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High-quality, chromosome-level reference genomes of the viviparous Caribbean skinks Spondylurus nitidus and S. culebrae

New World mabuyine skinks are a diverse radiation of morphologically cryptic lizards with unique reproductive biologies. Recent studies examining population-level data (morphological, ecological, and genomic) have uncovered novel biodiversity and phenotypes, including the description of dozens of new species and insights into the evolution of their highly complex placental structures. Beyond the potential for this diverse group to serve as a model for the evolution of viviparity in lizards, much of the taxonomic diversity is concentrated in regions experiencing increasing environmental instability from climate and anthropogenic change. Consequently, a better understanding of genome structure and diversity will be an important tool in the adaptive management and conservation of this group. Skinks endemic to Caribbean islands are particularly vulnerable to global change with several species already considered likely extinct and several remaining species either endangered or threatened. Combining PacBio long-read sequencing, Hi-C, and RNAseq data, here we present the first genomic resources for this group by describing new chromosome-level reference genomes for the Puerto Rican Skink Spondylurus nitidus and the Culebra Skink S. culebrae . Results indicate two high quality genomes, both ∼1.4 Gb, assembled nearly telomere to telomere with complete mitochondrion assembly and annotation.

Genome Biology and Evolution

Associations of stream geomorphic conditions and prevalence of alternative reproductive tactics among sockeye salmon populations

In many species, males may exhibit alternative life histories to circumvent the costs of intrasexual competition and female courtship. While the evolution and underlying genetic and physiological mechanisms behind alternative reproductive tactics are well studied, there has been less consideration of the ecological factors that regulate their prevalence. Here, we examine six decades of age composition records from thirty‐six populations of sockeye salmon ( Oncorhynchus nerka ) to quantify associations between spawning habitat characteristics and the prevalence of precocious sneakers known as ‘jacks’. Jack prevalence was independent of neutral genetic structure among stream populations, but varied among habitat types and as a function of continuous geomorphic characteristics. Jacks were more common in streams relative to beaches and rivers, and their prevalence was negatively associated with stream width, depth, elevation, slope and area, but positively related to bank cover. Behavioural observations showed that jacks made greater use of banks, wood and shallows than guard males, indicating that their reproductive success depends on the availability of such refuges. Our results emphasize the role of the physical habitat in shaping reproductive tactic frequencies among populations, likely through local adaptation in response to variable fitness expectations under different geomorphic conditions.

Journal of Evolutionary Biology

Nanopore amplicon sequencing reveals molecular convergence and local adaptation of rhodopsin in Great Lakes salmonids

Local adaptation can drive diversification of closely related species across environmental gradients and promote convergence of distantly related taxa that experience similar conditions. We examined a potential case of adaptation to novel visual environments in a species flock (Great Lakes salmonids, genus Coregonus ) using a new amplicon genotyping protocol on the Oxford Nanopore Flongle and MinION. We sequenced five visual opsin genes for individuals of C. artedi , C. hoyi , C. kiyi , and C. zenithicus. Comparisons revealed species-specific differences in a key spectral tuning amino acid in rhodopsin (Tyr261Phe substitution), suggesting local adaptation of C. kiyi to the blue-shifted depths of Lake Superior. Ancestral state reconstruction demonstrates that parallel evolution and “toggling” at this amino acid residue has occurred several times across the fish tree of life, resulting in identical changes to the visual systems of distantly related taxa across replicated environmental gradients. Our results suggest that ecological differences and local adaptation to distinct visual environments are strong drivers of both evolutionary parallelism and diversification.

Lake Superior

Evolution of large body size in abalones (Haliotis): Patterns and implications

Kelps and other fleshy macroalgae - dominant reef-inhabiting organisms in cool - seasmay have radiated extensively following late Cenozoic polar cooling, thus triggering a chain of evolutionary change in the trophic ecology of nearshore temperate ecosystems. We explore this hypothesis through an analysis of body size in the abalones (Gastropoda; Haliotidae), a widely distributed group in modern oceans that displays a broad range of body sizes and contains fossil representatives from the late Cretaceous (60-75 Ma). Geographic analysis of maximum shell length in living abalones showed that small-bodied species, while most common in the Tropics, have a cosmopolitan distribution, whereas large-bodied species occur exclusively in cold-water ecosystems dominated by kelps and other macroalgae. The phylogeography of body size evolution in extant abalones was assessed by constructing a molecular phylogeny in a mix of large and small species obtained from different regions of the world. This analysis demonstrates that small body size is the plesiomorphic state and largeness has likely arisen at least twice. Finally, we compiled data on shell length from the fossil record to determine how (slowly or suddenly) and when large body size arose in the abalones. These data indicate that large body size appears suddenly at the Miocene/Pliocene boundary. Our findings support the view that fleshy-algal dominated ecosystems radiated rapidly in the coastal oceans with the onset of the most recent glacial age. We conclude with a discussion of the broader implications of this change. ?? 2005 The Paleontological Society. All rights reserved.

