Koen Sabbe

Historical processes constrain patterns in global diatom diversity

There is a long-standing belief that microbial organisms have unlimited dispersal capabilities, are therefore ubiquitous, and show weak or absent latitudinal diversity gradients. In contrast, using a global freshwater diatom data set, we show that latitudinal gradients in local and regional genus richness are present and highly asymmetric between both hemispheres. Patterns in regional richness are explained by the degree of isolation of lake districts, while the number of locally coexisting diatom genera is highly constrained by the size of the regional diatom pool, habitat availability, and the connectivity between habitats within lake districts. At regional to global scales, historical factors explain significantly more of the observed geographic patterns in genus richness than do contemporary environmental conditions. Together, these results stress the importance of dispersal and migration in structuring diatom communities at regional to global scales. Our results are consistent with predictions from the theory of island biogeography and metacommunity concepts and likely underlie the strong provinciality and endemism observed in the relatively isolated diatom floras in the Southern Hemisphere.

Experimental Studies on Sexual Reproduction in Diatoms

The diatoms are the most speciose group of algae, having global ecological significance in the carbon and silicon cycles. They are almost unique among algae in being diplontic, and sexual reproduction is an obligate stage in the life cycle of most diatom species. It is unclear which are the principal factors that have fostered the evolutionary success of diatoms, but the unique life cycle (which is correlated with a curious wall structure and cell division mechanism) and size-dependent control of sexuality must have played an important part. Progress in understanding life cycle dynamics and their interrelationships with population biology and evolution will depend on how successfully sex can be initiated and manipulated experimentally, and our review provides a foundation for such work. Relevant data are scattered in time and come mostly from non-English publications, producing a false impression of diatoms as recalcitrant with respect to sexualization. Recent advances dependent on experimental cultures include the discovery of widespread heterothallism (including some complex types of behavior) in pennate diatoms, sexual diversity among clones of centric diatoms, more flexible size restitution strategies in centric diatoms than had been suspected, and use of reproductive isolation as a criterion in diatom taxonomy. We identify unsolved problems in the life history of diatoms, including aspects of sexualization, cell-cell recognition, sexual reproduction, and the development of the special expanding cell (the auxospore), which is crucial to morphogenesis in this group. Some of these problems are being addressed using modern molecular genetic tools, and progress will be facilitated when whole-genome sequences are published (e.g., for Thalassiosira pseudonana). Problems of culture maintenance and methods for manipulating the life cycle are discussed.

Limits to gene flow in a cosmopolitan marine planktonic diatom

The role of geographic isolation in marine microbial speciation is hotly debated because of the high dispersal potential and large population sizes of planktonic microorganisms and the apparent lack of strong dispersal barriers in the open sea. Here, we show that gene flow between distant populations of the globally distributed, bloom-forming diatom species Pseudo-nitzschia pungens (clade I) is limited and follows a strong isolation by distance pattern. Furthermore, phylogenetic analysis implies that under appropriate geographic and environmental circumstances, like the pronounced climatic changes in the Pleistocene, population structuring may lead to speciation and hence may play an important role in diversification of marine planktonic microorganisms. A better understanding of the factors that control population structuring is thus essential to reveal the role of allopatric speciation in marine microorganisms.

Were the Larsemann Hills ice-free through the Last Glacial Maximum?

Lake sediments in the Larsemann Hills contain a great diversity of biological and physical markers from which past environments can be inferred. In order to determine the timing of environmental changes it is essential to have accurate dating of sediments. We used radiometric (210Pb and 137Cs), radiocarbon (AMS 14C) and uranium series (238U) methods to date cores from eleven lakes. These were sampled on coastal to inland transects across the two main peninsulas, Broknes and Stornes, together with a single sample from the Bolingen Islands. Radiometric dating of recent sediments yielded 210Pb levels below acceptable detection limits. However, a relatively well-defined peak in 137Cs gave a date marker which corresponds to the fallout maximum from the atmospheric testing of atomic weapons in 1964/65. Radiocarbon (AMS 14C) measurements showed stratigraphical consistency in the age-depth sequences and undisturbed laminae in some cores provides evidence that the sediments have remained undisturbed by glacial action. In addition, freshwater surface sediments were found to be in near-equilibrium with modern 14CO2 and not influenced by radiocarbon contamination processes. This dating program, together with geomorphological records of ice flow directions and glacial sediments, indicates that parts of Broknes were ice-free throughout the Last Glacial Maximum and that some lakes have existed continuously since at least 44 ka bp. Attempts to date sediments older than 44 ka bp using 128U dating were inconclusive. However, supporting evidence for Broknes being ice-free is provided by an Optically Stimulated Luminescence date from a glaciofluvial deposit. In contrast, Stornes only became ice-free in the mid to late Holocene. This contrasting glacial history results from the Dålk Glacier which diverts ice around Broknes. Lakes on Broknes and some offshore islands therefore contain the oldest known lacustrine sediment records from eastern Antarctica, with the area providing an ice-free oasis and refuge for plants and animals throughout the Last Glacial Maximum. These sediments are therefore well placed to unravel a unique limnological sequence of environmental and climate changes in East Antarctica from the late Pleistocene to the present. This information may help better constrain models of current climate changes and ensure the adequate protection of these lakes and their catchments from the impacts of recent human occupation.

