Continental diatom biodiversity discovery and description in China: 1848 through 2019.

In this paper we inventory the continental diatom taxa described from inland waters in China, from the first species descriptions dating back to 1848 through 2019. China's geography and hydrography are complex, including the world's highest mountains, many large rivers, salty lakes, and large karst regions. From this area, a total of 1128 taxa have been described from China over this time period. We examine the number of taxa described in ca. 20-year intervals and note the periods of time of no to few descriptions, versus time intervals with many taxon descriptions. Early on, taxon descriptions of freshwater diatoms from China were done by mostly by Europeans working alone, and the time frame of 1948 to 1967 had few descriptions, as a devasting famine and the cultural revolution impacted scientific work and productivity. B.V. Skvortzov produced a large number of taxon descriptions, during his time in residence in Harbin, later while in Sao Paulo, Brazil, and even posthumously. More recently, a wide range of labs and collaborations across China, and with a diverse array of international partners, is ushering in a new, robust era of research on the biodiversity of continental diatoms. A few areas of research and work for the future are discussed.

Extensive chloroplast genome rearrangement amongst three closely related Halamphora spp. (Bacillariophyceae), and evidence for rapid evolution as compared to land plants.

Diatoms are the most diverse lineage of algae, but the diversity of their chloroplast genomes, particularly within a genus, has not been well documented. Herein, we present three chloroplast genomes from the genus Halamphora (H. americana, H. calidilacuna, and H. coffeaeformis), the first pennate diatom genus to be represented by more than one species. Halamphora chloroplast genomes ranged in size from ~120 to 150 kb, representing a 24% size difference within the genus. Differences in genome size were due to changes in the length of the inverted repeat region, length of intergenic regions, and the variable presence of ORFs that appear to encode as-yet-undescribed proteins. All three species shared a set of 161 core features but differed in the presence of two genes, serC and tyrC of foreign and unknown origin, respectively. A comparison of these data to three previously published chloroplast genomes in the non-pennate genus Cyclotella (Thalassiosirales) revealed that Halamphora has undergone extensive chloroplast genome rearrangement compared to other genera, as well as containing variation within the genus. Finally, a comparison of Halamphora chloroplast genomes to those of land plants indicates diatom chloroplast genomes within this genus may be evolving at least ~4-7 times faster than those of land plants. Studies such as these provide deeper insights into diatom chloroplast evolution and important genetic resources for future analyses.

Local habitat heterogeneity determines the differences in benthic diatom metacommunities between different urban river types.

Benthic diatoms are useful indicators of the ecological state of river systems. To understand the factors determining benthic diatom metacommunity composition in urban rivers, we studied in situ surface sediment diatom communities from 23 rivers in Shanghai City. Based on our study results on the metacommunity structure of benthic diatoms and hierarchical cluster analysis (HCA), we found substantial differences between restored (G1) and unrestored rivers (G2-G4) in taxa richness, relative abundances, and dominant and indicator taxa of benthic diatoms. The epiphytic diatoms Cocconeis placentula and Amphora libyca var. baltica were representative of the restored rivers (G1), where aquatic macrophytes were more abundant and the water was clearer. The motile epipelic diatoms Navicula recens and Navicula germainii dominated the moderately polluted rivers (G2). The eutrophic taxa Cyclotella meneghiniana, Aulacoseira granulata, and Cyclostephanos tholiformis dominated in G3, which comprised relatively heavily polluted rivers with low organic matter sediment and high disturbance. The polysaprobic taxon Nitzschia palea and the halophilous taxon Fallacia pygmaea represented relatively heavily polluted rivers with a comparatively higher sedimentary salinity (SSal) (G4). Redundancy analysis (RDA) revealed that total phosphorus (TP), dissolved organic carbon (DOC), silicon dioxide (SiO2), dissolved oxygen (DO), Secchi depth (SD), SSal, and the ratio of carbon to nitrogen (C/N) in the sediment were important environmental factors explaining variation among benthic diatom metacommunity composition. Partial RDA (pRDA) implied that the relative importance of environmental factors in structuring benthic diatom metacommunity was much higher than spatial factors. Classification and regression trees (CART) further indicated that DOC, the sediment C/N ratio, and SSal were the key local environmental factors affecting grouping patterns of benthic diatom metacommunities. Our study proposes that benthic diatom metacommunities respond to the complex characteristics of local environment in urban rivers and provides useful knowledge for consideration in the ecological monitoring of urban river systems.

Comparative analysis of the mitochondrial genomes of six newly sequenced diatoms reveals group II introns in the barcoding region of cox1.

Diatoms are the most diverse lineage of algae and at the base of most aquatic food webs, but only 11 of their mitochondrial genomes have been described. Herein, we present the mitochondrial genomes of six diatom species, including: Melosira undulata, Nitzschia alba, Surirella sp., Entomoneis sp., Halamphora coffeaeformis, and Halamphora calidilacuna. Comparison of these six genomes to the 11 currently published diatom mitochondrial genomes revealed a novel ubiquitous feature block consisting of tatC-orf157-rps11. The presence of intronic retrotransposable elements in the barcoding region of cox1 in the Halamphora genomes may explain historic difficulty (especially PCR) with cox1 as a universal barcode for diatoms. Our analysis suggests that high rates of variability in number and position of introns, in many commonly used coding sequences, prevent these from being universally viable as barcodes for diatoms. Therefore, we suggest researchers examine the chloroplast and/or nuclear genomes for universal barcoding markers.