Paleobiology

Thermal and energetic constraints on ectotherm abundance: A global test using lizards

Population densities of birds and mammals have been shown to decrease with body mass at approximately the same rate as metabolic rates increase, indicating that energetic needs constrain endotherm population densities. In ectotherms, the exponential increase of metabolic rate with body temperature suggests that environmental temperature may additionally constrain population densities. Here we test simple bioenergetic models for an ecologically important group of ectothermic vertebrates by examining 483 lizard populations. We find that lizard population densities decrease as a power law of body mass with a slope approximately inverse to the slope of the relationship between metabolic rates and body mass. Energy availability should limit population densities. As predicted, environmental productivity has a positive effect on lizard density, strengthening the relationship between lizard density and body mass. In contrast, the effect of environmental temperature is at most weak due to behavioral thermoregulation, thermal evolution, or the temperature dependence of ectotherm performance. Our results provide initial insights into how energy needs and availability differentially constrain ectotherm and endotherm density across broad spatial scales. ?? 2008 by the Ecological Society of America.

Ecology

North American Bird Banding and quantitative population ecology

Early bird-banding programs in North America were developed to provide descriptions of bird migration and movement patterns. This initial interest in description quickly evolved into more quantitative interests in two ways. There was (1) interest in quantifying migration and movement patterns, and (2) rapid recognition that re-observations of marked birds provided information about other parameters relevant to population dynamics. These included survival rate, recruitment rate, and population size. The evolution of methods for estimating population size, survival, recruitment, and movement is reviewed and we show it to be closely tied to bird-banding data. These estimation methods have been used with bird-banding data to draw important inferences about evolutionary ecology, population ecology, and population management. Illustrative examples of such inferences are provided.

Book chapter

The role of ecology in allopatric speciation of darters in the Central Highlands, USA

Allopatric speciation is the predominant mode of speciation in riverine fishes. However, the relative importance of genetic drift versus natural selection in the allopatric speciation of these fishes remain uncertain. Here, we present a case study that demonstrates the role of ecology in the diversification of a group of imperiled freshwater fishes from the central United States. We integrate a phylogenomic dataset with analyses of streamwise distance, environmental variables, meristic and morphological traits, and diet to investigate the ecological context and outcomes of allopatric speciation within a species complex comprising the Slenderhead Darter Percina phoxocephala (Nelson), Ouachita Darter Percina brucethompsoni (Robison, Cashner, and Near), and Longnose Darter Percina nasuta (Bailey). We find that two of the species traditionally delimited based on disparity in snout length, P. phoxocephala and P. nasuta , are polyphyletic, revealing three instances of the parallel evolution of snout length disparity. We propose a revised taxonomy including the delimitation of six new species based on disparity in phenotypic traits and phylogenomic analyses. We find that morphological differences are not correlated with genetic divergence but are congruent with variations in diet and environmental niches, suggesting a role for ecological factors in allopatric speciation of riverine fishes.

Arkansas, Kansas, Missouri, Oklahoma

Data set incongruence and correlated character evolution: An example of functional convergence in the hind-limbs of stifftail diving ducks