Benthic diatom flora of freshwater and saline lakes in the Larsemann Hills and Rauer Islands, East Antarctica

A floristic and taxonomic survey was made of the diatom communities of sediments and microbial mats in 66 freshwater and saline lakes and pools in the Larsemann Hills, Rauer Islands and Bølingen Islands (continental eastern Antarctica). A total of 31 taxa were distinguished, 10 of which could not be identified to species (nine) or even generic (one) level, either because they have most probably not yet been described or because they belong to species complexes that are in need of revision. Four new combinations are proposed; three species are reported for the first time from continental Antarctica, while another three are confirmed for eastern Antarctica for the first time. Analysis of literature data on Antarctic lacustrine diatoms shows that taxonomic practice has a profound influence on the assessment of distribution patterns. Force-fitting of European and North American names to Antarctic taxa and erroneous identifications have contributed to an underestimation of endemism in the diatom flora of Antarctic inland waters. In addition, changing concepts on species boundaries during the last decade influence the interpretation of biogeographic patterns. The application of a more fine grained taxonomy will almost certainly reveal a higher degree of endemism in Antarctica, and especially continental Antarctica. The present case study shows that in the Larsemann Hills Antarctic endemics account for about 40% of all freshwater and brackish taxa, while the biogeographic distribution of about 26% is unknown, mainly due to their uncertain taxonomic identity. This contradicts the view that cosmopolitanism prevails in Antarctic diatoms.

Hidden levels of phylodiversity in Antarctic green algae: further evidence for the existence of glacial refugia

Recent data revealed that metazoans such as mites and springtails have persisted in Antarctica throughout several glacial–interglacial cycles, which contradicts the existing paradigm that terrestrial life was wiped out by successive glacial events and that the current inhabitants are recent colonizers. We used molecular phylogenetic techniques to study Antarctic microchlorophyte strains isolated from lacustrine habitats from maritime and continental Antarctica. The 14 distinct chlorophycean and trebouxiophycean lineages observed point to a wide phylogenetic diversity of apparently endemic Antarctic lineages at different taxonomic levels. This supports the hypothesis that long-term survival took place in glacial refugia, resulting in a specific Antarctic flora. The majority of the lineages have estimated ages between 17 and 84 Ma and probably diverged from their closest relatives around the time of the opening of Drake Passage (30–45 Ma), while some lineages with longer branch lengths have estimated ages that precede the break-up of Gondwana. The variation in branch length and estimated age points to several independent but rare colonization events.

EFFECTS OF DEPTH, SALINITY, AND SUBSTRATE ON THE INVERTEBRATE COMMUNITY OF A FLUCTUATING TROPICAL LAKE

Salinity is generally considered to be the dominant environmental factor regulating aquatic community structure in hydrologically closed lakes and wetlands, but it is not well known whether community response to long-term trends in hydrological balance is driven primarily by the direct physiological effect of salinity stress or by the habitat restructuring that accompanies changes in lake level and salinity. Attempts to separate the effects of various environmental factors on invertebrate populations in shallow fluctuating lakes through field study are hampered by the typically large temporal and spatial variation in species abundances and the long time scale of climate-driven habitat restructuring relative to the period of study. We used paleolimnological techniques to document long-term dynamics of the benthic invertebrate community inhabiting a shallow fluctuating lake in Kenya where during the period ∼1870–1991 lake depth fluctuated between 4 and 19 m, and lakewater conductivity between ∼250 and 14000...

