes across all Lymantria species, none shared between species. CONCLUSIONS/SIGNIFICANCE: This study demonstrates the efficacy of DNA barcodes for diagnosing species of Lymantria and reinforces the view that the approach is an under-utilized resource with substantial potential for biosecurity and surveillance. Biomonitoring agencies currently employing the NB restriction digest system would gather more information by transitioning to the use of DNA barcoding, a change which could be made relatively seamlessly as the same gene region underlies both protocols.

been handled alive, in addition to a smaller number of freshly killed specimens.Three trained birds have been under constant observation for periods of one, two, and three ymrs, respectively.A series of over 25 aeries or aerie sites has been visited and ob- serve& of which ten, on Langara Island, have been re-visited and observed in five different years.Acknu&dgments.-This study was made posdble by the cooperation and direction of Dr. G. Clifford Carl, Director, Provincial

This worldwide biosystematic study of Corethrellidae, with its single genus Corethrella Coquillett, provides a complete compilation of all that is known for the group, both taxonomically and bionomically. Descriptions of each species are based primarily on the adults, summarize all bionomic information, and provide a map showing its distribution. Keys to the species of each region are provided. A total of 97 extant species is recognized, with 52 of these being new. Seven fossil species are described with two of these being new to science. All species, including 13 new synonyms, are cataloged in a table for easy reference. Seven extant species are of uncertain status because of damaged or missing types. Lectotypes and, depending on the species, some paralectotypes, are designated for the following species: C. inepta (Annandale), C. pallitarsis Edwards, C. picticollis Edwards, C. ananacola Dyar, C. calathicola Edwards, and C. brakeleyi (Coquillett). A cladistic analysis interprets most extant and fossil species (some are not interpretable at the present time) and provides the basis for zoogeographic and bionomic interpretation. Worldwide, Corethrella species are found between 50°N and 50°S but most are found between 30°N and 30°S and below 1500 meters in elevation. Because female adults are attracted to the call of male frogs and feed on their blood, species are restricted to areas where there are frogs. Phylogenetic patterns suggest Gondwanan connections for earlier lineages within the genus. At least one lineage has dispersed from the New World to southeast Asia and some species are located on volcanic islands in the Caribbean, indicating further instances of dispersal. It is certain that many more species are yet to be discovered. Phylogenetic patterns indicate that the immatures of Corethrella species have repeatedly moved between ground-dwelling habitats and phytotelmata, with the plesiotypic habitat likely being ground-dwelling. Some lineages have diversified within phytotelmata. Fossil, cladistic and morphological evidence indicates that Corethrella females have been feeding on calling frogs since at least the Early Cretaceous. Females likely hear their frog hosts using the Johnston’s Organ. There is some evidence of host specificity as well as selection of particular biting sites for some species of Corethrella. The females of at least some species of Corethrella transmit Trypanosoma Gruby between calling frogs and this association is also likely an ancient one.Este estudio biosistemático de Corethrellidae a nivel Mundial, con su único género Corethrella, proporciona una completa recopilación de todo lo conocido para el grupo, tanto desde el punto de vista taxonómico como bionómico. Las descripciones de cada especie se realizan primariamente sobre la base de adultos, resumen toda la información bionómica y proveen un mapa donde se muestra su distribución. Se brindan claves para especie de cada región. Se reconoce un total de 97 especies, 52 de las cuales son nuevas. Se describen siete especies fósiles, siendo dos de ellas nuevas para la Ciencia. Para una fácil referencia, todas las especies son catalogadas en una tabla, incluyendo 13 nuevos sinónimos. Debido a que sus tipos se hallan dañados o perdidos, siete especies actuales ostentan un status incierto. De acuerdo a la especie, son designados lectotipos o paralectotipos de las siguientes especies: C. inepta (Annandale), C. pallitarsis Edwards, C. picticollis Edwards, C. ananacola Dyar, C. calathicola Edwards, y C. brakeleyi (Coquillett). El análisis cladístico interpreta la mayoría de las especies actuales y fósiles (algunas no pueden ser interpretadas actualmente) y provee la base para interpretaciones zoogeográficas y bionómicas. Las especies de Corethrella se hallan entre 50°N y 50°S, aunque la mayoría se encuentran entre 30°N y 30°S y por debajo de 1500 metros de elevación. Debido a que las hembras adultas son atraídas por el llamado de ranas macho y se alimentan de su sangre, las especies están restringidas a las áreas donde se hallan ranas. Los patrones filogenéticos sugieren conecciones Gondwánicas para los linajes más antiguos del género. Al menos un linaje se ha dispersado desde el Nuevo Mundo hacia el sudeste de Asia, y algunas especies se hallan en islas volcánicas del Caribe, indicando otras instancias de dispersión. Con seguridad aún quedan muchas más especies por ser descubiertas. Los patrones filogenéticos indican que los inmaduros de las especies de Corethrella se han movido repetidamente entre habitats ubicados a nivel del suelo y fitotelmata, siendo probablemente

We reviewed the availability of cytochrome c oxidase subunit I (COI) sequences for 2534 North American freshwater invertebrate genera in public databases (GenBank and Barcode of Life Data Systems) and assessed representation of genera commonly encountered in the Canadian Aquatic Biomonitoring Network (CABIN) database. COI sequence records were available for 61.2% of North American genera and 72.4% of Insecta genera in public databases. Mollusca (73.9%) and Nematoda (15.4%) were the best and worst represented groups, respectively. In CABIN, 85.4% of genera had COI sequence records, and 95.2% of genera occurring in >1% of samples were represented. Genera absent from CABIN tended to be uncommon or members of groups not routinely used for biomonitoring purposes. On average, 94.1% of genera in well-identified samples had associated sequence data. To leverage the full potential of genomics approaches, we must expand DNA-barcode reference libraries for poorly described components of freshwater food webs. Some genera appear to be well represented (e.g., Eukiefferiella), but deposited sequences represent few sampling localities or few species and lead to underestimation of sequence diversity at the genus level and reduced confidence in identifications. Public COI libraries are sufficiently populated to permit routine application of genomics tools in biomonitoring, and ongoing quality assurance/quality control should include re-evaluation as new COI reference sequences are added or taxonomic hierarchies change. Next, we must understand whether and how established biomonitoring approaches can capitalize on high-throughput sequencing tools. Biomonitoring approaches that use genomics data to facilitate structural and functional assessments are fertile ground for future investigation and will benefit from continued improvement of publicly available sequence libraries.