The unwitting inclusion of convergent characters in phylogenetic estimates poses a serious problem for efforts to recover phylogeny. Convergence is not inscrutable, however, particularly when one group of characters tracks phylogeny and another set tracks adaptive history. In such cases, convergent characters may be correlated with one or a few functional anatomical units and readily identifiable by using comparative methods. Stifftail ducks (Oxyurinae) offer one such opportunity to study correlated character evolution and function in the context of phylogenetic reconstruction. Morphological analyses place stifftail ducks as part of a large clade of diving ducks that includes the sea ducks (Mergini), Hymenolaimus, Merganetta , and Tachyeres , and possibly the pochards (Aythyini). Molecular analyses, on the other hand, place stifftails far from other diving ducks and suggest, moreover, that stifftails are polyphyletic. Mitochondrial cytochrome b gene sequences of eight stifftail species traditionally supposed to form a clade were compared with each other and with sequences from 50 other anseriform and galliform species. Stifftail ducks are not the sister group of sea ducks but lie outside the typical ducks (Anatinae). Of the four traditional stifftail genera, monophyly of Oxyura and its sister group relationship with Nomonyx are strongly supported. Heteronetta probably is the sister group of that clade, but support is weak. Biziura is not a true stifftail. Within Oxyura , Old World species ( O. australis, O. leucocephala, O. maccoa ) appear to form a clade, with New World species ( O. jamaicensis, O. vittata ) branching basally. Incongruence between molecules and morphology is interpreted to be the result of adaptive specialization and functional convergence in the hind limbs of Biziura and true stifftails. When morphological characters are divided into classes, only hind-limb characters are significantly in conflict with the molecular tree. Likewise, null models of synonymous and nonsynonymous substitution based on patterns of codon-degeneracy and chemical dissimilarity indicate that the nucleotide and amino acid changes postulated by the molecular tree are more plausible than those postulated by the morphological tree. These findings teach general lessons about the utility of highly adaptive characters (in particular those related to foraging ecology) and underscore the problems that convergence can pose for attempts to recover phylogeny. They also demonstrate how the concept of natural data partitions and simple models of evolution (e.g., parsimony, likelihood, neutrality) can be used to test the accuracy of independent phylogenetic estimates and provide arguments in favor of one tree topology over another.

Systematic Biology

Biomorphodynamics: Physical-biological feedbacks that shape landscapes

Plants and animals affect morphological evolution in many environments. The term "ecogeomorphology" describes studies that address such effects. In this opinion article we use the term "biomorphodynamics" to characterize a subset of ecogeomorphologic studies: those that investigate not only the effects of organisms on physical processes and morphology but also how the biological processes depend on morphology and physical forcing. The two-way coupling precipitates feedbacks, leading to interesting modes of behavior, much like the coupling between flow/sediment transport and morphology leads to rich morphodynamic behaviors. Select examples illustrate how even the basic aspects of some systems cannot be understood without considering biomorphodynamic coupling. Prominent examples include the dynamic interactions between vegetation and flow/sediment transport that can determine river channel patterns and the multifaceted biomorphodynamic feedbacks shaping tidal marshes and channel networks. These examples suggest that the effects of morphology and physical processes on biology tend to operate over the timescale of the evolution of the morphological pattern. Thus, in field studies, which represent a snapshot in the pattern evolution, these effects are often not as obvious as the effects of biology on physical processes. However, numerical modeling indicates that the influences on biology from physical processes can play a key role in shaping landscapes and that even local and temporary vegetation disturbances can steer large-scale, long-term landscape evolution. The prevalence of biomorphodynamic research is burgeoning in recent years, driven by societal need and a confluence of complex systems-inspired modeling approaches in ecology and geomorphology. To make fundamental progress in understanding the dynamics of many landscapes, our community needs to increasingly learn to look for two-way, biomorphodynamic feedbacks and to collect new types of data to support the modeling of such emergent interactions. Copyright 2008 by the American Geophysical Union.

Water Resources Research

Undeveloped and developed phases in the centennial evolution of a barrier-marsh-lagoon system: The case of Long Beach Island, New Jersey