Historical and Current Perspectives on the Systematics of the 'Enigmatic' Diatom Genus Rhoicosphenia (Bacillariophyta), with Single and Multi-Molecular Marker and Morphological Analyses and Discussion on the Monophyly of 'Monoraphid' Diatoms.

This study seeks to determine the phylogenetic position of the diatom genus Rhoicosphenia. Currently, four hypotheses based on the morphology of the siliceous valve and its various ultrastructural components, sexual reproduction, and chloroplasts have been proposed. Two previous morphological studies have tentatively placed Rhoicosphenia near members of the Achnanthidiaceae and Gomphonemataceae, and no molecular studies have been completed. The position of Rhoicosphenia as sister to 'monoraphid' diatoms is problematic due to the apparent non-monophyly of that group, so hypotheses of 'monoraphid' monophyly are also tested. Using an analysis of morphological and cytological features, as well as sequences from three genes, SSU, LSU, and rbcL, recovered from several freshwater Rhoicosphenia populations that have similar morphology to Rhoicosphenia abbreviata (Agardh) Lange-Bertalot, we have analyzed the phylogenetic position of Rhoicosphenia in the context of raphid diatoms. Further, we have used topology testing to determine the statistical likelihoods of these relationships. The hypothesis that Rhoicosphenia is a member of the Achnanthidiaceae cannot be rejected, while the hypothesis that it is a member of the Gomphonemataceae can be rejected. In our analyses, members of the Achnanthidiaceae are basal to Rhoicosphenia, and Rhoicosphenia is basal to the Cymbellales, or a basal member of the Cymbellales, which includes the Gomphonemataceae. Hypothesis testing rejects the monophyly of 'monoraphid' diatoms.

The Hawaiian freshwater algae biodiversity survey (2009-2014): systematic and biogeographic trends with an emphasis on the macroalgae.

BACKGROUND: A remarkable range of environmental conditions is present in the Hawaiian Islands due to their gradients of elevation, rainfall and island age. Despite being well known as a location for the study of evolutionary processes and island biogeography, little is known about the composition of the non-marine algal flora of the archipelago, its degree of endemism, or affinities with other floras. We conducted a biodiversity survey of the non-marine macroalgae of the six largest main Hawaiian Islands using molecular and microscopic assessment techniques. We aimed to evaluate whether endemism or cosmopolitanism better explain freshwater algal distribution patterns, and provide a baseline data set for monitoring future biodiversity changes in the Hawaiian Islands. RESULTS: 1,786 aquatic and terrestrial habitats and 1,407 distinct collections of non-marine macroalgae were collected from the islands of Kauai, Oahu, Molokai, Maui, Lanai and Hawaii from the years 2009-2014. Targeted habitats included streams, wet walls, high elevation bogs, taro fields, ditches and flumes, lakes/reservoirs, cave walls and terrestrial areas. Sites that lacked freshwater macroalgae were typically terrestrial or wet wall habitats that were sampled for diatoms and other microalgae. Approximately 50% of the identifications were of green algae, with lesser proportions of diatoms, red algae, cyanobacteria, xanthophytes and euglenoids. 898 DNA sequences were generated representing eight different markers, which enabled an assessment of the number of taxonomic entities for genera collected as part of the survey. Forty-four well-characterized taxa were assessed for global distribution patterns. This analysis revealed no clear biogeographic affinities of the flora, with 27.3% characterized as "cosmopolitan", 11.4% "endemic", and 61.3% as intermediate. CONCLUSIONS: The Hawaiian freshwater algal biodiversity survey represents the first comprehensive effort to characterize the non-marine algae of a tropical region in the world using both morphological and molecular tools. Survey data were entered in the Hawaiian Freshwater Algal Database, which serves as a digital repository of photographs and micrographs, georeferenced localities and DNA sequence data. These analyses yielded an updated checklist of the non-marine macroalgae of the Hawaiian Islands, and revealed varied biogeographic affinities of the flora that are likely a product of both natural and anthropogenic dispersal.

Molecular phylogeny of Amphora sensu lato (Bacillariophyta): an investigation into the monophyly and classification of the amphoroid diatoms.

Amphora sensu lato encompasses a large group of raphid diatoms, diverse in both form and ecology. The defining feature of this group has been an extreme asymmetry of the valve mantle and girdle bands, bringing both faces of the cell onto a single plane. Although this 'amphoroid' structure has long been the diagnostic feature and thus considered 'conservative' for the group, many have argued that the diversity of forms presently assigned to Amphora likely does not represent a monophyletic group. With the exception of several taxonomic transfers and the recent elevation of Halamphora to the level of genus, much of Amphora classification has remained unchanged for over 100 years. This study presents a phylogenetic analysis of Amphora s.l. based on a concatenated molecular alignment including the nuclear marker SSU rDNA and the chloroplast markers rbcL and psbC. These results are discussed within the framework of the current classification system of Amphora and Halamphora and lay the groundwork for a taxonomic revision of the group based on monophyly. The results of this analysis demonstrate that the genus Amphora is polyphyletic and that lineages assigned to the genus are distributed widely across the raphid diatom tree of life. The feature of amphoroid symmetry appears to have evolved independently several times. We discuss the nature of conservative characters in the raphid diatoms and their usefulness as a guide to phylogenetic relationships.