Morphological, genetic and mating diversity within the widespread bioindicator Nitzschia palea (Bacillariophyceae)

Trobajo R., Clavero E., Chepurnov V.A., Sabbe K., Mann D.G., Ishihara S. and Cox E.J. 2009. Morphological, genetic and mating diversity within the widespread bioindicator Nitzschia palea (Bacillariophyceae). Phycologia 48: 443–459. DOI: 10.2216/08-69.1. Nitzschia palea (Kützing) W. Smith is believed to be a widely distributed diatom and is common in various lotic and lentic freshwater habitats. However, it is also taxonomically problematic. As part of a multidisciplinary study of this diatom, 25 clones identified morphologically as N. palea were isolated from different freshwater habitats around the world (Belgium, Brazil, Egypt, India, Japan, Paraguay, Spain, Sri Lanka and the United Kingdom). Morphological and genetic diversity (using the hypervariable D1–D2 domains of LSU rDNA) were investigated, and an almost complete set of interclonal crossing experiments was carried out. Results indicate that N. palea is not a simple, homogeneous taxon and that N. palea will probably have to be split into three or more species. Molecular and mating groups do not separate along the traditional morphological boundaries among N. palea varieties, in particular between vars palea and debilis, two taxa that are commonly used to discriminate between different degrees of water pollution. At least two of the putative species within the N. palea complex appear to be geographically widespread. Because of the complexity of variation revealed by the LSU, mating and morphometric data, it is clear that further work, using extra genetic markers and new isolates, will be needed to determine the full extent of cryptic and pseudocryptic speciation in N. palea and to investigate whether the segregate species are ecologically differentiated and have value as indicators.

Spring phytoplankton assemblages in and around the maximum turbidity zone of the estuaries of the Elbe (Germany), the Schelde (Belgium/The Netherlands) and the Gironde (France)

In spring 1993 phytoplankton samples were taken along the longitudinal salinity gradient of the estuaries of the Elbe, Schelde and Gironde. A minimum in phytoplankton abundance, biomass and diversity was observed in the inner to upper reaches of all estuaries, suggesting massive mortality in these zones. Due to a combination of high turbidity and high river runoff, phytoplankton of the Gironde estuary differed in species composition and had lower cell abundancies and biomass compared with Elbe and Schelde. Phytoplankton structure and biomass of the polyhaline reaches of the Elbe and Schelde were quite comparable. They were characterized by the diatoms Skeletonema costatum and several Thalassiosira spp. and most probably developed in situ. The presence of tychoplanktonic species in these reaches indicates exchange with the intertidal flats. On the other hand, some marked differences were observed in the upper and tidal freshwater reaches of these two estuaries. In the oligohaline Schelde stations, phytoplankton populations were dominated by the diatom Cyclotella meneghiniana. The diatom Stephanodiscus hantzschii was dominant in the limnetic stations of the Elbe estuary. Despite the unfavourable light climate, the presence of the halophilous diatom C. meneghiniana in the Schelde estuary might indicate in situ development of phytoplankton populations. This could be attributed to the high residence time and the absence of grazers due to oxygen stress. In the Elbe, river runoff is too high (at least in spring) to permit in situ development of phytoplankton in the upper and freshwater tidal reaches.

SEXUAL REPRODUCTION, MATING SYSTEM, AND PROTOPLAST DYNAMICS OF SEMINAVIS (BACILLARIOPHYCEAE)1

Cell division, the mating system, and auxosporulation were studied in the marine epipelic diatom Seminavis cf. robusta Danielidis & D. G. Mann. The interphase protoplast contains two girdle‐appressed chloroplasts, each with an elongate bar‐like pyrenoid, and also a central nucleus, located in a bridge between two vacuoles. Before cell division, the chloroplasts divide transversely and translocate onto the valves. The nucleus relocates to the ventral side for mitosis. After cytokinesis and valve formation, the chloroplasts move back to the girdle, showing a constant clockwise movement relative to the epitheca of the daughter cell. Seminavis cf. robusta is dioecious, and sexual reproduction is possible once cells are less than 50 μm. In crosses of compatible clones, gametangia pair laterally, without the formation of a copulation envelope, and produce two gametes apiece. The intensity of sexualization increases as cells reduce further in size below the 50‐μm threshold. At plasmogamy, the gametangia dehisce fully and the gametes, which were morphologically and behaviorally isogamous, fuse in the space between the gametangial thecae. The auxospore forms a transverse and longitudinal perizonium. After expansion is complete, there is an unequal contraction of the protoplast within the perizonium, creating the asymmetrical shape of the vegetative cell. Apart from this last feature, almost all characteristics exhibited by the live cell and auxospores of Seminavis agree with what is found in Navicula sensu stricto, supporting the classification of both in the Naviculaceae. Haploid parthenogenesis and polyploid auxospores were found, lending support to the view that change in ploidy may be a significant mechanism in diatom evolution.