Barrier islands and their associated backbarrier environments protect mainland population centers and infrastructure from storm impacts, support biodiversity, and provide long-term carbon storage, among other ecosystem services. Despite their socio-economic and ecological importance, the response of coupled barrier-marsh-lagoon environments to sea-level rise is poorly understood. Undeveloped barrier-marsh-lagoon systems typically respond to sea-level rise through the process of landward migration, driven by storm overwash and landward mainland marsh expansion. Such response, however, can be affected by human development and engineering activities such as lagoon dredging and shoreline stabilization. To better understand the difference in the response between developed and undeveloped barrier-marsh-lagoon environments to sea-level rise, we perform a local morphologic analysis that describes the evolution of Long Beach Island (LBI), New Jersey, over the last 182 years. We find that between 1840 and 1934 the LBI system experienced landward migration of all five boundaries, including 171 meters of shoreline retreat. Between the 1920s and 1950s, however, there was a significant shift in system behavior that coincided with the onset of groin construction, which was enhanced by beach nourishment and lagoon dredging practices. From 1934 to 2022 the LBI system experienced ~22 meters of shoreline progradation and a rapid decline in marsh platform extent. Additionally, we extend a morphodynamic model to describe the evolution of the system in terms of five geomorphic boundaries: the ocean shoreline and backbarrier-marsh interface, the seaward and landward lagoon-marsh boundaries, and the landward limit of the inland marsh. We couple this numerical modeling effort with the map analysis during the undeveloped phase of LBI evolution, between 1840 and 1934. Despite its simplicity, the modeling framework can describe the average cross-shore evolution of the barrier-marsh-lagoon system during this period without accounting for human landscape modifications, supporting the premise that natural processes were the key drivers of morphological change. Overall, these results suggest that anthropogenic effects have played a major role in the evolution of LBI over the past century by altering overwash fluxes and marsh-lagoon geometry; this is likely the case for other barrier-marsh-lagoon environments around the world.

New Jersey

A Lake Charr pangenome reveals highly conserved Ohnologs as drivers of phenotypic diversity

Whole-genome duplication (WGD) is hypothesized to spur evolutionary diversification by producing genome-wide duplicate gene sets (Ohnologs) that are initially functionally redundant but can diverge markedly as the effects of relaxed selection accumulate over time. However, the underlying mechanisms remain unclear, in part because genomic studies often reconstruct Ohnolog evolution over millions of years, during which subsequent mutations can obscure deep-time signals. Investigating the relationship between Ohnolog evolution and diversification on a contemporary timescale offers clearer insights. We explore this relationship in Lake Charr ( Salvelinus namaycush ), where ∼10% of genes are retained highly conserved polyploid duplicates following the Salmonid-Specific Fourth Round WGD. Using 31 chromosome-level assemblies of Lake Charr from morphologically and ecologically diverse populations, joined into a pangenome graph, we characterized 189,555 structural variants (SVs) that were significantly less likely to affect genes retained as sequence-conserved Ohnolog pairs, nuancing the hypothesis that gene redundancy, relaxed selection, and functional diversification are intertwined. However, we found that SVs affecting such conserved Ohnologs may be potent drivers of adaptive evolution. Notably, we identified a putative 938-Kb interchromosomal translocation containing 25 genes with highly conserved Ohnologs in a paralogous (but untranslocated) genomic block. This putative translocation appears to have facilitated Ohnolog divergence in ankrd11 and hp , genes putatively linked to craniofacial and lipid metabolic diversity in sympatric Lake Superior morphs. This research reveals that conserved Ohnologs previously presumed to be redundant remain a reservoir for adaptive change.

BioRxiv

International Watershed Technology: Improving Water Quality and Quantity at the Local, Basin, and Regional Scales

This article introduces the five papers in the “International Watershed Technology” collection. These papers were selected from 60 technical presentations at the fifth biennial ASABE 21st Century Watershed Technology Conference and Workshop: Improving the Quality of Water Resources at Local, Basin, and Regional Scales, held in Quito, Ecuador, on 3-9 December 2016. The conference focused on solving spatial and temporal water quality and quantity problems and addressed topics such as watershed management in developing countries, aquatic ecology and ecohydrology, ecosystem services, climate change mitigation strategies, flood forecasting, remote sensing, and water resource policy and management. While diverse, the presentation topics reflected the continuing evolution of the “data mining” and “big data” themes of past conferences related to geospatial data applications, with increasing emphasis on practical solutions. The papers selected for this collection represent applications of spatial data analyses toward practical ends with a theme of “tools and techniques for sustainability.” The papers address a range of topics, including the matching of crops with water availability, and assessing the environmental impacts of agricultural production. The papers identify some of the latest tools and techniques for improving sustainability in watershed resource management that are relevant to both developing and developed countries.

Transactions of the ASABE

Active mountain building and the distribution of core Maxillariinae species in tropical Mexico and Central America

The observation that southeastern Central America is a hotspot for orchid diversity has long been known and confirmed by recent systematic studies and checklists. An analysis of the geographic and elevation distribution demonstrates that the most widespread species of “core” Maxillariinae are all adapted to life near sea level, whereas the most narrowly endemic species are largely distributed in wet highland environments. Drier, hotter lowland gaps exist between these cordilleras and evidently restrict the dispersal of the species adapted to wetter, cooler conditions. Among the recent generic realignments of “core” Maxillariinae based on molecular phylogenetics, the Camaridium clade is easily the most prominent genus in Central America and is largely restricted to the highlands of Costa Rica and Panama, indicating that this region is the ancestral home of this genus and that its dispersal limits are drier, lowland cordilleran gaps. The mountains of Costa Rica and Panama are among the geologically youngest topographic features in the Neotropics, reflecting the complex and dynamic interactions of numerous tectonic plates. From consideration of the available geological evidence, I conclude that the rapid growth of the mountain ranges in Costa Rica and Panama during the late Cenozoic times created, in turn, very rapid ranges in ecological life zones and geographic isolation in that part of the isthmus. Thus, I suggest that these recent geologic events were the primary drivers for accelerated orchid evolution in southeastern Central America.

Central America

[Book Review] Biology of marine birds

A text devoted to the biology and ecology of marine birds has not been published in the last 15 years. Although a number of more taxa-specific texts have been produced during that period, there has not been a single publication that attempted to review our knowledge of all the major seabird orders since the works of Nelson (1979), Croxall (1987), and Furness and Monaghan (1987). Following the publication of those works, a large and impressive body of literature has been produced. Given the rapid expansion of the field in the last two decades, the time was ripe for production of an extensive compendium on the biology, ecology, and conservation of the world's seabirds. E. A. Schreiber and J. Burger are editors of this CRC publication, Biology of Marine Birds . The book consists of 19 chapters that vary in length from 15 to 51 pages. There are also two extensive appendices: (1) a list of seabird species (restricted to the orders Sphenisciformes, Procellariiformes, Pelecaniformes, and Charadriiformes, the latter limited to Stercorariidae, Laridae, Rhynchopidae, and Alcidae) and their IUCN status, and (2) a very useful table of species-specific life-history traits. The 19 chapters were prepared by 26 authors, among them some of the most respected and published seabird scientists in the world. A brief preface introduces the book, its objective (to provide an examination and summary of the research on seabirds), its audience (researchers, conservationists, managers, and policy-makers), and the taxa covered. The editors coauthored the introductory chapter, Seabirds in the Marine Environment. The authors describe distinctive characteristics of seabird life-histories in comparison to other taxa, hypotheses for why those lifestyles evolved and the potential role of energy limitation in the evolution of seabird life-histories. Along with a discussion of other common seabird traits, such as a propensity for colonial breeding, the authors also suggest directions for future research in seabird ecology.

The Auk

A climate for speciation: rapid spatial diversification within the Sorex cinereus complex of shrews

The cyclic climate regime of the late Quaternary caused dramatic environmental change at high latitudes. Although these events may have been brief in periodicity from an evolutionary standpoint, multiple episodes of allopatry and divergence have been implicated in rapid radiations of a number of organisms. Shrews of the Sorex cinereus complex have long challenged taxonomists due to similar morphology and parapatric geographic ranges. Here, multi-locus phylogenetic and demographic assessments using a coalescent framework were combined to investigate spatiotemporal evolution of 13 nominal species with a widespread distribution throughout North America and across Beringia into Siberia. For these species, we first test a hypothesis of recent differentiation in response to Pleistocene climate versus more ancient divergence that would coincide with pre-Pleistocene perturbations. We then investigate the processes driving diversification over multiple continents. Our genetic analyses highlight novel diversity within these morphologically conserved mammals and clarify relationships between geographic distribution and evolutionary history. Demography within and among species indicates both regional stability and rapid expansion. Ancestral ecological differentiation coincident with early cladogenesis within the complex enabled alternating and repeated episodes of allopatry and expansion where successive glacial and interglacial phases each promoted divergence. The Sorex cinereus complex constitutes a valuable model for future comparative assessments of evolution in response to cyclic environmental change.

Molecular Phylogenetics and Evolution

Macroecological patterns of sexual size dimorphism in turtles of the world

Sexual size dimorphism (SSD) is a well-documented phenomenon in both plants and animals; however, the ecological and evolutionary mechanisms that drive and maintain SSD patterns across geographic space at regional and global scales are understudied, especially for reptiles. Our goal was to examine geographic variation of turtle SSD and to explore ecological and environmental correlates using phylogenetic comparative methods. We use published body size data on 135 species from nine turtle families to examine how geographic patterns and the evolution of SSD are influenced by habitat specialization, climate (annual mean temperature and annual precipitation) and climate variability, latitude, or a combination of these predictor variables. We found that geographic variation, magnitude and direction of turtle SSD are best explained by habitat association, annual temperature variance and annual precipitation. Use of semi-aquatic and terrestrial habitats was associated with male-biased SSD, whereas use of aquatic habitat was associated with female-biased SSD. Our results also suggest that greater temperature variability is associated with female-biased SSD. In contrast, wetter climates are associated with male-biased SSD compared with arid climates that are associated with female-biased SSD. We also show support for a global latitudinal trend in SSD, with females being larger than males towards the poles, especially in the families Emydidae and Geoemydidae. Estimates of phylogenetic signal for both SSD and habitat type indicate that closely related species occupy similar habitats and exhibit similar direction and magnitude of SSD. These global patterns of SSD may arise from sex-specific reproductive behaviour, fecundity and sex-specific responses to environmental factors that differ among habitats and vary systematically across latitude. Thus, this study adds to our current understanding that while SSD can vary dramatically across and within turtle species under phylogenetic constraints, it may be driven, maintained and exaggerated by habitat type, climate and geographic location.

Journal of Evolutionary Biology

Continental-scale, seasonal movements of a heterothermic migratory tree bat

Long-distance migration evolved independently in bats and unique migration behaviors are likely, but because of their cryptic lifestyles, many details remain unknown. North American hoary bats (Lasiurus cinereus cinereus) roost in trees year-round and probably migrate farther than any other bats, yet we still lack basic information about their migration patterns and wintering locations or strategies. This information is needed to better understand unprecedented fatality of hoary bats at wind turbines during autumn migration and to determine whether the species could be susceptible to an emerging disease affecting hibernating bats. Our aim was to infer probable seasonal movements of individual hoary bats to better understand their migration and seasonal distribution in North America. We analyzed the stable isotope values of non-exchangeable hydrogen in the keratin of bat hair and combined isotopic results with prior distributional information to derive relative probability density surfaces for the geographic origins of individuals. We then mapped probable directions and distances of seasonal movement. Results indicate that hoary bats summer across broad areas. In addition to assumed latitudinal migration, we uncovered evidence of longitudinal movement by hoary bats from inland summering grounds to coastal regions during autumn and winter. Coastal regions with nonfreezing temperatures may be important wintering areas for hoary bats. Hoary bats migrating through any particular area, such as a wind turbine facility in autumn, are likely to have originated from a broad expanse of summering grounds from which they have traveled in no recognizable order. Better characterizing migration patterns and wintering behaviors of hoary bats sheds light on the evolution of migration and provides context for conserving these migrants.

Ecological Applications

Fossil diatoms and neogene paleolimnology

Diatoms have played an important role in the development of Neogene continental biostratigraphy and paleolimnology since the mid-19th Century. The history of progress in Quaternary diatom biostratigraphy has developed as a result of improved coring techniques that enable sampling sediments beneath existing lakes coupled with improved chronological control (including radiometric dating and varve enumeration), improved statistical treatment of fossil diatom assemblages (from qualitative description to influx calculations of diatom numbers or volumes), and improved ecological information about analogous living diatom associations. The last factor, diatom ecology, is the most critical in many ways, but progresses slowly. Fortunately, statistical comparison of modern diatom assemblages and insightful studies of the nutrient requirements of some common freshwater species are enabling diatom paleolimnologists to make more detailed interpretations of the Quaternary record than had been possible earlier, and progress in the field of diatom biology and ecology will continue to refine paleolimnological studies. The greater age and geologic setting of Tertiary diatomaceous deposits has prompted their study in the contexts of geologic history, biochronology and evolution. The distribution of diatoms of marine affinities in continental deposits has given geologists insights about tectonism and sea-level change, and the distribution of distinctive (extinct?) diatoms has found utilization both in making stratigraphic correlations between outcrops of diatomaceous deposits and in various types of biochronological studies that involve dating deposits in different areas. A continental diatom biochronologic scheme will rely upon evolution, such as the appearance of new genera within a family, in combination with regional environmental changes that are responsible for the wide distribution of distinctive diatom species. The increased use of the scanning electron microscope for the detailed descriptions of fossil diatoms will provide the basis for making more accurate correlations and identifications, and the micromorphological detail for speculations about evolutionary relationships.

Palaeogeography, Palaeoclimatology, Palaeoecology